Invasion of maritime chaparral by the introduced succulentCarpobrotus edulis

Invasion by the alien succulent,Carpobrotus edulis, has become a common occurrence after fire in maritime chaparral in coastal California, USA. We studied post-burnCarpobrotus establishment in chaparral that lackedCarpobrotus plants before the fire and compared seedbank and field populations in adjacent burned and unburned stands.Carpobrotus seeds were abundant in deer scat and in the soil before burning. Burning did not enhance germination: many seeds were apparently killed by fire and seed bank cores taken after fire revealed no germinable seeds. Laboratory tests showed that temperatures over 105°C for five minutes killedCarpobrotus seeds. In a field experiment involving use of herbivore exclosures, we found that herbivory was an important source of mortality for seedlings in both burned and unburned chaparral. All seedlings, however, died outside of the burn regardless of the presence of cages. Establishment there is apparently limited by factors affecting plant physiology. In the burned area, seedlings that escaped herbivory grew very rapidly. Overall, it appears that herbivory limited seedling establishment in both burned and unburned sites but that the post-burn soil environment supportedCarpobrotus growth in excess of herbivore use, thus promoting establishment.     

Fruit fate,frugivory, and fruit characteristics: a study of the hawthorn,Crataegus monogyna (Rosaceae)

Summary The fate of fruits from a population of European hawthorn (Crataegus monogyna) in western Oregon, USA, was examined over a two-year period. Only one frugivore, the American robin (Turdus migratorius) foraged on the C. monogyna fruits, making this an unusually straightforward fruit-frugivore system. Dispersal efficiency was low, with an average 21% of seeds being dispersed (carried away from parent plants) each year; the most common fate of fruits was to simply fall. Robins dropped 20% of the fruits that they picked, and defecated/regurgitated 40% of the fruits (seeds) that they swallowed, beneath parent plants. One trait, gruit abundance, strongly affected the probability of bush visitation by robins, bushes with larger fruit displays being preferred. Both absolute dispersal success (number of seeds) and dispersal efficiency (proportion of seeds; success per propagule) were also found to be correlated most strongly with initial fruit abundance. Individual plant fecundity and fruit quality were found to vary little between years; as a result, dispersal efficiencies for individual plants were also annully consistent. Larger (older) plants produced more fruits and therefore had higher fitness. These results suggest that the optimal fruiting strategy for C. monogyna is therefore to get as big as possible as quickly as possible by delaying fruiting until later in life.     

Seed dispersal in Erythronium grandiflorum (Liliaceae)

Primary and secondary seed dispersal was investigated for the glacier lily Erythronium grandiflorum in the Colorado Rocky Mountains. These heavy seeds have no obvious adaptations for biotic or abiotic dispersal, but can be thrown short distances when the dehiscent fruits are shaken by wind. We used sticky traps to measure primary transport of seeds up to 1 m away from individual plants. A seed cafeteria experiment examined the role of ants and rodents in secondary seed transport. Primary dispersal by wind was positively skewed and median transport distances were influenced by variation in plant height. Secondary dispersal was negligible compared to Viola nuttallii, an elaiosome-bearing species. Thus, seed dispersal was highly restricted in E. grandiflorum, and a 1 m radius encompassed the modal section of the seed dispersal curve. The seed dispersal component of gene flow was quantified and combined with previous measurements of pollen flow to yield a more complete estimate of Wright's neighborhood size, N _e, for E. grandiflorum. The lack of a special seed dispersal mechanism in E. grandiflorum is discussed in terms of a source-sink model for seedling establishment with respect to distance from the parental plants.     

Early postfire seed dispersal, seedling establishment and seedling mortality of Pinus coulteri (D. Don) in central coastal California, USA

