Experimental Seed Predator Removal Reveals Shifting Importance of Predation and Dispersal Limitation in Early Life History Stages of Tropical Forest Trees

Recruitment limitation of trees in tropical forests can occur because of high rates of seed predation or low rates of seed dispersal, but the degree to which limitation is influenced by variation in seed predator abundance, and hence variation in seed predation and dispersal, is not well understood. We experimentally reduced the density of a granivorous small mammal (Heteromys desmarestianus) by 90 % to assess the degree to which its rates of seed predation and dispersal limit seed to seedling survival of nine species of trees in a Neotropical lowland forest. Overall, the proportion of seeds that germinated was influenced more by high rates of predation than by limited dispersal. Reduction in density of H. desmarestianus resulted in an order of magnitude decrease in fruit removal rates and an order of magnitude increase in both the absolute and relative numbers of seeds that germinated. However, the proportion of seeds that were cached remained relatively constant across all periods and between control grids and removal plots. In removal plots, H. desmarestianus dispersed and cached about 10 % of the fruits they handled, of which approximately 25 % germinated. This was 2 to 3 times greater than the germination rates of undispersed seeds, and for two species dispersed seeds were the only ones that germinated. The results suggest the simultaneous occurrence of both seed predation and dispersal limitation for trees with animal-dispersed seeds, but there may also be a hierarchy of importance in the relative strength of these two mechanisms that is determined by the dynamics of seed predator populations.     

Seed Dispersal of a High Quality Fruit by Specialized Frugivores: High Quality Dispersal?1

Dispersal quality, as estimated by the cumulative effects of dispersal, germination, seed predation, and seedling survival, was examined for Beilschmiedia pendula (Lauraceae) in Monteverde, Costa Rica. I determined the pattern of dispersal by finding seeds deposited by birds, protected the seeds from seed predators with cages to assess germination and seedling survival, and examined seed predation rates with marked seeds. Seed predation, germination, and seedling survival were compared between seeds naturally dispersed by birds and seeds placed at randomly located sites. Approximately 70 percent of seeds dispersed by birds (N= 244) were deposited <10 m from crown edges of fruiting B. pendula trees, although some seeds were dispersed at least 70 m away. Larger seeds were more likely to be dispersed under or close to the parent trees, and larger seeds produced larger seedlings. Seed size was not correlated directly with seedling survival, but larger seedlings at three months were most likely to survive one year. Seed predation by mammals and insects and seedling mortality due to fungal pathogens were concentrated beneath the crowns of parent trees. Seedlings and saplings were more abundant beneath fruiting B. pendula trees, but individuals farther away were taller on average. Thus, dispersal is beneficial for B. pendula, but such benefits appear most pronounced at a small spatial scale; seeds dispersed >30 m from the crown edges actually had a lower probability of survival than those dispersed 10–20 m. Only 10 percent of B. pendula. seeds received high‐quality dispersal in terms of landing in the zone with the highest per seed probability of seedling survival 10–20 m from parental crowns.     

Small rodents as significant dispersers of tree seeds in a Neotropical forest

Abstract. Through seed dispersal and predation, terrestrial mammals should be an important component of the mechanisms that determine patterns of tree recruitment in tropical forests. Despite their great abundance and ubiquity in Neotropical forests, small rodents as seed predators and dispersers remain largely forgotten. To investigate the fates of seeds in a hunted primary forest in Belize, we tagged seeds of Astrocaryum mexicanum (Palmae), Ampelocera hottlei (Ulmaceae), and Pouteria sapota (Sapotaceae) and placed them into open plots, exclosures accessible only to small mammals, and exclosures accessible to medium-sized and small mammals. The exclosure experiments and fates of the seeds show that the spiny pocket mouse, Heteromys desmarestianus (Heteromyidae), was the dominant handler of seeds of the first two species and also removed a significant proportion of the very large-seeded Pouteria. Most of the seeds were killed immediately upon removal, but many of the seeds (3–18 %) of the first two species were scatterhoarded (dispersed and buried in the soil) by Heteromys. Some of the scatterhoarded seeds (29%) remain buried and therefore protected from predation by other animals. Agoutis (Dasyprocta punctata), a caviomorph rodent, buried 13 % of the seeds of Pouteria, and Heteromys consumed and dispersed but did not bury Pouteria seeds. Results of this study support predictions by some researchers that small rodents are dominant terrestrial granivores in Neotropical forests. The role of small rodents as seed dispersers, however, has never been fully appreciated.     

