Parent Canopy Research Articles

How do birds and ants contribute to the recruitment of a tropical tree?

AbstractIn most tropical ecosystems, birds play a crucial role as primary dispersers (Phase 1) by removing seeds from the plant crown. Following primary dispersal, ground‐dwelling animals, notably ants, often disperse seeds secondarily (Phase 2). However, the relative contributions of each phase to seed dispersal effectiveness until plant establishment remains unclear. We combined observational and manipulative experiments to quantify the contributions of birds and ants to seed dispersal effectiveness of the ornithochoric tree Xylopia sericea in the Brazilian Atlantic Rainforest. Birds dispersed 17.2% of diaspores, with a dispersal distance (6.6 m ± 6.7 m) twenty times greater than that of ants (0.3 m ± 0.5 m). Notably, birds often dispersed seeds beyond the parental canopy, where seedling survival is higher. Ants were quantitatively more important to seed dispersal, with relative diaspore removal 221.3% higher than birds. Despite the limited dispersal‐distance by ants, proximity to ant nests increased seedling survival. Phase 1 led to the establishment of 16.26 seedlings per 100 diaspores produced, and Phase 2 increased the number of seedlings established by 28.6% (N = 20.91). Ants enhanced seed recruitment and improved seedling survival both away and near the parental plant, likely due to ant nests providing favorable microsites for establishment. Conversely, plants distant from ant nests performed better when away from the parental plant, underscoring the importance of birds in seed dispersal. Birds promote long‐distance dispersal and ants redistribute diaspores to microsites that increases establishment, so the dispersal of X. sericea is highly dependent on both dispersers.Abstract in Portuguese is available with online material.

A Comparison of Seed Predation, Seed Dispersal, and Seedling Herbivory in Oak and Hickory: Species with Contrasting Regenerating Abilities in a Bluegrass Savanna—Woodland Habitat

Abstract Quercus (oak) regeneration failure threatens many forest and savanna communities worldwide, where preservation of vegetation structure and composition depends on acorns germinating and surviving into adulthood. However, predation on the acorns and browsing of seedlings limits oak regeneration. To better understand the effects of these 2 mechanisms on oak recruitment in the endangered Bluegrass savanna—woodland of Kentucky, we compared seed predation and herbivory on Quercus muehlenbergii (Chinquapin Oak) with Carya laciniosa (Shellbark Hickory), a successfully regenerating tree species. Compared to hickory nuts, acorns were predated more, cached less, and dispersed shorter distances. Neither the distribution of the seedlings under the parent canopy nor browse damage differed between the 2 species. Our results suggest that seed-predation prevents regeneration of oaks in this endangered community.

Breaking and remaking a seed and seed predator interaction in the introduced range of Scotch Broom (Cytisus scoparius) in New Zealand

Summary Introduced plants may initially experience enemy release, but some of those interactions may be reinstated through biological control. These cases provide opportunities to explore the dynamics of broken and remade consumer–resource interactions. The European shrub broom (Cytisus scoparius) was introduced to New Zealand without a specialist seed predator (Bruchidius villosus) until a biological control programme reinstated this interaction in 1988. Broom produces substantially larger seeds throughout its non‐native range, and there are differences in seedling establishment beneath broom canopies between the native and introduced ranges. We hypothesized that large broom seeds produce seedlings with a survival advantage in shaded conditions, consistent with establishment under broom canopies in the introduced range. We also predicted that the seed‐feeding beetle would experience increased fitness in larger seeds. We found that seedling survival was correlated with seed weight in shaded conditions, consistent with the observed establishment of seedlings beneath broom stands in the non‐native range. Beetle size was positively correlated with seed weight, and female fecundity increased with body size. Comparison of the size distributions of beetle populations before and after hibernation provided indirect evidence that overwintering survival of beetles is also size‐dependent. These findings are consistent with observed higher levels of abundance of B. villosus in New Zealand, compared to native habitats. Selection gradients for seed size‐dependent seedling survival on broom occupancy were explored using the Rees–Paynter population model, demonstrating that a fitness benefit to the plant of large seed size through enhanced seedling survival beneath the parental canopy is only likely under low‐disturbance scenarios and moderate fecundity levels. Synthesis. The dynamics of biocontrol systems provide an underexploited opportunity to examine the impact of removal and reinstatement of species interactions. Differences in seed size between native and non‐native ranges may result from varying intensities of conflicting selection pressures acting on seed size. It remains to be seen whether a decline in seed size towards that seen in the native range will result from the biocontrol agent introduction, as we have demonstrated that selection pressures on seed size vary with disturbance‐mediated population turnover and fecundity.