Seedling recruitment in many highly serotinous populations of Pinus coulteri on California's central coast depends almost entirely on periodic, stand-replacing fire. Compared to serotinous pines of the Mediterranean Basin, little detailed information is available on the postfire demography of California closed-cone pines, including P. coulteri. In September 1996 a wildfire burned the 760-ha American Canyon Research Natural Area (RNA). Using aerial photography, we mapped burn severity of P. coulteri-chaparral woodlands and forests within the RNA. From May to September of 1997, we also quantified seedling establishment and mortality in relation to biophysical site characteristics including fire severity. Seventy-six percent of P. coulteri forests and woodlands experienced high-severity burns, 9% moderate-severity burns, and 15% low-severity or unburned. Of the 53 plots used for seedling counts, 70% were high-severity, 26% moderate-severity, and 4% low-severity. Seedling densities 13 months postfire were low (0.21 m^–2), but seedling mortality also was low (8.4%). Aerial seed bank size increased from north-facing to south-facing slopes and from high-severity to low-severity burns. Seedling recruitment was unrelated to burn severity and increased with the size of the canopy seed bank (cone density). Many seedlings established from rodent seed caches; 23% of the seedlings established in clumps from seeds cached by Dipodomys agilis, Chaetodipus californicus and Peromyscus maniculatus. Pinus coulteri seeds have low potential for dispersal by wind, but secondary dispersal by rodents moves seeds away from source trees and into neighboring chaparral. We discuss the potential importance of rodent seed caching to postfire demography of California and Mediterranean serotinous pines.     

Fruit removal and postdispersal survivorship in the tropical dry forest shrub Erythroxylum havanense: ecological and evolutionary implications

We studied the relationship between the removal rate and the spatiotemporal availability of ripe fruits of the tropical deciduous shrub Erythroxylum havanense in western Mexico. We also evaluated the effects of dispersal on seed survival during the first stages of establishment. Fast and early dispersal should be favored in E. havanense, since propagules have more time to grow and accumulate resources before the beginning of the severe dry season. In general, high rates of fruit removal imply faster and earlier dispersal. Thus, plants producing large crops should benefit from high removal rates, which will increase the probability of successful establishment by their progeny. To characterize both individual and population fruiting patterns, we made daily counts of fruits on 51 plants arranged in six clumps of different sizes. The daily number of fruits removed per plant was higher for plants with larger initial crop sizes and larger numbers of ripe fruits on a given day, but decreased as clump size increased. Additionally, we monitored postdispersal survival and germination in an experiment manipulating seed density, distance from adult plants, and seed predation. Early establishment was independent of density or distance, and vertebrate seed predation was the main agent of seed mortality. Our results indicate that the critical variable with respect to fruit removal is the number of fruits a plant produces, large plants having higher dispersal rates. Large plants are also more likely to have more seeds escaping postdispersal seed predation.     

Fruit removal efficiency and success: influence of crop size in a neotropical treelet

The reproductive success of animal-dispersed plants is closely linked to the number of seeds that they are able to disperse. The fruit crop size hypothesis states that a plant with large fruit crop size will attract more dispersers than a plant with a smaller fruit crop, which may result in more seeds being dispersed from the foremost. In this study, we experimentally examined the effect of crop size and other factors on primary seed dispersal in a neotropical shrub/tree, Casearia corymbosa (Flacourtiaceae). We used two predictive variables of reproductive success, which produce an accurate picture of seed dispersal ratio: fruit removal efficiency (proportion of a fruit crop removed by frugivores) and fruit removal success (relative contribution of each individual tree to the number of fruits removed in the population). We established two levels of fruit crop size at the C. corymbosa individuals, using plants with large (150 fruits) and small crops (50 fruits). We found that individual plants with larger crops had significantly higher values of fruit removal efficiency (92.6%) and success (5%) than plants with smaller crops (69.3% and 1.3%, respectively). Fruit removal efficiency was related to vegetation type, plant height and fruit width, but the variance explained by these variables was low ( < 8%). Fruit removal success was significantly related to crop size ( > 90% of the variance explained). These results suggest that fruit removal efficiency and success are strongly related to fruit crop size of C. corymbosa plants.     

Comparative water relations of adjacent california shrub and grassland communities

Summary Much of the coastal mountains and foothills of central and southern California are covered by a mosaic of grassland, coastal sage scrub, and evergreen sclerophyllous shrubs (chaparral). In many cases, the borders between adjacent plant communities are stable. The cause of this stability is unknown. The purpose of our study was to examine the water use patterns of representative grasses, herbs, and shrubs across a grassland/chaparrel ecotone and determine the extent to which patterns of water use contribute to ecotone stability. In addition, we examined the effects of seed dispersal and animal herbivory. We found during spring months, when water was not limited, grassland species had a much higher leaf conductance to water vapor diffusion than chaparral plants. As the summer drought progressed, grassland species depleted available soil moisture first, bare zone plants second, and chaparral third, with one chaparral species (Quercus durata) showing no evidence of water stress. Soil moisture depletion patterns with depth and time corresponded to plant water status and root depth. Rabbit herbivory was highest in the chaparral and bare zone as indicated by high densities of rabbit pellets. Dispersal of grassland seeds into the chaparral and bare zone was low. Our results support the hypothesis that grassland species deplete soil moisture in the upper soil horizon early in the drought, preventing the establishment of chaparral seedlings or bare zone herbs. Also, grassland plants are prevented from invading the chaparral because of low seed dispersability and high animal herbivory in these regions.     