Combining distances of ballistic and myrmecochorous seed dispersal in Adriana quadripartita (Euphorbiaceae)

The separate contributions of different vectors to net seed dispersal curves of diplochorous systems have rarely been characterised. In Australia, myrmecochory is a common seed dispersal syndrome and in the majority of such systems, seeds are initially dispersed ballistically. We measured ballistic and myrmecochorous seed dispersal distances in relation to canopies of Adriana quadripartita (Euphorbiaceae) and used a simulation model to estimate the net dispersal curve. We also compared seed removal rates and ant abundances under, and outside, plant canopies to examine how foraging patterns by ants may affect net dispersal.Overall ant abundance did not show a significant numerical response to seedfall; however, the abundance of the main seed dispersing ant, Rhytidoponera ‘metallica’ did. Despite this, seed removal rates did not differ significantly between canopy and open locations. Rhytidoponera ‘metallica’ account for 93% of observed seed dispersal events. On average, the ants dispersed seeds 1.54 m and in doing so, moved seed a mean radial distance of 0.76 m away from canopy edges. This contribution to net dispersal distance by ants is considerable since ballistic dispersal moved seeds a median distance of 7.5 cm. Our simulation model indicated that the combination of ballistic and ant seed dispersal is expected to result in seeds being transported a median net radial dispersal distance of 1.05 m from the canopy edge.Thus in this system, an important function of diplochory may simply be to move a higher proportion of seeds from under the canopy of parent plants than is possible by ballistic dispersal alone. This ‘dispersal-for-distance’ may result in reduced parent–offspring competition or may increase the probability that seeds reach rare safe sites for germination and recruitment.     

Effects of disperser abundance, seed type, and interspecific seed availability on dispersal distance

Seed dispersal distance is influenced by a variety of seed properties and functional responses of dispersers. However, to our knowledge, how and why seed dispersal distances are determined remains poorly understood. In the present study, seeds of sympatric tree species, Pinus koraiensis, Corylus mandshurica, Corylus heterophylla, and Quercus mongolica were released to investigate the effects of rodent abundance, seed type, and seed availability on seed dispersal. Our results showed that seeds of P. koraiensis were dispersed further than those of C. heterophylla and C. mandshurica regardless of the ambient rodent and seed abundances, reflecting a consistent effect of seed type on seed dispersal distances. Seed dispersal distance was greatly facilitated by lower per-capita seed abundance (the ratio of seeds to rodents); however, seed caching and cache survival were benefited from higher per-capita seed abundance. Although seed dispersal and seed caching of a particular tree species can be enhanced by its own seed availability, no consistent influence was detected at interspecific levels, reflecting different interspecific effects of seed availability on seed dispersal of sympatric seed species. Our results provide evidences that the effect of seed availability on seed dispersal should be evaluated in terms of per-capita seed abundance and interspecific effects, rather than the independent influence of seed or disperser abundances.     