Study of the Young Growth of Main Forest-forming Coniferous Species in the Primary Forests of the Sikhote-Alin Nature Reserve

This paper presents the results of studies of the young growth of main forest-forming coniferous species in four types of native cedar forest of the Sikhote-Alin nature reserve. The following types of young growth were measured: Manchurian fir (Abies nephrolepis), Ajan spruce (Picea ajanensis), Japanese yew (Taxus cuspidata), and Korean pine (Pinus koraiensis). The research was conducted during 5 field seasons from 2008 to 2012 at 4 permanent study areas. For each species, the common silvicultural parameters, distribution of height, age of young growth and dynamics of the average apical growth data for the twenty-year period were identified and analysed. It was established that the size and strength of young growth are affected by complex forest site conditions that are determined by: the composition of the parent canopy, stand age, and position in the landscape. The growing conditions for young growth conifers are more favourable in mountain cedar forests compared to cedar forests in a valley. A parent canopy of a specific phytocenosis forms a unique microclimate, significantly weakening and softening the impact of macro-climate indicators of the growing area on the value of the apical growth of young growth. Spruce young growth is the most responsive to changes in climatic parameters and Korean pine young growth is the least responsive.

What would happen to the trees and lianas if apes disappeared?

AbstractApes, like many frugivorous animals, are crucial allies for the reproduction of several fruiting tree species. Almost all apes, however, including bonobos Pan paniscus, are threatened with extinction. How will this affect tree conservation? How can plants that are adapted to seed dispersal by apes reproduce without their dispersal vectors? At LuiKotale, in an evergreen tropical forest of the Democratic Republic of Congo, the recruitment of 22 plant species in the absence of seed dispersal was investigated under the parental canopy, where a proportion of seeds fall without horizontal dissemination. Most bonobo-dispersed plant species (95% of 19 species) were unable to self-recruit under the canopy. As 40% of the tree species (65% of trees) at LuiKotale are dispersed by bonobos there is a risk of ecosystem decay and simplification (reduced biodiversity) if Pan paniscus disappears from its natural range. The extinction of other apes from their forests could have similar consequences. The conservation of tree species, therefore, must encompass conservation of pollinators, seed dispersal vectors and other species that provide ecological services to the trees and other fruiting plants.

Low redundancy in seed dispersal within an island frugivore community.

The low species diversity that often characterizes island ecosystems could result in low functional redundancy within communities. Flying foxes (large fruit bats) are important seed dispersers of large-seeded species, but their redundancy within island communities has never been explicitly tested. In a Pacific archipelago, we found that flying foxes were the sole effective disperser of 57 % of the plant species whose fruits they consume. They were essential for the dispersal of these species either because they handled >90 % of consumed fruit, or were the only animal depositing seeds away from the parent canopy, or both. Flying foxes were especially important for larger-seeded fruit (>13 mm wide), with 76 % of consumed species dependent on them for dispersal, compared with 31 % of small-seeded species. As flying foxes decrease in abundance, they cease to function as dispersers long before they become rare. We compared the seed dispersal effectiveness (measured as the proportion of diaspores dispersed beyond parent crowns) of all frugivores for four plant species in sites where flying foxes were, and were not, functionally extinct. At both low and high abundance, flying foxes consumed most available fruit of these species, but the proportion of handled diaspores dispersed away from parent crowns (quality) was significantly reduced at low abundance. Since alternative consumers (birds, rodents and land crabs) were unable to compensate as dispersers when flying foxes were functionally extinct, we conclude that there is almost no redundancy in the seed dispersal function of flying foxes in this island system, and potentially on other islands where they occur. Given that oceanic island communities are often simpler than continental communities, evaluating the extent of redundancy across different ecological functions on islands is extremely important.

The seed ecology of an ornamental wattle in South Africa — Why has Acacia elata not invaded a greater area?