Reproduction and growth of the chaparral geophyte, Zigadenus fremontii (Liliaceae), in relation to fire

Zigadenusfremontii is often a striking component of the flora following firein the chaparral. Like other geophytes, it produces large numbers of flowers inthe first spring after a burn. Although these plants are most conspicuous intheearly postfire environment, the question that remains is, how do they persistinthe interval between fires? To address this we investigated differences in thegrowth and reproduction of Z. fremontii in burned andunburned chaparral. We monitored marked individuals for nine years at threesites: two that were burned in 1990 and one in the same area that was inunburned mature chaparral. We measured leaf area, and production of flowers andfruits. We also conducted seed experiments in the field to determine the ratesand timing of germination. We found that reproduction occurs only in theimmediate postfireperiod: flowering and production of fruits and seeds in the first yearfollowingfire, and seedling establishment by year 3. There was a cost of reproduction;plants that flowered (in the burn area) had negative growth rates the followingyear. In contrast, plants in unburned chaparral, which did not flower, hadpositive growth rates over the same period. Moreover, plants that produced themost flowers had the lowest growth rates. In the unburned chaparral site,plantswere not dormant as predicted from previous literature; instead they producedleaves nearly every year. In most years the average leaf area per plant wasgreater than that in the burned sites. Our results indicate that postfirereproduction depends on growth and carbohydrate storage in the inter-fireperiod. We also suggest that this species is relatively long-lived for aherbaceous perennial.     

REPRODUCTIVE ATTRITION IN THE RARE CHAPARRAL SHRUB FREMONTODENDRON DECUMBENS LLOYD (STERCULIACEAE)

We examined reproductive attrition in Fremontodendron decumbens to characterize sexual reproduction in this rare California shrub. Reproductive individuals produced an average of 2,900 flower buds in a season, with no significant difference in bud production between two seasons. Because of intense insect predation, <;2% of initiated flower buds became mature fruits. A threefold decrease in predation of flower buds between seasons resulted in an increase in seed output the second season, indicating that seed production was partially predator-limited. Most seeds (97.8%) were dormant due to an impermeable seed coat. Breaking of the coat, mechanically or by heat, allowed high levels of germination. Chamise charate and ash added to the potting medium resulted in the highest level of germination and emergence. Rodents were more important than birds as seed predators, destroying 90% of seeds under parent shrub canopies within 8–10 months. Seeds already integrated into the seed bank were comparatively safe from predation, relative to newly added seeds. If predation was prevented, seeds were long-lived under field conditions (>;80% survived after 5.75 years). Most seedlings produced in unburned chaparral by planting heat-treated seeds in openings between shrubs were destroyed by predators (rodents and insects). All seedlings that escaped predation died during the summer drought. We concluded that sexual reproduction was limited by (in order of importance): 1) lack of fire, 2) predehiscence predation by insects, and 3) postdehiscence predation by rodents. Size distributions from two populations revealed that, despite the apparent absence of sexual reproduction in unbumed chaparral, two unbumed sites contained a large proportion of individuals in small size classes. Excavation of several small individuals demonstrated they were sprouts from the roots of nearby larger shrubs. Because asexual reproduction by rootsprouting circumvents the high attrition of sexual reproductive effort on unbumed sites, rootsprouting may be a significant reproductive strategy of some ‘sprouter’ species in chaparral.     

