The Role of Arboreal Seed Dispersal Groups on the Seed Rain of a Lowland Tropical Forest1

Most tropical plants produce fleshy fruits that are dispersed primarily by vertebrate frugivores. Behavioral disparities among vertebrate seed dispersers could influence patterns of seed distribution and thus forest structure. This study investigated the relative importance of arboreal seed dispersers and seed predators on the initial stage of forest organization–seed deposition. We asked the following questions: (1) To what degree do arboreal seed dispersers influence the species richness and abundance of the seed rain? and (2) Based on the plant species and strata of the forest for which they provide dispersal services, do arboreal seed dispersers represent similar or distinct functional groups? To answer these questions, seed rain was sampled for 12 months in the Dja Reserve, Cameroon. Seed traps representing five percent of the crown area were erected below the canopies of 90 trees belonging to nine focal tree species: 3 dispersed by monkeys, 3 dispersed by large frugivorous birds, and 3 wind‐dispersed species. Seeds disseminated by arboreal seed dispersers accounted for ca 12 percent of the seeds and 68 percent of the seed species identified in seed traps. Monkeys dispersed more than twice the number of seed species than large frugivorous birds, but birds dispersed more individual seeds. We identified two distinct functional dispersal groups, one composed of large frugivorous birds and one composed of monkeys, drop dispersers, and seed predators. These groups dispersed plants found in different canopy strata and exhibited low overlap in the seed species they disseminated. We conclude it is unlikely that seed dispersal services provided by monkeys could be compensated for by frugivorous birds in the event of their extirpation from Afrotropical forests.     

Early herbaceous succession along a topographical gradient on forest clear-felling sites in mountainous terrain,central Japan

Early successional patterns of herbaceous communities in forest clear-felling sites were investigated along a topographical gradient, which included ridge, slope and valley habitat types, in warm-temperate evergreen forest regions of central Japan for 5 years. Three dominant species with wind-dispersed seeds played a major role in the succession: an annual,Crassocephalum crepidioides, a biennial,Erigeron canadensis, and a perennial,Miscanthus sinensis. Pioneer herbs that have a seed-bank strategy, which are common in old field succession, were not found in the mountainous sites. The persistence of the annual or biennial dominants during the very early stages of secondary succession was different in the three topographical habitat types. In the ridge habitat,M. sinensis dominated from the first year state, andC. crepidioides andE. canadensis were less prominent.Crassocephalum crepidioides andE. canadensis became dominant as one moved down the slope. In the valley habitat,C. crepidioides dominated in the first year stage, was succeeded byE. canadensis in the second year, and thenM. sinensis gradually replaced it in later years. As all three wind-dispersed dominants simultaneously invaded in all the habitat types after clear-felling, the different successional patterns along the topographical gradient might have resulted from differences in the establishment ability and the growth rate of the three dominants depending on the three habitat types.     

Seed Dispersal by Monkeys and the Fate of Dispersed Seeds in a Peruvian Rain Forest1

Primary seed dispersal by two species of monkeys and the effects of rodents and dung beetles on the fate of dispersed seeds are described for a rain forest in southeastern Perú. During the six-month study period (June–November 1992) spider monkeys (Ateles paniscus) dispersed the seeds of 71 plant species, whereas howler monkeys (Alouatta seniculus) dispersed seeds of 14 species. Spider and howler monkeys also differed greatly in their ranging behavior and defecation patterns, and as a consequence, produced different seed rain patterns. Monkey defecations were visited by 27 species of dung beetles (Scarabaeidae). Dung beetles buried 41 percent of the seeds in the dung, but the number of seeds buried varied greatly, according to seed size. Removal rates of unburied seeds by rodents varied between 63–97 percent after 30 d for 8 plant species. The presence of fecal material increased the percentage of seeds removed by seed predators, but this effect became insignificant with time. Although seed predators found some seeds buried in dung balls (mimicking burial by dung beetles), depth of burial significantly affected the fate of these seeds. Less than 35 percent of Brosimum lactescens seeds buried inside dung balls at a depth of 1 cm remained undiscovered by rodents, whereas at least 75 percent of the seeds escaped rodent detection at a depth of 3 cm and 96 percent escaped at 5 cm. Both dung beetles and rodents greatly affected the fate of seeds dispersed by monkeys. It is thus important to consider postdispersal factors affecting the fate of seeds when assessing the effectiveness of frugivores as seed dispersers.     