Australian Acacia species introduced to South Africa as ornamentals have notably smaller invasive ranges than those introduced for forestry or dune stabilization. We asked whether the relatively small invasive extent of Acacia elata, a species used widely for ornamental purposes, is due to low rates of reproduction. Age at reproductive maturity, seed dispersal, annual seed production, seed bank dynamics and seed germination and viability were assessed at five sites in the Western Cape. Results indicate that A. elata has similar traits to other invasive Australia Acacia species: annual seed input into the leaf litter was high (up to 5000seedsm−2); large seed banks develop (>20,000seedsm−2) in established stands; seed germinability is high (>90%); seeds accumulate mostly in the top soil layers but can infiltrate to depths of 40cm; and seed germination appears to be stimulated by fire. However the age at the onset of reproduction (~4years) is longer than most widespread invaders (~3years) and dispersal is fairly limited (seeds fell up to distances of 6m from the parent canopy; the highest density of seed rain was found directly under the canopy with over 20% of seeds falling directly under the terminal branches). We suggest that the current limited distribution of invasive A. elata populations is the result of the relatively small size of initial populations (cf. large plantations and widespread plantings for forestry and dune stabilization species), the species' apparent lack of secondary dispersal vectors, and the planting of trees in gardens and urban settings offer limited opportunities for recruitment, proliferation and spread. The species is, however, increasing in abundance and range. We propose methods to improve management of invasions of the species. Management to reduce seed production of this species through classical biological control, as has been achieved for other Australian Acacia species in South Africa, should be prioritised.

Small-scale spatial structure within patterns of seed dispersal

A large proportion of dispersing propagules land near their maternal plant, even in species that have evolved structures which enhance dispersal. For these propagules, their post-dispersal spatial pattern is likely to reflect the overall shape and scale of the parental plant canopy and, especially in poorly dispersing species, aggregation of propagules on the plant prior to dispersal. Localised patterns within seed shadows are also likely to be affected by secondary movement after dispersal, leading to either more or less small-scale aggregation, depending on the mechanism. Our general aim was to study the small-scale spatial structure within patterns of seed dispersal of Raphanus raphanistrum L. to generate hypotheses about the sequence of processes and events leading to the spatial pattern of dispersal in this species. More specifically, we determined the sizes of small-scale structures within the seed shadows on the ground after dispersal and the extent to which these match the sizes of pre-dispersal aggregations within the parental canopy. Variation in plant size and shape was provided by four levels of inter-specific competition resulting from differing wheat crop densities. Positions of propagules were determined using a three-dimensional digitizer, and the data for each plant were analysed using point pattern analysis. Not surprisingly, larger plants, growing at lower plant density, had larger seed shadows, showing an overall influence of maternal plant size. The pattern of propagules exhibited significant small-scale aggregates, with similar sizes on the plant and on the ground. There was no evidence that aggregation size was greater on the ground or increased with time, but the strength of the aggregation increased with length of time on the ground.

Structural development and productivity of replanted mangrove plantations in Kenya

Forest structure and productivity was investigated in a 12-year-old Rhizophora mucronata Lam. plantation at Gazi Bay, Kenya. Sampling was carried out in 22, 10 m × 10 m quadrats laid along belt transects perpendicular to the waterline. Within each quadrat all trees with stem diameter greater than 2.5 cm were identified, position marked and counted. Vegetation measurements included tree height (m), canopy cover (%) and stem diameter measured at 1.3 m above the ground ( D 130); from which were derived basal area (m 2/ha); stand density (stems/ha) and biomass (t/ha). Information regarding composition and distribution of juveniles was derived using linear regeneration sampling (LRS). The replanted forest had a stand density of 5132 stems/ha; with a mean canopy height and stem diameter of 8.4 ± 1.1 m (range: 3.0–11.0 m) and 6.2 ± 1.87 cm (range: 2.5–12.4 cm), respectively. The total juvenile density was 4886 saplings per hectare; 78.6% of which constituted the parental canopy. The standing biomass for the 12-year-old R. mucronata plantation was 106.7 ± 24.0 t/ha, giving a biomass accumulation rate of 8.9 t/(ha year).

