Seasonal rhythms of seed rain and seedling emergence in two tropical rain forests in southern Brazil

Seasonal tropical forests show rhythms in reproductive activities due to water stress during dry seasons. If both seed dispersal and seed germination occur in the best environmental conditions, mortality will be minimised and forest regeneration will occur. To evaluate whether non-seasonal forests also show rhythms, for 2 years we studied the seed rain and seedling emergence in two sandy coastal forests (flooded and unflooded) in southern Brazil. In each forest, one 100 x 30-m grid was marked and inside it 30 stations comprising two seed traps (0.5 x 0.5 m each) and one plot (2 x 2 m) were established for monthly monitoring of seed rain and a seedling emergence study, respectively. Despite differences in soil moisture and incident light on the understorey, flooded and unflooded forests had similar dispersal and germination patterns. Seed rain was seasonal and bimodal (peaks at the end of the wetter season and in the less wet season) and seedling emergence was seasonal and unimodal (peaking in the wetter season). Approximately 57% of the total species number had seedling emergence 4 or more months after dispersal. Therefore, both seed dormancy and the timing of seed dispersal drive the rhythm of seedling emergence in these forests. The peak in germination occurs in the wetter season, when soil fertility is higher and other phenological events also occur. The strong seasonality in these plant communities, even in this weakly seasonal climate, suggests that factors such as daylength, plant sensitivity to small changes in the environment (e.g. water and nutrient availability) or phylogenetic constraints cause seasonal rhythms in the plants.     

The role of avian ‘seed predators’ as seed dispersers

Seed dispersal is a central process in plant ecology with consequences for species composition and habitat structure. Some bird species are known to disperse the seeds they ingest, whereas others, termed ‘seed predators’, digest them and apparently play no part in dispersal, but it is not clear if these are discrete strategies or simply the ends of a continuum. We assessed dispersal effectiveness by combining analysis of faecal samples and bird density. The droppings of seed dispersers contained more entire seeds than those of typical seed predators, but over a quarter of the droppings of seed predators contained whole seeds. This effect was further magnified when bird density was taken into account, and was driven largely by one frequent interaction: the Chaffinch Fringilla coelebs, a typical seed predator and the most abundant bird species in the area and dispersed seeds of Leycesteria formosa, a non‐native plant with berry‐like fruits. These results suggest the existence of a continuum between seed predators and seed dispersers.     

Dung beetles in a Central Amazonian rainforest and their ecological role as secondary seed dispersers

Abstract 1. The role of several factors that affect the composition of the dung beetle assemblages in an Amazonian rainforest was quantified, together with the effect of these factors on the role that dung beetles play as secondary seed dispersers. 2. A total of 61 dung beetle species was captured during 3360 h of trapping. During nocturnal trapping periods, more dung beetles, of larger mean size, and more species were captured per trap than during diurnal trapping periods. 3. During the rainy season, more dung beetle species were captured per trap than during the dry season, but the number of individuals and their mean size did not vary between seasons. 4. Bait size had a significant effect on the mean number of beetles and mean number of species but not on mean beetle size. As bait size increased from 5, 10, 25, to 50 g, more beetles and more species were captured per trap. 5. Between 6 and 73% of plastic beads, used as seed mimics, were buried by dung beetles at depths that ranged from 0.5 to 7 cm. Both the proportion of beads buried and burial depth decreased with increasing bead size, and increased with increasing amounts of dung surrounding each bead (5, 10, and 25 g). 6. The proportion of buried seeds for three species varying in size between 5 and 27 mm, increased with increasing dung beetle size, using beetles of seven sizes, varying between 10 and 25 mm. 7. Seeds surrounded by dung were buried more often and more deeply when placed on the forest floor during the late afternoon than when placed during the early morning. Seeds were also buried more often when placed on the forest floor during the rainy season than when placed during the dry season, but season had no effect on burial depth. 8. Forests in Central Amazonia hold a rich dung beetle community that plays an active role in secondary seed dispersal, and consequently in plant regeneration. The interaction between seeds and beetles is complex because it is affected by many factors.     