药用植物种子休眠类型及休眠解除研究进展

种子休眠是植物在长期发育过程中形成的对不良环境条件主动适应的现象。介绍了常见药用植物种子休眠的原因,根据休眠原因对种子休眠与萌发的影响将种子休眠划分的几种类型,总结了对不同休眠类型的种子应分别采用的休眠解除方法,并评述了目前的研究进展。     

Frugivory and Seed Dispersal by Four Species of Primates in Madagascar's Eastern Rain Forest1

In the unique faunal assemblage of the Malagasy rain forest, lemurs appear to play particularly important roles as seed-dispersing frugivores. A three-month study of feeding ecology and seed dispersal by four species of lemurs in Madagascar's eastern rain forest found that three species, Eulemur rubriventer, Eulemur fulvus, and Varecia variegata were seed dispersers, and the fourth, Propithecus diadema, was a seed predator. In germination trials, seeds passed by lemurs sprouted significantly faster and in greater numbers than those not passed by lemurs. Analysis of fruit morphologies of 69 local plant taxa producing fleshy fruits during the study period found that these fruits fell into two well-defined color categories that correlated significantly with fruit size. Seventy seven percent of fleshy fruits greater than 10 mm in diameter were colored green, brown, tan, purplish, or black, while all fruits less than 10 mm in diameter were colored red, yellow, orange, pink, blue, or white. Three introduced exotic plant species provided exceptions to this pattern, producing fruits which were larger than 10 mm and pink or orange. Fruits chosen by the primates in this study were usually larger than 10 mm in diameter and were in nearly all cases colored green, brown, tan, purplish, red, or some combination of these colors. Morphological traits shared by fruits of multiple plant taxa in the diets of seed-dispersing lemurs suggest possible coevolved relationships between Malagasy rain forest plants and lemurs.     

Natural Regeneration in Plantations of Native Trees in Lowland Brazilian Atlantic Forest: Community Structure,Diversity, and Dispersal Syndromes

Plantations of native‐tree species are often recommended for ecological restoration, but the understanding of how these techniques catalyze natural ecological processes is limited. We investigated natural regeneration in five plantations of native trees in the Poço das Antas Biological Reserve (PABR) in the state of Rio de Janeiro, Brazil. The plantations were 9–11 years old, and contained 8–14 native‐tree species with different compositions and relative density of species. We analyzed floristic composition, structure (density and basal area) of overstory and understory strata, as well as other ecological attributes (dispersal syndromes, fruit or seed size, and the availability of fruit for frugivores). Zoochorous species comprised 77% of the community, with a prevalence of the two smallest size classes of propagules (< 0.6 and 0.6–1.6 cm) in natural regeneration. The density of zoochorous plants in the understory was positively correlated with their density in the overstory, indicating their influence on natural regeneration (r² = 0.36; p < 0.0002). Fruit availability for frugivores (density and richness of plants fruiting during the year) was also positively correlated with the density of stems in the understory. Therefore, attributes such as dispersal syndrome and fruiting season should be considered in selecting species to be planted. The differences in natural regeneration observed in each of the native‐tree plantations indicated that the performance of plantations as a restoration strategy may differ, depending on initial species composition, planting density, and site conditions.     