Linking seed dispersal, germination and seedling recruitment in the invasive species Berberis darwinii (Darwin’s barberry)

Seedling recruitment is a multi-phased process involving seed production, dispersal, germination, seedling establishment and subsequent survival. Understanding the factors that determine success at each stage of this process is of particular interest to scientists and managers seeking to understand how invasive species spread and persist, and identify critical stages for management. To understand the factors and processes influencing recruitment of the invasive species Berberis darwinii Hook. (Darwin’s barberry), temporal and spatial patterns of seed dispersal, germination and seedling establishment were examined. Seed dispersal from a large source population was measured over two fruiting seasons, and subsequent patterns of seedling emergence and survival within each cohort were measured. Seed longevity was tested under both natural and artificial conditions. Seeds were widely dispersed by birds, up to 450 m from the source population. Dispersal was essential to seedling establishment, as few seedlings survived beneath the parent canopy. Seeds were relatively short-lived in the soil under both field and glasshouse conditions, with few surviving for more than 1 year. Patterns of newly emerged seedlings largely reflected patterns of seed rain, but seedling survival was significantly affected by distance from source population, seedling density and light environment. These results suggest that recruitment of B. darwinii is dependent on dispersal of seeds to favourable microsites. Management priorities should include the removal of fruiting plants, and seedling control in highlight areas.

Offspring spatial patterns in Picconia excelsa (Oleaceae) in the Canarian laurel forest

We studied the spatial patterns of seedlings and seeds in isolated Picconia excelsa (Oleaceae) trees in the laurel forest of Anaga, Tenerife (Canary Islands). By finding isolated trees we assessed the correlation of seed and seedling bank traits and parent trees by removing the confounding effects of proximity (<100 m radius) of conspecific fruiting trees. We counted all the seedlings per age (height) class within its parental range, and sampled the seed number along transects departing from beneath the parent canopy at regular intervals. We mapped all seedlings per age class and plotted seed and seedling profiles in relation to distance to parent trees. Older Picconia seedlings tended to clump significantly further from parent trees than younger seedlings, which clumped just beneath the parents. We found significant differences among distances to parent tree in numbers of seedlings per age class. The seedling bank area was significantly correlated with maximum distance of seedlings to parent trees. The majority of seeds were deposited within the first 4 m below the parent crown. Seedlings amount at further distances from the trees is larger than seeds/fruits as counted on the ground. Our results suggest that disseminated, older seedlings have occupied germination sites far from the parent tree because there is probably lower seedling–seedling and parent–seedling competition for resources, and perhaps no intraspecific allelopathy and predation/disease.

Recruitment of Pterocarpus angolensis in the wild

Pterocarpus angolensis is a hardwood species subject to heavy exploitation throughout miombo woodlands of south-central Africa. Rates of natural recruitment measured in western Tanzania were found to be low, with only a third of parent trees generating any seedlings despite considerable seed production. Although heavily protected areas supported large parent trees, these adults had very few seedlings of >5 cm DBH nearby, possibly suggesting mortality caused by high densities of browsing ungulates. Outside these well protected areas, cutting has removed most large trees and remaining adults are small producing relatively few seeds. Factors affecting recruitment include location and elevation perhaps indicative of higher precipitation, whereas short grass and reduced parent canopy cover are associated with more seedlings suggesting competition for light. Given these recruitment rates, the prognosis for sustainable exploitation of P. angolensis looks bleak.

COMPARATIVE SEED SHADOWS OF BIRD-, MONKEY-, AND WIND-DISPERSED TREES

Although spatial patterns of seed distribution are thought to vary greatly among plant species dispersed by different vectors, few studies have directly examined this assumption. We compared patterns of seed rain of nine species of trees disseminated by large birds, monkeys, and wind in a closed canopy forest in Cameroon. We used maximum-likelihood methods to fit seed rain data to four dispersal functions: inverse power, negative exponential, Gaussian, and Student t. We then tested for differences in dispersal characteristics (1) among individuals within species, and (2) among species dispersed by the same vector. In general, an inverse power function best described animal-dispersed species and the Gaussian and Student t functions best described wind-dispersed species. Animal-dispersed species had longer mean dispersal distances than wind-dispersed species, but lower fecundities. In addition to these distinct differences in average dispersal distance and functional form of the seed shadow between animal- and wind-dispersed species, seed shadows varied markedly within species and vector, with conspecifics and species within vector varying in their dispersal scale, fecundity, and clumping parameters. Dispersal vectors determine a significant amount of variation in seed distribution, but much variation remains to be explained. Finally, we demonstrate that most seeds, regardless of vector, fall directly under the parent canopy. Long-distance dispersal events (>60 m) account for a small proportion of the seed crop but may still be important in terms of the absolute numbers of dispersed seeds and effects on population and community dynamics.