Reliability of macaques as seed dispersers

Seed dispersal is an ecological process crucial for forest regeneration and recruitment. To date, most studies on frugivore seed dispersal have used the seed dispersal effectiveness framework and have documented seed-handling mechanisms, dispersal distances and the effect of seed handling on germination. In contrast, there has been no exploration of “disperser reliability” which is essential to determine if a frugivore is an effective disperser only in particular regions/years/seasons or across a range of spatio-temporal scales. In this paper, we propose a practical framework to assess the spatial reliability of frugivores as seed dispersers. We suggest that a frugivore genus would be a reliable disperser of certain plant families/genera if: (a) fruits of these plant families/genera are represented in the diets of most of the species of that frugivore, (b) these are consumed by the frugivore genus across different kinds of habitats, and (c) these fruits feature among the yearly staples and preferred fruits in the diets of the frugivore genus. Using this framework, we reviewed frugivory by the genus Macaca across Asia to assess its spatial reliability as seed dispersers. We found that the macaques dispersed the seeds of 11 plant families and five plant genera including at least 82 species across habitats. Differences in fruit consumption/preference between different groups of macaques were driven by variation in plant community composition across habitats. We posit that it is essential to maintain viable populations of macaques across their range and keep human interventions at a minimum to ensure that they continue to reliably disperse the seeds of a broad range of plant species in the Anthropocene. We further suggest that this framework be used for assessing the spatial reliability of other taxonomic groups as seed dispersers.     

Characterization of seven microsatellite marker loci in a genus of Malagasy lemurs (Propithecus)

The sifakas (genus Propithecus) are large, diurnal, lemurs endemic to the island of Madagascar. Throughout the island, lemur species are particularly vulnerable to extinction due to relatively small geographical ranges (Martin 2000). Thus, understanding the taxonomy of these highly endangered primates is essential for taking appropriate conservation action. For this study we collected tissue samples from 68 wild sifakas representing various Propithecus species. Genomic DNA was isolated and polymerase chain reaction (PCR) amplification was carried out. We describe seven microsatellites found to be polymorphic and useful for analysis of the population genetics of sifakas.     

Small mammals as potential seed dispersers in New Zealand

Weed invasion success is strongly influenced by availability of seed dispersal vectors, which may include animals. We examined the potential of several small introduced mammals (mice, kiore, ship rats and possums) to disperse germinable seeds in New Zealand. Captive animals were fed fleshy fruit of weeds (Berberis glaucocarpa, Cotoneaster spp., Crataegus monogyna, Ilex aquifolium, Leycesteria formosa, Ligustrum sinense, Lonicera japonica, Passiflora mollissima, Pyracantha angustifolia, Sorbus hupehensis) and native species (Coprosma spp., Prumnopitys ferruginea and Solanum aviculare). We recorded the percentage of fruit consumed, seed ingested and gut passage time. Faeces were collected and the seeds extracted and tested for germination potential in an unheated glasshouse (two weed species) or under controlled conditions (11 species). The smallest rodents (mice and kiore) generally destroyed all seeds eaten. Large numbers of viable seeds of the small‐seeded (<1 mg) species, L. formosa and S. aviculare, passed through ship rats. Possums consumed the seeds of all adventive and native fruits except P. ferruginea. The proportion of seeds recovered intact from possum faeces varied with plant species and ranged from 6 to 83%. The time required for 50% of all seeds to be passed by possums ranged from 2.5 to 5.5 days with an average of 3.7 days, and was generally unrelated to simple fruit parameters such as percentage pulp and moisture content. For seeds where germination also occurred in the uneaten controls, the germination of seed from possums ranged from 3 to 78%. Germination was mostly lower in seeds from possums than in the controls, where differences were significant. Possums have major potential to disperse a wide range of fleshy fruit‐producing native and introduced plant species. Ship rats have the potential to disperse those with very small seeds.     