Seed Dispersal by Bats and Birds in Forest and Disturbed Habitats of Chiapas,Mexico1

We examined seed dispersal by bats and birds in four habitats of the Selva Lacandona tropical rain forest region, Chiapas, Mexico. The four habitats represented a disturbance gradient: active cornfield, ten-year-old abandoned cornfield, cacao plantation, and forest. Using seed traps examined before sunrise (0400 h) and before sunset (1800 h), we compared volant vertebrate seed dispersal, assuming that seeds found at the end of the night were dispersed by bats and those found at the end of the day were dispersed by birds. We did not find seeds from other frugivores such as monkeys or opossums. In all habitats bats dispersed more seeds than birds. In most months bats also dispersed more seeds than birds, except in December when no seeds were found in the traps. Bats also consistently dispersed more species of seeds than birds, although a x2 comparison showed differences not to be significant. Fifty percent of the species represented in the dispersed seeds in all habitats were pioneer species. Cecropia seeds represented a high percentage (up to 87% of those dispersed by bats and up to 83% by birds) of dispersed seeds that fell in our traps. The influence of bats and birds on secondary successional processes is likely to be fundamental for the establishment of vegetation. Since bats dispersed more seeds than birds (primarily to disturbed areas and consisting primarily of pioneer species), they are likely to play an important role in successional and restoration processes among habitats as structurally and vegetationally different as cornfields, old fields, cacao plantations, and forest.     

Forest Fragmentation Reduces Seed Dispersal of Duckeodendron cestroides, a Central Amazon Endemic

Fragmentation that alters mutualistic relationships between plants and frugivorous animals may reduce the seed dispersal of trees. We examined the effects of forest fragmentation on the distributions of seeds and seedlings of a Central Amazon endemic tree, Duckeodendron cestroides . In the dry seasons of 2002–2004, seeds and first-year seedlings were counted within wedge-shaped transects centered around Duckeodendron adults in fragments and nearby continuous forests at the Biological Dynamics of Forest Fragmentation Project. Analyses showed that fragmentation reduced seed dispersal quantity and quality. The percent and distance of dispersed seeds were both twice as great in continuous forest as in fragments. The distances of each tree's five furthest dispersed seeds were three times greater in continuous forest than fragments. Over the 3-yr study, 20 times more seeds were dispersed more than 10 m from parent crowns in continuous forest than fragments. A regression analysis showed more dispersed seeds at all distances in continuous forest than fragments. Dispersal differences were strong in 2002 and 2004, years of large fruit crops, but weak or absent in 2003, when fruit production was low. As distance from parent crowns increased, the number of seedlings declined more rapidly in fragments than continuous forest. Distance-dependent mortality between the seed and seedling stages appeared to be more important in continuous forest than fragments. This research provides ample, indirect evidence demonstrating that forest fragmentation can result in the breakdown of a seed dispersal mutualism, potentially jeopardizing the reproduction of a rare, tropical tree.     

水曲柳种子次生休眠的预防和解除

以解除休眠和经高温诱导产生次生休眠的水曲柳种子为材料,探讨预防和解除水曲柳种子次生休眠的方法。结果表明：GA3和乙烯利不能阻止已解除休眠水曲柳种子在25℃下萌发时诱导的次生休眠。干燥和短时间低温解除水曲柳种子次生休眠的效果不明显,较长时间（大于2周）的低温对解除水曲柳种子的次生休眠有一定的效果。综合来看,10^-3mol&#183;L^-1的乙烯利或GA4＋7解除水曲柳种子次生休眠的效果较好。     

Forest Regeneration in a Chronosequence of Tropical Abandoned Pastures: Implications for Restoration Ecology

During the mid‐1900s, most of the island of Puerto Rico was deforested, but a shift in the economy from agriculture to small industry beginning in the 1950s resulted in the abandonment of agricultural lands and recovery of secondary forest. This unique history provides an excellent opportunity to study secondary forest succession and suggest strategies for tropical forest restoration. To determine the pattern of secondary succession, we describe the woody vegetation in 71 abandoned pastures and forest sites in four regions of Puerto Rico. The density, basal area, aboveground biomass, and species richness of the secondary forest sites were similar to those of the old growth forest sites (>80 yr) after approximately 40 years. The dominant species that colonized recently abandoned pastures occurred over a broad elevational range and are widespread in the neotropics. The species richness of Puerto Rican secondary forests recovered rapidly, but the species composition was quite different in comparison with old growth forest sites, suggesting that enrichment planting will be necessary to restore the original composition. Exotic species were some of the most abundant species in the secondary forest, but their long‐term impact depended on life history characteristics of each species. These data demonstrate that one restoration strategy for tropical forest in abandoned pastures is simply to protect the areas from fire, and allow natural regeneration to produce secondary forest. This strategy will be most effective if remnant forest (i.e., seed sources) still exist in the landscape and soils have not been highly degraded. Patterns of forest recovery also suggest strategies for accelerating natural recovery by planting a suite of generalist species that are common in recently abandoned pastures in Puerto Rico and throughout much of the neotropics.     