Spatial distribution of seeds and juveniles of Enterolobium glaziovii Bentham (Leguminosae, Mimosoideae) in the Atlantic forest, Brazil

The spatial distribution of fruits and juveniles of Enterolobium glaziovii Bentham were evaluated in four circular plots of 30 m radius, centered around reproductive individuals to determine whether the size of the parental crowns, topography, and canopy coverage influence these patterns. Each crown was subdivided into eight irregular triangles and the area of each was calculated. Each plot was subdivided into four sub-plots, two of them in an up-slope direction, and the other two in a down-slope direction. The canopy coverage for each plot was estimated at breast height (1.30 m) and at ground level, every three months during 1998. The numbers of fruits and juveniles of E. glaziovii were recorded. Although there were more fruit under the parental crown (F1,12 = 0.01; p = 0.92), there was no difference in the number of juveniles under and beyond the reproductive trees (F1,12 = 0.88; p = 0.36). There was a positive correlation between the areas of the crowns and the numbers of fruits and juveniles under them (r = 0.62; df = 14; p &lt; 0.05). Most of the fruits and juveniles were found in the down-slope directions. The canopy coverage was lower in spring, but higher under parental crowns (only at breast height). These results indicate that the parental crown, topography and canopy coverage influenced the pattern of spatial distribution and probably also the dynamics of E. glaziovii seeds and juveniles.

UV‐A/BLUE LIGHT–INDUCED REACTIVATION OF SPORE GERMINATION IN UV‐B IRRADIATED ULVA PERTUSA (CHLOROPHYTA)1

Recent reduction in the ozone shield due to manufactured chlorofluorocarbons raised considerable interest in the ecological and physiological consequences of UV‐B radiation (λ=280–315 nm) in macroalgae. However, early life stages of macroalgae have received little attention in regard to their UV‐B sensitivity and UV‐B defensive mechanisms. Germination of UV‐B irradiated spores of the intertidal green alga Ulva pertusa Kjellman was significantly lower than in unexposed controls, and the degree of reduction correlated with the UV doses. After exposure to moderate levels of UV‐B irradiation, subsequent exposure to visible light caused differential germination in an irradiance‐ and wavelength‐dependent manner. Significantly higher germination was found at higher photon irradiances and in blue light compared with white and red light. The action spectrum for photoreactivation of germination in UV‐B irradiated U. pertusa spores shows a major peak at 435 nm with a smaller but significant peak at 385 nm. When exposed to December sunlight, the germination percentage of U. pertusa spores exposed to 1 h of solar radiation reached 100% regardless of the irradiation treatment conditions. After a 2‐h exposure to sunlight, however, there was complete inhibition of germination in PAR+UV‐A+UV‐B in contrast to 100% germination in PAR or PAR+UV‐A. In addition to mat‐forming characteristics that would act as a selective UV‐B filter for settled spores under the parental canopy, light‐driven repair of germination after UV‐B exposure could explain successful continuation of U. pertusa spore germination in intertidal settings possibly affected by intense solar UV‐B radiation.

Effects of canopy and settlement density on the performance of the brown seaweed Fucus serratus germlings

Effects of the settlement density of germlings and canopy on settled germlings of Fucus serratus were investigated on the rocky shore of the Isle of Man. The survival of transplanted germlings was mainly determined by parent canopy rather than by initial settlement density of germlings. However, germling growth was greater at low density than at high density and enhanced by canopy removal. Recruitment by natural propagules was stimulated at high settlement density and maximal recruits occurred on caged slides under the canopy. On the experimental slides, tiny snails and sedimentation were found. The number of snails was positively related with the settlement density of germlings indicating that they fed the germlings. Sedimentation and snail number were greater with canopy removal treatments than in canopy intact ones. These indicate that canopy sweeping gives benefits to germlings by removing sediment from substrata and protecting them from herbivores. In conclusion, the survival of settled F. serratus germlings is mainly determined by canopy sweeping and their growth is retarded in the presence of a canopy and at high settlement density.