Leaf-cutting ant nests support less dense and impoverished seed assemblages in a human-modified Caatinga dry forest

Regenerating forests make up an increasingly large portion of tropical landscapes worldwide and regeneration dynamics may be influenced by leaf-cutting ants (LCA), which proliferate in disturbed areas and collect seeds for fungus culturing. Here, we investigate how LCA influences seed fate in human-modified areas of Caatinga dry forest. We evaluate the seed deposition and predation on Atta opaciceps nests, foraging habitat surrounding nest and control habitat away of nest influence of 15 colonies located along a forest cover gradient during the rainy and dry seasons. For each habitat, four 50-cm² plots were established and all seeds on the soil surface were collected along 1 year. We recorded 13,628 seeds distributed among 47 species and 36.57% of the total seeds did not show any sign of predation. Nest mound habitats supported low-density and species-poor seed assemblages, which were taxonomically distinct from the control habitats. These effects only occurred in the rainy season. The proportion of undamaged seeds were similar across the habitats. While forest cover did not influence seed assemblage in terms of species richness or seed predation, it did interact with habitat type via increments in seed abundance as forest cover increased across the nests. Forest cover also affected seed composition, but only in the rainy season. These results indicate that LCA decrease seed deposition in areas under their influence, particularly on the nest mounds. As LCA profit from human disturbance in the Caatinga, their role as seed ‘sinks’ should be enhanced in disturbed Caatinga patches, particularly during the rainy season, when most of the plant recruitment occurs. Our findings reinforce the importance of LCA as drivers of forest dynamics and resilience in human-modified landscapes.     

Feeding ecology and seed dispersal of sympatric cheirogaleid lemurs (Microcebus murinus, Cheirogaleus medius, Cheirogaleus major) in the littoral rainforest of south-east Madagascar

The lemurs of Madagascar are known for their extraordinary species diversity. The mechanisms that allow the coexistence of these species are still poorly known. Here feeding patterns were investigated for three small nocturnal lemur species of Cheirogaleidae ( Microcebus murinus , Cheirogaleus medius and Cheirogaleus major ) occurring sympatrically in a littoral rainforest in south-east Madagascar. During three rainy seasons, the plant species eaten by these three lemurs were described in relation to morphological and biochemical characteristics. All three species were mainly frugivorous and fed on 68 different plant species with small- and medium-sized fruits. A total of 91% of these forage plant species was visited by all three lemur species. Fruits larger than 25–30 mm were avoided. Seeds of a total of 51 food plant species were swallowed and passed the gut unharmed. Thus, even these smaller lemur species play an important role in seed dispersal. There were no differences in the morphological and biochemical characteristics of fruits eaten between the three species, but the feeding height was significantly different between the species. Thus, competition avoidance and niche separation are presumably not based on different feeding patterns of M. murinus , C. medius and C. major in the littoral rainforest, but on different habitat utilization.     

Bushmeat and the Fate of Trees with Seeds Dispersed by Large Primates in a Lowland Rain Forest in Western Amazonia

In Neotropical forests, large fruit-eating primates play important ecological roles as dispersal agents of large seeds. Bushmeat hunting threatens to disrupt populations of primates and large-seeded trees. We test the hypothesis that otherwise intact Neotropical forests with depressed populations of large primates experience decline in recruitment of large-seeded trees. We quantify the proportion of small juveniles (> 0.5 m tall–1 cm diameter at breast height, DBH) of large primate-dispersed tree species found underneath heterospecifc trees that are also dispersed by large primates at two protected sites in Manu National Park and one hunted site outside Manu N.P. in southeastern Peru. The forests are comparable in edaphic and climatic qualities, successional stage, and adult tree species composition. We found that hunting locally exterminates populations of large primates, and reduced primates of intermediate body size (hereafter "medium primates") by 80 percent. Moreover, tree species richness was 55 percent lower and density of species dispersed by large and medium-bodied primates 60 percent lower in hunted than in protected sites. In addition, richness and density of abiotically dispersed species and plants dispersed by non-game animals are greater in hunted sites. Overhunting threatens to disrupt the ecological interactions between primates and the plants that rely on them for seed dispersal and recruitment. Sustainable wildlife management plans are urgently needed, because protected areas are at risk of becoming "island" parks if buffer zones become empty of animals and have impoverished flora.     