Mammal Abundances and Seed Traits Control the Seed Dispersal and Predation Roles of Terrestrial Mammals in a Costa Rican Forest

In Neotropical forests, mammals act as seed dispersers and predators. To prevent seed predation and promote dispersal, seeds exhibit physical or chemical defenses. Collared peccaries (Pecari tajacu) cannot eat some hard seeds, but can digest chemically defended seeds. Central American agoutis (Dasyprocta punctata) gnaw through hard‐walled seeds, but cannot consume chemically defended seeds. The objectives of this study were to determine relative peccary and agouti abundances within a lowland forest in Costa Rica and to assess how these two mammals affect the survival of large seeds that have no defenses (Iriartea deltoidea, Socratea exorrhiza), physical defenses (Astrocaryum alatum, Dipteryx panamensis), or chemical defenses (Mucuna holtonii) against seed predators. Mammal abundances were determined over 3 yrs from open‐access motion‐detecting camera trap photos. Using semi‐permeable mammal exclosures and thread‐marked seeds, predation and dispersal by mammals for each seed species were quantified. Abundances of peccaries were up to six times higher than those of agoutis over 3 yrs, but neither peccary nor agouti abundances differed across years. Seeds of A. alatum were predominantly dispersed by peccaries, which did not eat A. alatum seeds, whereas non‐defended and chemically defended seeds suffered high levels of predation, mostly by peccaries. Agoutis did not eat M. holtonii seeds. Peccaries and agoutis did not differ in the distances they dispersed seeds. This study shows that seed fates are contingent upon many factors such as seed defenses, frugivore–granivore abundances, and seed‐handling capabilities. Mammal–seed interactions are complex; the outcomes of these interactions depend on the inherent characteristics of seeds and their potential dispersers.     

Primate and Dung Beetle Communities in Secondary Growth Rain Forests: Implications for Conservation of Seed Dispersal Systems

Conservation efforts are often aimed at one or a few species. However, habitat sustainability relies on ecological interactions among species, such as seed dispersal. Thus, a community-scale conservation strategy may be more valuable in some settings. We describe communities of primary (primates) and secondary (dung beetles) seed dispersers from 5 sites in the Brazilian Amazon. We estimate community biomass of these taxa and, using multivariate ordination, examine the potential for natural reforestation at each site, given the communities of seed dispersers present. Since disturbed habitat is increasingly common and increasingly the focus of conservation efforts, we also examine differences among seed disperser communities between primary forest and secondary growth at each site. Analyses of faunal biomass in different localities and habitats indicate that secondary growth receives nearly as much use by primates as primary forest; given the dominant groups of dung beetles in secondary growth, disturbed habitat should show a pattern of seed burial that is clumped and deep. Areas with high biomass of Alouatta spp. and the large nocturnal dung beetle species may have the greatest potential for natural reforestation of secondary growth particularly for large seeded species. The data suggest that knowledge of the biomass of primary and secondary dispersing fauna facilitates predictions for the likelihood of disturbed habitat to regenerate and comparisons of sites in broader geographical areas e.g., Neotropical vs. Paleotropical forests.     

Effects of Selective Vegetation Thinning on Seed Removal in Secondary Forest Succession1

Seed removal was assessed for two tree species in three forest types: (1) secondary forest with and (2) without selective vegetation thinning, and (3) mature forest. Selective vegetation thinning meant the removal of all stems ≤3 cm in diameter of secondary‐forest species and was intended as a management technique to accelerate succession toward mature forest. Thinning did not have an effect on seed removal. One of the species showed lower seed removal in mature forest compared to secondary forest.     

