The consumption of figs (the fruit of Ficus spp.; Moraceae) by vertebrates is reviewed using data from the literature, unpublished accounts and new field data from Borneo and Hong Kong. Records of frugivory from over 75 countries are presented for 260 Ficus species (approximately 30% of described species). Explanations are presented for geographical and taxonomic gaps in the otherwise extensive literature. In addition to a small number of reptiles and fishes, 1274 bird and mammal species in 523 genera and 92 families are known to eat figs. In terms of the number of species and genera of fig-eaters and the number of fig species eaten we identify the avian families interacting most with Ficus to be Columbidae, Psittacidae, Pycnonotidae, Bucerotidae, Sturnidae and Lybiidae. Among mammals, the major fig-eating families are Pteropodidae, Cercopithecidae, Sciuridae, Phyllostomidae and Cebidae. We assess the role these and other frugivores play in Ficus seed dispersal and identify fig-specialists. In most, but not all, cases fig specialists provide effective seed dispersal services to the Ficus species on which they feed. The diversity of fig-eaters is explained with respect to fig design and nutrient content, phenology of fig ripening and the diversity of fig presentation. Whilst at a gross level there exists considerable overlap between birds, arboreal mammals and fruit bats with regard to the fig species they consume, closer analysis, based on evidence from across the tropics, suggests that discrete guilds of Ficus species differentially attract subsets of sympatric frugivore communities. This dispersal guild structure is determined by interspecific differences in fig design and presentation. Throughout our examination of the fig-frugivore interaction we consider phylogenetic factors and make comparisons between large-scale biogeographical regions. Our dataset supports previous claims that Ficus is the most important plant genus for tropical frugivores. We explore the concept of figs as keystone resources and suggest criteria for future investigations of their dietary importance. Finally, fully referenced lists of frugivores recorded at each Ficus species and of Ficus species in the diet of each frugivore are presented as online appendices. In situations where ecological information is incomplete or its retrieval is impractical, this valuable resource will assist conservationists in evaluating the role of figs or their frugivores in tropical forest sites.

Fig-eating by vertebrate frugivores: a global review.

People have hunted mammals in tropical Asian forests for at least 40,000 yr. This period has seen one confirmed global extinction (the giant pangolin, Manis palaeojavanica) and range restrictions for several large mammals, but there is no strong evidence for unsustainable hunting pressure until the last 2000‐3000 yr, when elephants, rhinoceroses, and several other species were progressively eliminated from the large parts of their ranges. Regional declines in most species have occurred largely within the last 50 yr. Recent subsistence hunting has typically focused on pigs and deer (hunted with dogs and spears or with snares), monkeys and other arboreal mammals (often caught with blowpipes), and porcupines and other rodents (smoked or dug out of burrows). Over the last 50 yr, the importance of hunting for subsistence has been increasingly outweighed by hunting for the market. The hunted biomass is dominated by the same species as before, sold mostly for local consumption, but numerous additional species are targeted for the colossal regional trade in wild animals and their parts for food, medicines, raw materials, and pets. Many populations of mammalian dispersers of large seeds and understory browsers have been depleted or eliminated, while seed predators have had a more variable fate. Most of this hunting is now illegal, but the law enforcement is generally weak. However, examples of successful enforcement show that hunting impacts can be greatly reduced where there is sufficient political will. Ending the trade in wild animals and their parts should have the highest regional conservation priority. HABITAT LOSS AND DEGRADATION are such massive and visible threats to biodiversity in tropical Asia that the impact of hunting is sometimes considered secondary, at least in comparison with Africa and the Neotropics (Primack & Corlett 2005, Sodhi & Brook 2006). However, hunting impacts almost all remaining forest in the region and few areas now support the vertebrate fauna that they potentially could if hunting were prevented. This reduction in vertebrate populations may, in turn, slow vegetation recovery through its impact on seed dispersal, particularly since hunters favor the large vertebrate species that disperse large fruits with large seeds (Corlett 1998, 2002). Hunting, moreover, is potentially more easily controlled than most other conservation problems in tropical Asian forests. In most areas it is a relatively marginal economic activity, involving relatively few people. Hunting has become a massive problem because of high human population densities and a generally well-developed infrastructure that not only makes most forest areas easily accessible, but also gives access to distant urban markets for luxury (often medicinal) products. This review covers the hunting (including trapping) of mammals in closed-canopy forest habitats of tropical and subtropical Asia west of Wallace’s line, i.e., the Oriental or Indomalayan region. This region excludes eastern Indonesia, with its very different forest fauna, and much of the more arid western part of the Oriental region. The area covered has a relatively uniform mammalian fauna at the genus and family level, with many widespread species. Information on hunting impacts was obtained from several sources, including: archaeological data; studies of contemporary hunters; observations of the wildlife trade and markets; and changes in forest faunas over time. Mammalian nomenclature follows Wilson and