A new hypothesis for the importance of seed dispersal in time

Most studies on seed dispersal in time have focused on seed dormancy and the physiological triggers for germination. However, seed dispersed by animals with low metabolic and moving rates, and long gut-passage times such as terrestrial turtles, could be considered another type of dispersal in time. This study tests the hypothesis that seeds dispersed in time may lower predation rates. We predicted that seeds deposited below parent trees after fruiting fall has finished is advantageous to minimize seed predators and should show higher survival rates. Four Amazonian plant species, Dicranostyles ampla, Oenocarpus bataua, Guatteria atabapensis and Ocotea floribunda, were tested for seed survival probabilities in two periods: during fruiting and 10-21 days after fruiting. Experiments were carried out in two biological stations located in the Colombian Amazon (Caparú and Zafire Biological Stations). Seed predation was high and mainly caused by non-vertebrates. Out of the four plant species tested, only Guatteria atabapensis supported the time escape hypothesis. For this species, seed predation by vertebrates after the fruiting period increased (from 4.1% to 9.2%) while seed predation by non-vertebrates decreased (from 54.0% to 40.2%). In contrast, seed predation by vertebrates and by non-vertebrates after the fruiting period in D. ampla increased (from 7.9% to 22.8% and from 40.4% to 50.6%, respectively), suggesting predator satiation. Results suggest that for some species dispersal in time could be advantageous to avoid some type of seed predators. Escape in time could be an additional dimension in which seeds may reach adequate sites for recruitment. Thus, future studies should be address to better understand the survival advantages given by an endozoochory time-dispersal process.     

Experimental field test of spatial variation in rodent predation of nuts relative to distance and seed density

The spatial context in which seed predation occurs may modify the spatial structure of recruitment generated by seed dispersal. The Janzen–Connell (J-C) model predicts that granivores will exert greater pressure on the parent plant or at those sites where the density of dispersed seeds is higher. We have investigated how the probability of post-dispersal survival of Juglans australis varies with nut density across a hierarchy of spatial scales. We experimentally evaluated the survival of 3,120 nuts at three spatial scales: meso-scale (≤1.5 ha), as forest sites with two densities of fruiting J. australis individuals; intermediate scale (<0.2 ha), as individual trees with two experimental crop sizes; small scale (<0.1 m²), as microsites with two factors (number of nuts and distance from source). Nut removal coincided with seed predation, a condition that allowed us to test the density-dependent seed predation hypothesis. We found that the probability of nut survival was greater at forest sites with higher J. australis density. Nut survival was not affected by nut density in the seed shadow of individual specimens: at sites where J. australis density was greater, the proportion of surviving nuts did not differ between microsites located at different distances from the parent plant, but it was greater at microsites with greater initial nut density. Nut survival depended on the scale at which rodents responded to nut density, being negatively density dependent at the meso-scale and spatially random at intermediate and small scales. At the meso-scale, excess nut supply increased the probability of nut survival, which is in agreement with a model of granivore satiation near the seed source. Rodent satiation at the meso-scale may favour maintenance of sites with high J. australis density, where individual trees may have greater probabilities of passing their genes onto the next stage of the dispersal cycle.     

Scatter Hoarding of Seeds Confers Survival Advantages and Disadvantages to Large-Seeded Tropical Plants at Different Life Stages

Scatter hoarding of seeds by animals contributes significantly to forest-level processes, including plant recruitment and forest community composition. However, the potential positive and negative effects of caching on seed survival, germination success, and seedling survival have rarely been assessed through experimental studies. Here, I tested the hypothesis that seed burial mimicking caches made by scatter hoarding Central American agoutis (Dasyprocta punctate) enhances seed survival, germination, and growth by protecting seeds from seed predators and providing favorable microhabitats for germination. In a series of experiments, I used simulated agouti seed caches to assess how hoarding affects seed predation by ground-dwelling invertebrates and vertebrates for four plant species. I tracked germination and seedling growth of intact and beetle-infested seeds and, using exclosures, monitored the effects of mammals on seedling survival through time. All experiments were conducted over three years in a lowland wet forest in Costa Rica. The majority of hoarded palm seeds escaped predation by both invertebrates and vertebrates while exposed seeds suffered high levels of infestation and removal. Hoarding had no effect on infestation rates of D. panamensis, but burial negatively affected germination success by preventing endocarp dehiscence. Non-infested palm seeds had higher germination success and produced larger seedlings than infested seeds. Seedlings of A. alatum and I. deltoidea suffered high mortality by seed-eating mammals. Hoarding protected most seeds from predators and enhanced germination success (except for D. panamensis) and seedling growth, although mammals killed many seedlings of two plant species; all seedling deaths were due to seed removal from the plant base. Using experimental caches, this study shows that scatter hoarding is beneficial to most seeds and may positively affect plant propagation in tropical forests, although tradeoffs in seed survival do exist.     

