Naohiko Noma

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

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.

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

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.

Cheek-pouch Dispersal of Seeds by Japanese Monkeys (Macaca fuscata yakui) on Yakushima Island, Japan

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.

Environmental determinants of the altitudinal variations in relative group densities of Japanese macaques on Yakushima

Altitudinal variations in relative group densities of the Japanese macaques on Yakushima were studied. This is an ideal place for studying resource limitations because it avoids various complicating factors that are difficult to quantify but might affect animal densities, such as predation, interspecific competition, and past catastrophes. The relative group density was high in the coastal forest (0–400 m), while it did not differ among the higher zones (400–800, 800–1200 and 1200–1886 m). To examine this variation, three habitat variables were analyzed: total basal area of food trees per unit area, seasonal variations in fruit abundance, and total annual fleshy fruit production. All of these variables indicate that fruit and seeds are most available in the coastal forest. Thus, altitudinal variations in the density of Japanese macaques on Yakushima are determined by the total annual food abundance.

Dispersal of Canarium euphyllum (Burseraceae), a large-seeded tree species, in a moist evergreen forest in Thailand

We investigated the dispersal of a large-seeded tree species, Canarium euphyllum (Burseraceae), in the moist evergreen forests of the Khao Yai National Park in Thailand. By combining direct observations of fruit consumption in tree canopies (543 h) and the camera-trapping observations of fallen fruit consumption on the forest floor (175 camera-days), we identified the frugivore assemblage that foraged on the fruits of C. euphyllum and assessed their role in seed dispersal and seed predation. In the canopy, our results showed that seeds were dispersed by a limited set of frugivores, one pigeon and four hornbill species, and predated by two species of squirrel. On the forest floor, seven mammal species consumed fallen fruits. A combination of high rates of fruit removal and short visiting times of mountain imperial pigeons (Ducula badia) and hornbills (Buceros bicornis, Aceros undulatus, Anorrhinus austeni and Anthracoceros albirostris) led us to conclude that these large frugivorous birds provide effective seed dispersal for this tree species, in terms of quantity. These frugivorous species often have low tolerance to negative human impacts and loss of these dispersers would have severe deleterious consequences for the successful regeneration of C. euphyllum.

Pattern and impact of hornbill seed dispersal at nest trees in a moist evergreen forest in Thailand

Following the entire process of frugivore seed dispersal, from intake of seeds to seed deposition, is a difficult task. One alternative is to monitor areas of heavy seed rain deposited by animals. We quantified the number of seeds deposited by hornbills and followed the fates of these seedlings for 3 y to evaluate the effectiveness of hornbill seed dispersal at nest trees, on the basis of seedling survival. For 14 mo, fallen fruits and seeds were collected in traps established around four nest trees of each of two hornbill species ( Aceros undulatus and Anthracoceros albirostris ) and the seedlings were monitored in adjacent quadrats. Seedfall and seedlings of species represented in hornbill diets occurred at significantly higher densities in the traps/quadrats in front of nest cavities than in other traps/quadrats. Fewer seedling species and individuals germinated under nest trees than expected from the composition of the seedfall. Our results suggest that the quality of hornbill seed dispersal might be poor at nest trees due to the highly concentrated seedfall, which results in high seed and seedling mortality. Although seed deposition at nest trees is a useful guide to hornbill diet during the breeding season, it is clearly not of benefit to the plants involved. However, the pattern and consequences of hornbill seed dispersal at nest sites is likely very different from that during the non-breeding season.

Relationship between leaf characteristics, tree sizes and species distribution along a slope in a warm temperate forest

The relationships between some leaf characteristics, tree size and species distribution were investigated for evergreen tree species along a slope in a warm temperate forest in Japan. Tree species were classified into three groups based on their dominance on the slope: ‘ridge species’ that were aggregated in an uppersite, ‘valley species’ that were aggregated in a lowersite, and ‘uniform species’ that were distributed almost uniformly. The ridge species had a more positive leaf carbon isotope ratio than the valley species, which suggests that the ridge species have larger water use efficiency than the valley species. This may give some advantage to the ridge species over the valley species in the uppersite where water availability would be limited. However, the ridge species had smaller leaf nitrogen content on a mass basis and larger leaf mass per area than the valley species, which suggests that the ridge species had a smaller mass-based leaf photosynthetic capacity than the valley species. This may be disadvantageous to ridge species in the lowersite, because smaller leaf photosynthetic capacities cause lower leaf carbon gain and thus lower growth than the valley species. These differences in leaf characteristics between the ridge and the valley species were affected by microenvironments, and were also partly affected by the difference in species specific responses to microenvironments on the slope.