Variation in bradyspory and seedling recruitment without fire among populations of Banksia serrata (Proteaceae)

Abstract Long-term canopy storage of seeds in fruits (bradyspory) varies among species and is thought to be related to the fire regime experienced by the plant community. This study sought to quantify intraspecific variation in bradyspory among nine populations of Banksia serrata (Proteaceae) on the south coast of New South Wales. Marked differences were found between populations, ranging from more than 30% of follicles open at two coastal sites, to less than 5% of follicles open at two inland heath sites. Low levels of bradyspory occurred particularly in coastal sites which also had Banksia integrifolia, a non-bradysporous species, present. In addition to spontaneous follicle opening without fire, fall of infructes-cences to the ground after breaking of branches was observed to be a major contributor to seed release. At three sites where fire had not occurred for many years, the amount of seedling establishment since the last fire corresponded well with levels of bradyspory; there were no inter-fire recruits at the site with the strongest bradyspory, while at the site with the weakest bradyspory there were numerous inter-fire plants, some with lignotubers large enough to be able to tolerate the next fire. It is argued that populations of B. serrata exhibit variable patterns of recruitment, depending on the site characteristics, ranging from complete dependence on fire for population maintenance to bet-hedging (spreading potential recruitment between fire events and inter-fire periods). Variations in populations and reproductive characteristics among sites make the application of demographic models to management decisions habitat-specific.     

Allometric allocation in fruit and seed packaging conditions the conflict among selective pressures on seed size

Selective pressures on seed size could vary among the different stages of plant life cycles, so no simple relation could explain a priori its evolution. Here, we determined the relationships between seed size and two fitness components—seed dispersal and survival from predation—in a bird-dispersed tree, Crataegus monogyna. We interpret these relationships in relation to the patterns of mass allocation to fruit and seed components. Selection patterns were assessed at two levels (1) selection pressures on the parent tree; comparing seed dispersal efficiency among individual plants and (2) selection pressures at the individual seed level; comparing seed size variation (i) before and after dispersal, and (ii) before and after postdispersal seed predation. Dispersal efficiency (percentage of seed crop dispersed) was positively correlated with fruit mass and fruit width. Differences in crop size did not offset this effect, and larger seeds were overrepresented in the seed rain relative to the seed pool before dispersal. However, the advantage of larger seeds during the dispersal stage was cancelled later by an opposite selection pressure exerted by seed predators. As a result, smaller seeds had a higher probability of surviving postdispersal seed predation, establishing an evolutionary conflict imposed by the need for dispersal and the danger of being predated. Birds and rodents preferentially selected highly profitable fruits and seeds in terms of the relative proportion of their components. Larger fruits had a higher pulp to seed proportion than smaller ones, and all seeds had the same proportion of coat relative to the embryo-plus-endosperm fraction. Hence, although predator pressures were stronger than disperser ones, larger seeds invested proportionally less in structural defense than in dispersal.     

Indirect interactions between browsers and seed predators affect the seed bank dynamics of a chaparral shrub

Interactions between herbivores and seed predators may have long-term consequences for plant populations that rely on persistent seed banks for recovery after unpredictable fires. We assessed the effects of browsing by deer and seed predation by rodents, ants and birds on the densities of seeds entering the seed bank of Ceanothus cuneatus var. rigidus, a maritime chaparral shrub in coastal California. Ceanothus produced many more seeds when protected from browsers in long-term experimental exclosures than did browsed plants, but the seed densities in the soil beneath browsed and unbrowsed Ceanothus were the same at the start of an intensive one-year study. The density of seeds in the soil initially increased in both treatments following summer seed drop: while densities returned to pre-drop levels within a few weeks under browsed plants, soil seed densities remained high for 5–8 months beneath unbrowsed plants. Rodent abundance (especially deer mice) was higher near unbrowsed plants than >30 m away, and rodents removed Ceanothus seeds from dishes in the experimental plots. At least in the short term, rodent density and rates of seed removal were inversely related to the intensity of browsing. Our data have management implications for maintaining viable Ceanothus populations by regulating the intensity of browsing and the timing, intensity and frequency of fires.Electronic Supplementary Material Supplementary material is available to authorised users in the online version of this article at .     

Long-term variability in seed size and seedling establishment of <Emphasis Type="Italic">Maianthemum bifolium</Emphasis>

Relatively few studies conducted in natural plants populations focus on the relationship between seed size and their germination ability and seedling establishment. Maianthemum bifolium is a perennial herb that spreads vegetatively through rhizomatous growth and reproduces through seeds. However, this species is characterized as seed and microsite limited, and under undisturbed conditions seedlings are not noted. The studies were conducted in two populations of M. bifolium in six subsequent seasons. The mean seed mass was negatively correlated both per ramet as well as in the fruit with the number of seeds, and positively with its height and the number of flowers. The long-term mean annual production of seeds in the populations was 37 and 56 seeds per m². The seeds from both populations had similarly high germination abilities that were approximately 90% under laboratory conditions, 60% in garden, and 55% in the natural habitat. Seeds from four size classes were sown and a positive correlation was noted between seedling establishment and the mass of the seeds from which they grew (r_S = 0.64). Seedling survival was also significantly correlated with seed mass.     