Nut predation and dispersal of Harland Tanoak Lithocarpus harlandii by scatter-hoarding rodents

Plants that use the propagule to co-opt animals as dispersal agents must balance the costs of seed predation with the benefits of dispersal. Successful post-dispersal germination is a key metric that reflects these costs and benefits. By tracking individual nuts with coded tin-tags over 3 years (2000–2003), this study quantified nut predation and dispersal of harland tanoak (Lithocarpus harlandii) by seed-caching rodents in a subtropical evergreen broadleaved forest in the Duiangyan Region of Sichuan Province, Southwest China. We found that tanoak seedlings established from rodent-generated caches in the primary stands over a 12-month post-dispersal period. Our results indicate that seed-caching rodents are effective dispersers of tanoak nuts, but dispersal effectiveness varies among years and stands, probably due to mast seeding of harland tanoak or community-level seed availability according to the predator satiation hypothesis. Some nut traits in tanoak species, e.g. large seed size, hard nut husk, lower tannin and mast seeding, are important characteristics for seed dispersal by scatter-hoarding rodents, compared with oak species with higher tannin content.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

Do Meyer’s Parrots Poicephalus meyeri benefit pollination and seed dispersal of plants in the Okavango Delta,Botswana?

Recent studies of new world parrots repeatedly document, with few exceptions, that parrots are wasteful and destructive predispersal seed predators that are unlikely to contribute towards pollination and seed dispersal. Few detailed studies, however, have assessed the contribution of African parrots to forest ecology by quantifying the potential net benefit of seed and flower predation by parrots for most tree species in their diet. Due to the incidence of pollen on the heads of Meyer’s Parrots when feeding on Leguminosae flowers and the dispersal of viable seeds to the ground during seed predation, we compared destruction rates, when feeding on pods, fruits and flowers, with dispersal rates of viable seeds to the ground and frequency of head contact with reproductive apparatus to estimate net benefit from Meyer’s Parrot feeding activity. Meyer’s Parrots were not implicated in endo‐ or epizoochory, but they dropped uneaten fruit pulp and seeds to the ground during feeding bouts, thus providing ripe, undamaged seeds to secondary seed dispersers. This link with forest recruitment was weak, as all tree species utilized by Meyer’s Parrots either had more significant primary dispersal agents or were primarily wind‐dispersed. In most cases, the negative effect of seed predation outweighed any positive effects in terms of dispersal, whereby almost three times more seeds were consumed or destroyed than were dispersed to the ground. Significantly, only Sclerocarya birrea caffra recorded marginal net dispersal benefit from utilization by Meyer’s Parrots. Due to low relative resource abundance and high destruction rate, feeding activity on Berchemia discolor may be significant enough to influence its spatial distribution and abundance. Utilization of flowers of Kigelia africana and Adansonia digitata by parrots likely had a significant negative impact on pollination. Feeding on Acacia nigrescens flowers, however, was potentially advantageous to their pollination. We conclude that Poicephalus parrots are net consumers of ripe, undamaged seeds and flowers, thus having an overall negative impact on forest recruitment in subtropical Africa.     