Mass mortality of Japanese macaques in a western coastal forest of Yakushima

The mass mortality of wild Japanese macaques (Macaca fuscata Blyth) was observed in a warm temperate forest of Yakushima, southern Japan. Demographic changes of eight troops between August 1998 and August 1999 were studied and 56% of macaques disappeared from the five intensively studied troops. Mortality varied among troops: two troops became extinct, while another troop did not decrease in size. The rate of mortality of the other troops was between 33 and 80%. The variation in mortality among the troops was either the outcome of local concentrations of mortality or of intertroop competition. The rate of mortality decreased with increasing distance from the two extinct troops and with increasing troop size; these two factors could not be separated statistically. The direct cause of death was diagnosed as pneumonia for four out of five fresh carcasses. The fleshy fruit production in autumn 1998 was the lowest in 14 years, and macaques had relied on leaves earlier than in usual years. It was exceptionally hot and dry in the summer of 1998. The exceptionally poor fruit production and hot summer of this year, with the resulting shortage of high-quality foods, was consistent with the scenario that mass mortality was due to the poor nutritional conditions. However, the possibility that epidemics caused the mass mortality cannot be ruled out. Our findings proved that primates in a seemingly stable habitat experience fluctuations in demographic parameters under natural conditions.

Flowering phenology and mating success of the heterodichogamous tree Machilus thunbergii Sieb. et Zucc (Lauraceae)

Heterodichogamy is defined as the presence of two flower morphs that exhibit male and female functions at different times among individuals within a population, and is regarded as an adaptation to promote outbreeding through enhanced intermorph pollination. In highly fragmented populations in which the morph frequency is biased, heterodichogamy may hamper population growth by reducing seed sets of the more numerous morph, and enhancing seed sets of the less numerous morph. In such situations, we hypothesize that individual plants experience greater seed sets if the opposite sexual morphs are nearby, and that individuals of a less numerous sexual morph have greater seed sets. After confirming heterodichogamy by observing flowering behavior and phenology, we tested these two hypotheses in a highly fragmented population of Machilus thunbergii, a putative heterodichogamous evergreen laurel tree. Our observations confirmed that M. thunbergii is heterodichogamous, consisting of two types of protogynous and bisexual flowers: a morning female (MF)–afternoon male morph and a morning male (MM)–afternoon female morph at the individual level. Sexual expression of the two morphs was highly synchronized and reciprocal. Investigation of seed-set rates revealed greater rates of both morphs if the opposite morph was nearby. The less numerous sexual morph (MF) showed a greater seed-set rate than the more numerous sexual morph (MM).

Development of microsatellites in Machilus thunbergii (Lauraceae), a warm-temperate coastal tree species in Japan

PREMISE OF THE STUDY Microsatellite markers were developed and characterized in a typically coastal, widespread, and dominant tree species of the evergreen broadleaf forests, Machilus thunbergii, for comparison of the genetic diversity and structure of inland populations surrounding the ancient Lake Biwa and coastal populations in Japan. METHODS AND RESULTS Eighteen polymorphic microsatellites of this species were isolated using an improved technique for isolating codominant compound microsatellite markers. These isolated loci provided compound simple sequence repeat (SSR) markers with polymorphisms of three to 19 alleles per locus, with an average of 10.9. The expected and observed within-population heterozygosities ranged from 0.16 to 0.86 and from 0.13 to 0.72, respectively. CONCLUSIONS These markers may be useful tools for further investigation of the population genetic structure and biogeographic history of M. thunbergii in the warm-temperate zone of East Asia.