Seed dispersal of East Asian coastal dune plants via seawater – short and long distance dispersal

Coastal plants provide precious and irreplaceable services to human and coastal ecosystems, but people are still rather unaware how coastal plants relate with sea tides. We assumed that their seeds may reflect some relationships with tides. Our objective was to understand seed dispersal of plants living in the upper coastal regions, not reached by normal tides, but inundated by storm surges. For this purpose, we observed seed characteristics, especially seed buoyancy, of coastal and non-inundated plants of the Shandong Peninsula, north China. Through field simulation, we studied how buoyancy affected the dispersal during tides, using twig segments as retrievable dummies of seeds, in order to collect evidences with concern, how seed dispersal could occur during storm surges, which are extremes of tides. Coastal plants had predominantly buoyant seeds, while inland-growing plants had largely non-buoyant seeds. Ninety percent of buoyant twig segments (mimicking seeds) were recaptured after tides on experimentally used coastal areas; only 1% of non-buoyant ones of such dummies were recaptured, whereas the rest probably had been swept away by wave action to deposit at the sea bottom. Buoyant twig segments (mimicking seeds) dispersed along the coastlines as tides surged towards the coast. About 10% of these buoyant twigs were swept away with off-shore currents, but they might land elsewhere, similar as it might happen with long-distance dispersal of seeds.     

Ecological benefits of myrmecochory for the endangered chaparral shrub Fremontodendron decumbens (Sterculiaceae)

Fremontodendron decumbens grows in a single county in central California, USA. Prior research showed that its elaiosome-bearing seeds are dispersed by the harvester ant Messor andrei. I tested several hypotheses regarding the positive role of ant-mediated dispersal to F. decumbens: (1) Does ant-mediated seed dispersal facilitate seed escape from rodent predation?; (2) Does ant processing of seeds stimulate germination?; (3) Are ant middens more suitable microsites for seed or seedling survival in unburned chaparral areas?; and (4) Do survival benefits of dispersal occur post-fire in the form of differences in seedling survival probabilities and, if so, why? Results of tests of each hypothesis were: (1) similar percentages of seeds placed on ant middens and under F. decumbens shrub canopies were destroyed by rodents, but seeds from which elaiosomes had been removed were more likely to escape rodent predation; (2) seeds processed by ants did not germinate more readily than seeds removed directly from shrub branches; (3) seedling predation was a major cause of mortality in unburned chaparral on both ant middens and under shrubs, and overall seedling survival did not differ between the two microsites; (4) post-burn seedling survival was significantly greater for seedlings dispersed away from F. decumbens shrub canopies, because dispersed seedlings were both less likely to be killed by predators and more likely to be growing in a gap created by the fire-caused death of an established shrub. I concluded that the major ecological benefit to F. decumbens of ant-mediated seed dispersal was elevated post-fire seedling survival resulting from enhanced escape by dispersed seedlings from both predation and competition.     

Birds and guelder rose <Emphasis Type="Italic">Viburnum opulus</Emphasis>: selective consumption and dispersal via regurgitation of small-sized fruits and seeds

The evolution of fleshy fruit size, in particular in bird-dispersed plants, is believed to be influenced by the size of seed-dispersing vertebrates through gape limitation. Also, it has been demonstrated that seed size correlates positively with fruit size, especially in single- or few-seeded fruits. However, there is little evidence of current selection pressure by disperser birds on fruit and seed size within populations of a particular plant species. In the present study, this aspect was investigated in guelder rose Viburnum opulus (Caprifoliaceae) fruit consumption by birds in an area in NW Spain. Guelder rose fruits are sub-globose drupes that can exceed 11 mm in width, with a single hard seed of up to 8.5 mm in width. Most of the seeds were dispersed by the robin Erithacus rubecula (gape width < 8 mm) and a small thrush, the song thrush Turdus philomelos (gape width < 11 mm), which swallowed the fruits whole, and some were destroyed by the bullfinch Pyrrhula pyrrhula. Most of the seeds were regurgitated rather than defecated by disperser birds, probably because seed size limited gut processing. The mean size of the ingested seeds was smaller than the mean diameter of Turdus droppings, which in turn was smaller than the mean size of the seeds of the fruits available on the plants. As winter progressed, only larger fruits and seeds remained on the plants (seed and fruit size were positively correlated), and the size of ingested seeds increased. Thus, the largest fruits were consumed less by seed-dispersing birds and were exposed to seed-predators (bullfinches and climbing rodents) for longer. Selection pressure on smaller guelder rose seeds must therefore be effective in the dispersal stage in the study area.     