Investigations in plant commonness and rarity: a comparison of seed removal patterns in the widespread Jatropha standleyi and the endemic J. chamelensis (Euphorbiaceae)

During early August to late September 1998 we examined seed dispersal in the tree species Jatropha standleyi (widespread) and J. chamelensis (endemic; Euphorbiaceae) at the Estación de Biologı́a Chamela, Jalisco, México. Using general linear models, we found that seed removal from beneath the parental canopy did not differ between the two species and that seed removal was correlated with both seed availability on the ground and rodent predation. We inferred that seed removal was largely mediated by rodents and determined that if a seed was not removed within 4 d (approx. 16% chance), it was highly unlikely to be removed at all (approx. 3.5% chance) for both species. We note that although differences in patterns of seed removal may not explain the observed differences in range size, differences between the two species in seed crop size are similar to patterns found in other endemic/widespread species pairs.

Investigations in plant commonness and rarity: a comparison of seed removal patterns in the widespread Jatropha standleyi and the endemic J. chamelensis (Euphorbiaceae)

During early August to late September 1998 we examined seed dispersal in the tree species Jatropha standleyi (widespread) and J. chamelensis (endemic; Euphorbiaceae) at the Estacion de Biologia Chamela, Jalisco, Mexico. Using general linear models, we found that seed removal from beneath the parental canopy did not differ between the two species and that seed removal was correlated with both seed availability on the ground and rodent predation. We inferred that seed removal was largely mediated by rodents and determined that if a seed was not removed within 4d (approx. 16% chance), it was highly unlikely to be removed at all (approx. 3.5% chance) for both species. We note that although differences in patterns of seed removal may not explain the observed differences in range size, differences between the two species in seed crop size arc similar to patterns found in other endemic/widespread species pairs.

The effect of bat (Rousettus aegyptiacus) dispersal on seed germination in eastern Mediterranean habitats.

The fruit-bat Rousettus aegyptiacus (Pteropodidae) in Israel consumes a variety of cultivated and wild fruits. The aim of this study was to explore some of its qualities as a dispersal agent for six fruit-bearing plant species. The feeding roosts of the fruit-bat are located an average of 30 m from its feeding trees and thus the bats disperse the seeds away from the shade of the parent canopy. The bat spits out large seeds but may pass some (2%) of the small seeds (<4 mg) through its digestive tract. However, neither the deposited seeds nor the ejected seeds (except in one case) had a significantly higher percentage germinating than intact seeds. Although the fruit-bat did not increase the percentage germinating, seeds of three plant species subject to different feeding behaviors (deposited in feces or spat out as ejecta) had a different temporal pattern of germination from the intact seeds. The combined seed germination distribution generated by these different treatments is more even over time than for each treatment alone. It is sugested that this increases asynchronous germination and therefore enhances plant fitness by spreading the risks encountered during germination, especially in eastern Mediterranean habitats where the pattern of rainfall is unpredictable.

Recruitment pattern of Rhus integrifolia populations in periods between fire in chaparral

AbstractPopulation structure, fruit production and dispersal, and recruitment of Rhus integrifolia, a gynodioecious sclerophy llous shrub living in coastal chaparral, were studied in two localities in southern California last burned 60 and 90 years ago. Though hermaphroditic plants produce some fruits, only the male‐sterile plants bore significant numbers. Among the sample plants, one individual accounted for 50% of the crop measured. Terrestrial animals, through the loss of some seeds taken for consumption, play a decisive role in the seed dispersal, promoting the establishment of seedlings outside the parent's canopy. Birds have not been demonstrated to contribute to long distance dispersal, but they are responsible for dropping 25% of the fruits collected below the R. integrifolia canopies, 36%) of which were completely destroyed. Allelopathic effects were not observed in laboratory assays, and germination is probably not influenced by the species beneath which seeds have been deposited. This pattern can explain the non‐aggregated distribution of individuals observed in the studied populations. We conclude that establishment of new individuals has been continuous at both localities and that the populations of R. integrifolia are increasing in these long unburned stands. This may be indicative of a successional trend in unburned chaparral and coastal scrub toward a sclerophy llous woodland.