啮齿类对植物种子的传播作用

种子植物是固着生活的有机体 ,如果能成功地将种子扩散到适宜的生境 ,将会在生存竞争中获得优势。在长期的进化过程中 ,不同的植物依赖不同的媒介传播种子 ,如风传播 ,水传播 ,或自身的力量传播 (重力、弹爆力等 )。在很多情况下也依靠动物完成种子扩散 ,即所谓的动物传播。根据依赖的动物对象不同可以分为蚁传播 ,鸟传播 ,哺乳类传播 ,以及鱼传播 ,爬行类传播等。哺乳类中传播种子的类群主要包括翼手类 ,灵长类和啮齿类。由于能够飞行 ,热带食果实的蝙蝠(属翼手类 )对种子的传播作用最明显 ,研究得也最多 ,源于蝙蝠类的传播特称为ｃｈｉｒ…     

动物与植物种子更新的关系Ⅱ．动物对种子的捕食、扩散、贮藏及与幼苗建成的关系

植物的繁殖体总是面临来自各类生物（如昆虫、脊椎动物、真菌）的捕食风险。因动物捕食引起的种子死亡率影响植物的适合度、种群动态、群落结构和物种多样性的保持。种子被捕食的时间和强度成为植物生活史中发芽速度、地下种子库等特征的主要选择压力,而种子大小、生境类型等因素也影响动物对植物种子的捕食。捕食者饱和现象被认为是植物和种子捕食者之间的高度协同进化作用的结果,是限制动物破坏种子、提高被扩散种子存活率的一种选择压力。大部分群落中的大多数植物种子被动物扩散。种子扩散影响种子密度、种子被捕食率、病原体攻击率、种子与母树的距离、种子到达的生境类型以及建成的植株将与何种植物竞争,从而影响种子和幼苗的存活,最终影响母树及后代植物的适合度。种子被动物扩散后的分布一般遵循负指数分布曲线,大多数种子并没有扩散到离母树很远的地方。捕食风险、生境类型、植被盖度均影响动物对种子的扩散。植物结实的季节和果实损耗的过程也体现了其对扩散机会的适应。许多动物有贮藏植物种子的行为。动物贮藏植物繁殖体的行为,一方面调节食物的时空分布,提高了贮食动物在食物缺乏期的生存概率;另一方面也为种子萌发提供了适宜条件,促进了植物的扩散。于是,植物与贮食动物形成了一种协同进化关系,这种关系可能是自然界互惠关系（mutualism)的一种。影响幼苗存活和建成的因子包括种子贮蒇点的微生境、湿度、坡向、坡度、林冠盖度等。许多果食性动物吃掉果肉后,再将完好的种子反刍或排泄出来。种子经动物消化道处理后,发芽率常有所提高。     

The effects of seed abundance on seed predation and dispersal by rodents in <Emphasis Type="Italic">Castanopsis fargesii</Emphasis> (Fagaceae)

Based on the animal dispersal hypothesis and the predator satiation hypothesis, we examined the effects of seed abundance at both population (i.e., mast seeding) and community levels on seed predation and dispersal of Castanopsis fargesii (Fagaceae), a rodent-dispersed mast species in Eastern Asia. In a subtropical evergreen broadleaved forest in the Dujiangyan region of Sichuan Province, China, individual seeds with coded tin tags were tracked in two contrasting stands (seed-poor and seed-rich) over two years (2000, a low-seed year; 2001, a high-seed year). Our results showed that: (1) small rodents did not harvest the tagged seeds of C. fargesii more rapid in the high-seed year than in the low-seed year in either stand. But, seed harvest was significantly faster in the seed-rich stand than in the seed-poor stand. (2) The removal proportion was significantly lower in the high-seed year than in the low-seed year for either stand, but the removal proportion was slightly higher in the seed-poor stand than in the seed-poor stand. This indicates that high seed abundance decreases seed removal (predator satiation hypothesis). (3) There were only small differences about seed caching, seed survival and seedling establishment of C. fargesii between years and stands. During the survey, no cached seeds survived to geminate in the spring for both stands and years. (4) Mean dispersal distances of the cached seeds are much shorter in the high-seed year (3.1 m) than in the low-seed year (8.1 m) in the seed-rich stand, though similar trend is not examined in the seed-poor stand. Our results indicate that seed predation and dispersal of C. fargesii are influenced by both mast seeding and community-level seed abundance, which is not completely consistent with either the animal dispersal hypothesis or the predator satiation hypothesis, but seems more related to the predator satiation hypothesis.     