The Impact of Hunting on the Mammalian Fauna of Tropical Asian Forests

Introduction. Ecologically Meaningful Germination Studies. Types of Seed Dormancy. Germination Ecology of Seeds with Nondeep Physiological Dormancy. Germination Ecology of Seeds with Morphophysiological Dormancy. Germination Ecology of Seeds with Physical Dormancy. Germination Ecology of Seeds in the Persistent Seed Bank. Causes of Within-Species Variations in Seed Dormancy and Germination Characteristics. A Geographical Perspective on Germination Ecology: Tropical and Subtropical Zones. A Geographical Perspective on Germination Ecology: Temperate and Arctic Zones. Germination Ecology of Plants with Specialized Life Cycles and/or Habitats. Biogeographical and Evolutionary Aspects of Seed Dormancy. Subject Index.

A Definitive Book on Seed Ecology@@@Seeds: Ecology, Biogeography, and Evolution of Dormancy and Germination

Seed dispersal in changing landscapes.

Effects of sika deer on vegetation in Japan: A review

Large frugivores are considered to be important seed dispersers for many tropical plant species. Their roles as seed dispersers are not well known in Southeast Asia, where degraded landscapes typically lack these animals. Interactions between 259 (65 families) vertebrate-dispersed fruits and frugivorous animals (including 7 species of bulbul, 1 species of pigeon, 4 species of hornbill, 2 species of squirrel, 3 species of civet, 2 species of gibbon, 1 species of macaque, 2 species of bear, 2 species of deer, and 1 species of elephant) were studied for 3 years in a tropical seasonal forest in Khao Yai National Park, Thailand. The purpose was to examine the dietary overlaps among the large frugivores and the characteristics of fruits they consumed. Most fruit species are eaten by various kinds of frugivores; no close relationship between a particular fruit and a frugivore was found. The number of frugivore groups that served a given plant species was negatively correlated with seed size. Additionally, the fruit/seed diameters consumed by bulbuls were significantly smaller than consumed by the other nine groups. These trends of fruit characteristics were consistent with those observed elsewhere in Southeast Asia: small fruits and large, soft fruits with many small seeds are consumed by a wide spectrum of frugivores while larger fruits with a single large seed are consumed by relatively few potential dispersers. Importantly, these large, single-seed fruits are not consumed by the small frugivores that thrive in small forest fragments and degraded areas in Southeast Asia. To insure the natural seed dispersal process in the forest, an evaluation of all frugivore groups in the forest is urgently needed in Southeast Asia.

Interactions between fleshy fruits and frugivores in a tropical seasonal forest in Thailand

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima, southwestern Japan, were studied for 2 years by means of fecal analysis. The altitudinal range of fecal samples collected was 30 m to 1,203 m above sea level, and it was divided into three zones: low-zone forest (0–399 m), middle-zone forest (400–799 m), and high-zone forest (800 m–1,230 m). There was a considerable altitudinal and seasonal variation in the macaques' diet. Seed/fruit and animal matter were eaten more in the lower zones, whereas more fiber and fungi were consumed in the higher zones. In all of the zones, they ate seed/fruits the most in autumn (September–November) and the least in spring (March–April). They ate fibrous food the most in spring and the least in autumn. Macaques relied on seed/fruits heavily in the lower zone for a longer period than in the higher zones. Macaques in the high-zone forest ate almost no seed/fruit foods from March to May. Altitudinal variations in availability of seed/fruit foods seem to have influenced the altitudinal variations in diet. Total basal area of seed/fruit-food trees, species richness of seed/fruit-foods, main seed/fruit-food types available, and annual fleshy-fruit production all decreased with increasing altitude. Both interannual variation and annual cyclicity of diet were found in all zones.

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima.