西双版纳热带森林土壤种子库的季节变化

通过萌发实验法对西双版纳地区的一类热带季节雨林 (番龙眼、千果榄仁 )和 2类次生林(白背桐林、中平树林 )的土壤种子库的季节变化进行了探讨。结果表明 :该地区的土壤种子库动态具有明显的季节性。季节雨林的土壤种子库储量相对稳定 ,土壤上层 ( 0～ 2 cm )的种子储量在雨季末期较大。2类次生林土壤种子库的变化则相反 ,土壤种子库中的种子种类在旱季末期较雨季末期多 ,土壤上层的种子储量在旱季末期较大。各样地均有一些种类只出现在旱季末期或雨季末期。种子在土壤种子库的动态与植物的繁殖物候和所处的环境紧密相关 ,不同种类植物的土壤种子库由于植物本身的生物学特性、传播方式和所处环境的影响而表现出不同的动态模式     

The role of seed dispersal in the natural regeneration of rain forest after strip-cutting in the Peruvian Amazon

Seed dispersal and forest regeneration were studied on a 30×150 m strip cleared by strip-cutting, a system of forest management designed for sustained yield (Hartshorn 1989), in high terrace rain forest in the Department of Loreto, Peru. After one year the strip was dominated by seedlings of a few bat- and bird-dispersed pioneer tree species (Cecropia spp., Melastomataceae, and Alchornea triplinervia); stump sprouts from cut trees and saplings that survived the clearing were less numerous. The density of saplings (>2 m in height) surviving the clearing was 903 per hectare; 94% of these survived the subsequent 18 months. About 30% of 417 stumps (>7.5 cm diameter at breast height) resprouted within 3 months, with an additional 10% sprouting in the subsequent 10 months. Sprouting frequency was greater for small stumps than large and varied greatly among plant families. Seed deposition over this year was much lower in the interior of the strip, both in species richness and numbers of seeds, than within the forest; strip edges were intermediate in richness and number. The decline in seed input from forest to edge to strip, both in species and in numbers of seeds, was most pronounced for bird-dispersed taxa (primarily Melastomataceae); bat- and wind- dispersed taxa were more evenly distributed. The similarity in bat species composition between the strip and nearby primary forest was higher than the similarity in bird species composition between these habitats, reflecting a failure of many forest bird species to venture into the strip. The predominance of Cecropia spp. and other pioneers of minimal commercial value in the regeneration question the sustainability of strip-cutting. Subsequent succession and future tree species composition on the cleared strip will depend not only on the survivorship and growth of sprouts, survivors, and seedlings, but also on responses of different seed-dispersing animal taxa to changes in the species composition and structure of the vegetation in the strip.     

The insect‐focused classification of fruit syndromes in tropical rain forests: An inter‐continental comparison

We propose a new classification of rain forest plants into eight fruit syndromes, based on fruit morphology and other traits relevant to fruit‐feeding insects. This classification is compared with other systems based on plant morphology or traits relevant to vertebrate fruit dispersers. Our syndromes are based on fruits sampled from 1,192 plant species at three Forest Global Earth Observatory plots: Barro Colorado Island (Panama), Khao Chong (Thailand), and Wanang (Papua New Guinea). The three plots differed widely in fruit syndrome composition. Plant species with fleshy, indehiscent fruits containing multiple seeds were important at all three sites. However, in Panama, a high proportion of species had dry fruits, while in New Guinea and Thailand, species with fleshy drupes and thin mesocarps were dominant. Species with dry, winged seeds that do not develop as capsules were important in Thailand, reflecting the local importance of Dipterocarpaceae. These differences can also determine differences among frugivorous insect communities. Fruit syndromes and colors were phylogenetically flexible traits at the scale studied, as only three of the eight seed syndromes, and one of the 10 colors, showed significant phylogenetic clustering at either genus or family levels. Plant phylogeny was, however, the most important factor explaining differences in overall fruit syndrome composition among individual plant families or genera across the three study sites. Abstract in Melanesian is available with online material.     

Carnivorous mammals as seed dispersers of Myrtus communis (Myrtaceae) in the Mediterranean shrublands

ABSTRACT

The role of carnivorous mammals as seed dispersers of Myrtus communis L. in the Mediterranean environment was studied. Seeds of M. communis were found in faeces of red fox (Vulpes vulpes L.) and carnivores from the genus Martes, at four sites on the Tyrrhenic coast of Southern Italy in the period autumn-winter 1993–1995. All the study sites are covered by Mediterranean shrublands with M. communis as one of the dominant species. Content analysis of fox scats showed that fruits of M. communis constitute a relevant component of the diet of these carnivores, although other food categories were also well represented. Further investigations on the effect of mammal digestion on seed germinability showed that carnivores not only moved the seeds away from the mother plant but also enhanced their germinability. This species was known to be bird- and ant-dispersed and the adaptive implications of a multi-vectorial seed dispersal process is discussed.     