The role of Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island, the Ryukyu Archipelago, Japan

The role of the Orii’s flying-fox (Pteropus dasymallus inopinatus) as a pollinator and a seed disperser on Okinawa-jima Island was investigated by direct observations and radio-tracking from October 2001 until January 2006. We found that Orii’s flying-fox potentially pollinated seven native plant species. Its feeding behavior and plant morphological traits suggested that this species is an important pollinator of Schima wallichii liukiuensis and Mucuna macrocarpa. The flying-fox also dispersed the seeds of 20 native plant species. The seeds of all plants eaten by the flying-fox were usually dropped beneath the parent tree, although large fruits of four plant species were occasionally brought to the feeding roosts in the mouth, with the maximum dispersal distance—for Terminalia catappa—estimated to be 126 m. Small seeds of 11 species (mostly Ficus species) were dispersed around other trees, during the subsequent feeding session, through the digestive tracts, with the mean dispersal distance for ingested seeds estimated at 150 ± 230.3 m (±SD); the maximum dispersal distance was 1833 m. A comparison of the seed dispersal of available fruits according to the size of flying-foxes and other frugivores suggested that the seed dispersal of eight plant species producing large fruits mostly depended on Orii’s flying-fox. On Okinawa-jima Island, the Orii’s flying-fox plays an important role as a pollinator of two native plants and as a long-distance seed disperser of Ficus species, and it functions as a limited agent of seed dispersal for plants producing large fruits on Okinawa-jima Island.     

Diversities in morphological characteristics and seed germination behavior in fruits ofSalsola komarovii Iljin

Two different types of dispersal units (called fruits in this study) were observed inSalsola komarovii Iljin. One is a fruit which has dark brown lignified tepals with long wings and a green seed and falls easily from the mother plant (long-winged type). Another has light brown lignified tepals with short wings and a yellow seed and attaches tightly to the mother plant (short-winged type). This difference of fruit type appeared independently from maturity of fruit. Seeds in the short-winged fruits were in dormancy for a longer period of time than those in the long-winged fruits. The germination rate was significantly higher in the seeds of long-winged fruits. The dormancy in seeds of the long-winged fruits was effectively terminated by reducing the temperature but the effect of chilling was very weak in seeds of the short-winged fruits. It was concluded from these observations that there exists a dimorphism in the fruits of, or a heterocarpy in,S. komarovii. The plants grown under water stress produced mostly short-winged fruits and those grown under well-watered conditions bore fruits of both types. Exogenously applied abscisic acid (ABA) tended to produce the short-winged fruits, suggesting that the heterocarpy was, at least partly, regulated by ABA.     

Two-phase seed dispersal: linking the effects of frugivorous birds and seed-caching rodents

Frugivorous birds disperse the seeds of many fruit-bearing plants, but the fate of seeds after defecation or regurgitation is often unknown. Some rodents gather and scatter hoard seeds, and some of these may be overlooked, germinate, and establish plants. We show that these two disparate modes of seed dispersal are linked in some plants. Rodents removed large (>25 mg) seeds from simulated bird feces (pseudofeces) at rates of 8–50%/day and scatter hoarded them in soil. Ants (Formica sibylla) also harvested some seeds and carried them to their nests. Rodents carried seeds 2.5±3.2 m to cache sites (maximum 12 m) and buried seeds at 8±7 mm depth. Enclosure studies suggest that yellow pine chipmunks (Tamias amoenus) and deer mice (Peromyscus maniculatus) made the caches. In spring, some seeds germinated from rodent caches and established seedlings, but no seedlings established directly from pseudofeces. This form of two-phase seed dispersal is important because each phase offers different benefits to plants. Frugivory by birds permits relatively long-range dispersal and potential colonization of new sites, whereas rodent caching moves seeds from exposed, low-quality sites (bird feces on the ground surface) to a soil environment that may help maintain seed viability and promote successful seedling establishment.     

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.