Feeding Behavior of Purple‐throated Fruitcrow (Querula purpurata: Cotingidae) in the Colombian Amazon and Its Implications for Seed Dispersal1

We studied some feeding behaviors of the purple‐throated fruitcrow (Querula purpurata) in two Colombian Amazonian forests, which affect the primary seed dispersal of the plants on which it feeds. Visit length, number of fruits removed and dispersed, feeding rates, and fruit‐handling times were compared to those obtained for two other cotingas feeding on the same fruiting trees. Querula purpurata exhibited shorter visits (98 sec) and fruit‐handling times (4 sec), and higher mean feeding rates (1.6 fruits/min) than Phoenicircus nigricollis and Cotinga cayana. In contrast, P. nigricollis dispersed the highest number of seeds of four of the five tree species studied. Although Q. purpurata and P. nigricollis exhibited feeding behaviors that increase seed dispersal, Q. purpurata may be more important in the transport of seeds between habitats, while P. nigricollis may be a major seed disperser within the primary forest.     

Chimpanzee (Pan troglodytes) Seed Dispersal in an Afromontane Forest: Microhabitat Influences on the Postdispersal Fate of Large Seeds1

We examined the postdispersal fate of large seeds (≥5 mm) dispersed by chimpanzees in an afromontane forest to evaluate aspects of the effectiveness of seed dispersal by chimpanzees, Pan troglodytes. We assessed the influence of six microhabitat characteristics on seed persistence and germination in seeds dispersed in chimpanzee feces and “wadges.” A total of 257 fecal samples and 56 wadges were located over a 4‐mo period by tracking a semi‐habituated chimpanzee community on day follows. Forty‐nine (19.1%) of the fecal samples contained large seeds from five different tree species. The majority of fecal samples with seeds contained seeds from the mature forest tree Olea capensis (Oleaceae) (83.7%). Forty‐two wadges (75%) contained seeds from the mature forest tree Syzygium guineense (Myrtaceae). Seeds were monitored at their deposition site for removal and germination up to 49 d following deposition. We collected data on the microhabitat surrounding each fecal and wadge sample. Multivariate analyses indicated that while fecal and wadge samples were not clustered into particular microhabitats, there was little overlap in the microhabitats in which wadges and fecal samples were deposited. Significantly more seeds persisted over 49 d in wadges (67.9%) than in feces (30.3%). Elevation was the only microhabitat variable determined to have a significant influence on seed persistence, whereas slope was determined to have a significant influence on germination.     

Frugivory and Seed Dispersal by Brown Lemurs in a Malagasy Tropical Dry Forest

In the Ankarafantsika tropical dry forest (northwestern Madagascar), the common brown lemur (Eulemur fulvus fulvus) is the largest frugivore and probably the sole disperser of large‐seeded plants (seed diameter > 10 mm). To investigate seed dispersal by this primate, I recorded the feeding activities of a troop; also conducted fecal analyses, germination trials on defecated seeds, and a vegetation survey over 1 yr (beginning Dec 2006). Brown lemurs mostly consumed fruit (68%). The fruit of Vitex beraviensis was the most exploited resource (21% of total feeding time). Among dung samples, 1126 contained intact seeds of 70 plant species, with a median of six seeds and two species per sample. These data indicate that the brown lemur population dispersed approximately 9854 seeds/km²/d. Although the number of annually defecated seeds was overwhelmingly the largest in Grewia triflora, many of the small seeds were often clumped in dung piles. In contrast, large seeds of V. beraviensis occurred in the largest number of dung samples. The rate and time of seed germination in V. beraviensis were improved by passage through brown lemur guts. Therefore, V. beraviensis may readily establish seedlings in sites of brown lemur fecal deposition. Vitex beraviensis and brown lemurs are probably involved in a strong mutualism. Twenty‐three large‐seeded plants were probably dependent on brown lemurs for seed dispersal and some of these species were common trees in the forest. Maintenance of these key plant–animal interactions will probably contribute to the conservation of species diversity and intact regeneration of the Ankarafantsika forest.     