Lowland tapirs facilitate seed dispersal in degraded Amazonian forests

The forests of southeastern Amazonia are highly threatened by disturbances such as fragmentation, understory fires, and extreme climatic events. Large‐bodied frugivores such as the lowland tapir (Tapirus terrestris) have the potential to offset this process, supporting natural forest regeneration by dispersing a variety of seeds over long distances to disturbed forests. However, we know little about their effectiveness as seed dispersers in degraded forest landscapes. Here, we investigate the seed dispersal function of lowland tapirs in Amazonian forests subject to a range of human (fire and fragmentation) and natural (extreme droughts and windstorms) disturbances, using a combination of field observations, camera traps, and light detection and ranging (LiDAR) data. Tapirs travel and defecate more often in degraded forests, dispersing much more seeds in these areas [9,822 seeds per ha/year (CI_95% = 9,106; 11,838)] than in undisturbed forests [2,950 seeds per ha/year (CI_95% = 2,961; 3,771)]. By effectively dispersing seeds across disturbed forests, tapirs may contribute to natural forest regeneration—the cheapest and usually the most feasible way to achieve large‐scale restoration of tropical forests. Through the dispersal of large‐seeded species that eventually become large trees, such frugivores also contribute indirectly to maintaining forest carbon stocks. These functions may be critical in helping tropical countries to achieve their goals to maintain and restore biodiversity and its ecosystem services. Ultimately, preserving these animals along with their habitats may help in the process of natural recovery of degraded forests throughout the tropics. Abstract in Portuguese is available with online material.     

The effect of seed mass and gap size on seed fate of tropical rain forest tree species in guyana

Abstract: For eleven tree species, differing in seed mass, germination success (emergence success for two small-seeded species) and the causes of failure to germinate were studied in the forest understorey and in logging gaps in the tropical rain forests of Guyana. In the forest understorey, germination success increased with seed mass. However, as gap size increased the difference between smaller and larger seeded species diminished because germination success of smaller-seeded species increased slightly, while that of larger-seeded species decreased dramatically. The negative effect of gap size on germination success of larger-seeded species was caused by an increased risk of desiccation with gap size, which was a far more important seed mortality agent for larger than for smaller-seeded species. Generally, seeds of smaller-seeded species suffered more from insect predation and were removed at higher rates than larger-seeded species. On the other hand, larger-seeded species were eaten more by mammals than smaller-seeded species. It is concluded that logging can result in shifts in the species composition in the tropical rain forests of Guyana which are dominated by species with large seeds, since germination success of larger-seeded species is dramatically reduced in large logging gaps.     

Tropical rain forest fragmentation,howler monkeys (Alouatta palliata), and dung beetles at Los Tuxtlas,Mexico

In Neotropical rain forests, fresh mammal dung, especially that of howler monkeys, constitutes an important resource used by dung beetles as food and for oviposition and further feeding by their larvae. Tropical rain forest destruction, fragmentation, and subsequent isolation causing reductions in numbers of and the disappearance of howler moneys may result in decreasing numbers of dung beetles, but this has not been documented. In this study, we present information on the presence of howlers and dung beetles in 38 isolated forest fragments and 15 agricultural habitats. Howler monkeys were censused by visual means, while dung beetles were sampled with traps baited with a mixture of howler, cow, horse, and dog dung. Results indicated that loss of area and isolation of forest fragments result in significant decrements in howlers and dung beetles. However, dung beetle abundance was found to be closely related to the presence of howler monkeys at the sites and habitats investigated. Scenarios of land management designed to reduce isolation among forest fragments may help sustain populations of howler monkeys and dung beetles, which may have positive consequences for rain forest regeneration. Am. J. Primatol. 48:253–262, 1999. © 1999 Wiley‐Liss, Inc.