Among wild Japanese macaques, which have clear reproductive seasonality, correspondence between fruit-food production in the mating season and birth rate in the following year was confirmed in two different habitats. One of the study areas was evergreen broad leaved forest on Yakushima Island, for which demographic and fruiting data for seven years were used. The other was a deciduous-coniferous mixed forest on Kinkazan Island in the cool temperate zone, for which 11 years of data were used. From the fruit-crop data, each year was classified as a good or bad fruiting year for each population. At both habitats, female macaques had fewer babies after bad fruiting years than after good fruiting years. In Yakushima, small troops had a lower birth rate than large troops and this tendency was clear after bad fruiting but not after good fruiting. On the other hand, in Kinkazan such differences due to troop size were not found. These findings were consistent with the observation that intertroop encounters occur more often and are more agonistic in Yakushima than in Kinkazan and large troops tend to be dominant to small troops in the Yakushima population. Thus annual fluctuations in fruit production appear to increase the difference in birth rates between troops of different sizes through intertroop competition in Yakushima, but not in Kinkazan.

Inter-annual variation of reproductive parameters and fruit availability in two populations of japanese macaques

We investigated the seed dispersal of Aglaia spectabilis, a large-seeded tree species in a moist evergreen forest of Khao Yai National Park in Thailand. Although one-to-one relationships between frugivores and plants are very unlikely, large-seeded plants having to rely on few large frugivores and therefore on limited disperser assemblages, might be vulnerable to extinction. We assessed both the frugivore assemblages foraging on arillate seeds of Aglaia spectabilis and dispersing them and the seed predator assemblages, thereby covering dispersal as well as the post-dispersal aspects such as seed predation. Our results showed that frugivores dispersing seeds were a rather limited set of four hornbill and one pigeon species, whereas two squirrel species were not dispersers, but dropped the seeds on the ground. Three mammal species were identified as seed predators on the forest floor. Heavy seed predation by mammals together with high seed removal rates, short visiting times and regurgitation of intact seeds by mainly hornbills lead us to the conclusion that hornbills show high effectiveness in dispersal of this tree species.

Dispersal of Aglaia spectabilis, a large-seeded tree species in a moist evergreen forest in Thailand

Seed dispersal by Japanese monkeys (Macaca fuscata yakui) via cheek-pouch was studied in a warm temperate evergreen forest on Yakushima Island. Plant list was compiled based on a study during 1986–1995, of which troops of monkeys have been habituated without artificial feeding. We followed the well-habituated monkeys in 1993 and 1994 to observe the feeding behavior and their treatments of fruits and seeds, and collected seeds dispersed by monkeys to record the distance carried from the mother trees. We checked the difference of germination ratio between seeds dispersed via cheek-pouch and seeds taken from mother trees by sowing experiments. Seeds and acorns of 22 species were observed to be dispersed via cheek-pouch of monkeys. Among them, three species with acorns were never dispersed via feces, and 15 species with drupes were seldom dispersed via feces. Plant species of which seeds are dispersed only via cheek-pouch had larger seeds than those of dispersed both via cheek-pouch and via feces, and typically had only one or two seeds in a fruit. As for one of cheek-pouch dispersal species,Persea thunbergii, the mean distance when seeds were carried from the mother trees via cheek-pouch was 19.7 m, and the maximum distance was as long as 105 m although more than 80% of seeds were dispersed within 30 m from mother trees. And 82% of seeds dispersed via cheek-pouch germinated. The easy separation of seeds from other parts of the fruit seems to facilitate cheek-pouch dispersal more than dispersal via feces. Cheek-pouch dispersal by monkeys has possibly enhanced the natural selection for larger seeds which bring forth larger seedlings with high shade-tolerance. In conclusion, cheek-pouch dispersal by monkeys is quite an important mode for trees in the mature stand in a warm temperate evergreen forest on Yakushima Island.

Naohiko Noma

Papers

Interactions between fleshy fruits and frugivores in a tropical seasonal forest in Thailand

Altitudinal and seasonal variations in the diet of Japanese macaques in Yakushima.

Inter-annual variation of reproductive parameters and fruit availability in two populations of japanese macaques

Dispersal of Aglaia spectabilis, a large-seeded tree species in a moist evergreen forest in Thailand

Cheek-pouch dispersal of seeds by Japanese monkeys (Macaca fuscata yakui) on Yakushima Island, Japan