Dung Beetle Communities and Seed Dispersal in Primary Forest and Disturbed Land in Amazonia1

Seeds from tropical fruiting trees ingested and defecated on the soil surface by primary dispersers (such as primates) are vulnerable to destruction from rodents, insects, and fungi. Burial by dung beedes as an incidental result of their feeding and nesting activities often provides these seeds with refugia from attack. To examine the effect of habitat disturbance on the dung beetle communities involved in this process, I surveyed dung beedes at three sites in the Amazon basin, in die states of Parí, Amazonas, and Rondónia, Brazil. Through principal component analysis on measurements of size and behavioral characters of beedes, I determined the relative quality of beetle species as seed dispersers (dispersal defined as horizontal or vertical movement of seeds) and ranked them into seed dispersal guilds. I used correspondence analysis to examine in what habitats (primary forest or varying degrees of disturbed habitat) these guilds were abundant. Most guilds decreased with increasing habitat disturbance, but one guild made up of large nocturnal burrowers (primarily Dichotomius) became more abundant with increasing disturbance (up to the level of highest disturbance surveyed), at which point all dung beede species became scarce. Clear–cuts had lower species richness, lower abundance, and lower biomass dian forested areas. These results imply that clear–cutting is detrimental to all seed dispersal dynamics in tropical rain forests, but that some levels of disturbance allow enough disperser activity to preserve this ecosystem function.     

Seed Dispersal and Demography of Pioneer Trees: The Case of Hortia arborea

Abstract: This study investigates the relationships among seed dispersal, patterns of seedling recruitment and the spatial distribution of a pioneer tree (Hortia arborea, Rutaceae) in the Brazilian Atlantic forest. The study was carried out at Dois Irmãos Reserve, a 387.4 ha reserve in northeastern Brazil. Fruit shadow concentrated beneath parental trees (74.6 % < 6 m distant) and fruit removal averaged 5.6 % (0 ‐ 25.5 %). Agoutis (Dasyprocta prymnolopha, Rodentia) ate fruits in the forest and in captivity and were the only vertebrates recorded feeding on Hortia fruits. Agoutis, however, destroyed 86.7 % of the seeds they ingested. In addition, only one seed from an expected number of 1980 germinated in a treefall gap after it passed through the digestive tract of agoutis. Both seed germination and seedling recruitment were restricted to gaps, and occurred among seeds manually released from the pulp or among those from fruits that naturally rotted in gaps. Moreover, exposed seedlings were taller than those covered by plants in a gap. H. arborea appears to self‐maintain populations in the same patches of forest, which are delimited by a pool of old and newly created gaps. More specifically, there is successful recruitment in patches occupied by parents, resulting in dense clumps of H. arborea.     

不同年龄香果树种子雨和种子库及其更新特征

对宝天曼自然保护区香果树种子雨、土壤种子库进行了观测,并进行不同微生境播种实验,研究其种子萌发及幼苗生长动态。结果表明:(1)不同龄级香果树种子雨持续时间及其高峰期有所不同,种子雨强度存在极显著差异。(2)20~50龄级香果树的种子饱满率、土壤种子库中种子密度均显著低于其他龄级,但其不同类型种子的水平和垂直分布规律一致;大约80%的香果树种子在其萌发前消失,剩余种子中以霉烂种子密度最高(9.81粒·m-2),饱满种子密度最低(1.94粒·m-2)。(3)野外育苗实验表明,不同龄级香果树所产饱满种子的萌发率及幼苗存活率差异不显著,其种子萌发率约为16.93%,但仅有3.86%的幼苗寿命超过5个月;不同微生境对香果树幼苗存活率产生显著影响,林窗是其最适宜微生境。研究认为,人工采集香果树种子,于次年4月份在原生境下播种;清理母树冠下和冠缘的地被物,增加林缘空地地被物覆盖度,以减少香果树种子的损失,改善香果树种子萌发和幼苗生长的光照条件,以利于幼苗根系的生长,促进香果树的自然更新。