The intermediate dispersal hypothesis: seed dispersal is maximized in areas with intermediate usage by hoarders

Seed dispersal and predation are paramount for tropical plant diversity. When encountered by scatter-hoarding frugivores, seeds can be either eaten, dispersed or ignored. But even after dispersal, seed caches are still subjected to predation. Many factors are known to influence these dynamics; however, how frequently hoarders use certain patches has seldom been related to cache predation rates. We used the interaction between agoutis (Dasyprocta leporina), a scatter-hoarding rodent, and Joannesia princeps, a tropical tree, as a model to investigate how the number of visits by hoarders in certain areas influences cache predation and seed fate. Camera-traps were used for 30 days in twenty different locations in Tijuca National Park to assess number of visits by agoutis. Thereafter, we placed seed piles on the same areas and determined their fate using the spool-and-line method to track seeds for over one hundred days. We found a non-linear relationship between how often an area is used by hoarders and the final proportion of dispersed seeds. At areas with a low number of visits, proportion of dispersed seeds was low due to low removal. As frequency of visits by hoarders increased, seed removal and the number of dispersal events increased but so did cache predation. Thus, in areas intensively used by hoarders, high cache predation resulted in a low number of dispersed seeds that remained alive in caches. As a result, dispersal was maximized in areas with intermediate use by scatter-hoarders, where there was a balance between primary seed dispersal and cache predation.

     

啮齿动物对不同林木种子的搬运和取食微生境选择机制

不同植物种子依靠不同的方式实现扩散,啮齿动物对林木种子搬运后在取食点微生境和贮藏方式的选择存在偏好,研究其贮藏行为与微生境的关系是探究幼苗建成的关键。在秦岭中段火地塘林区,采用标签标记法,以锐齿槲栎、华山松和油松种子为材料,探究了小型啮齿动物对松栎混交林建群种种子扩散过程的影响。结果表明:1)油松种子原地取食率显著高于锐齿槲栎和华山松种子,且啮齿动物更倾向于搬运后取食(60%)和埋藏(4.33%)华山松种子,搬运后取食距离也为华山松最大(2.49 m);锐齿槲栎小种子被搬运后埋藏的距离最大(4.92 m)。2)除华山松种子外,其他类型种子被搬运后单个取食的比例均在85%以上;油松种子不存在埋藏点,而其他类型种子90%以上均以单个形式被埋藏。3)大部分种子被啮齿动物搬运后选择在裸地丢弃;锐齿槲栎大种子(87.5%)、小种子(78.57%)和华山松种子(53.33%)较大比例被啮齿动物埋藏在灌丛下方,埋藏在裸地的种子较少。4)大部分种子在灌丛下方被取食,仅华山松种子被啮齿动物搬运到洞穴取食;除油松种子被大量原地取食外,其他类型种子被搬运到取食点的种子比例基本呈现由微生境植被复杂到简单(灌丛—草丛—灌丛边缘—裸地)而逐渐减小的趋势。种子的营养价值及取食和搬运过程中啮齿动物付出的成本是影响种子命运的关键性因子,且啮齿动物对种子埋藏和取食地点的微生境存在较明显的选择性。     

Seed dispersal by carnivores in temperate and tropical dry forests

The seed dispersal mechanisms and regeneration of various forest ecosystems can benefit from the actions of carnivores via endozoochory. This study was aimed to evaluate the role of carnivores in endozoochory and diploendozoochory, as well as their effect on seed viability, scarification, and germination in two forest ecosystems: temperate and tropical dry forest. We collected carnivore scat in the Protected Natural Area of Sierra Fría in Aguascalientes, Mexico, for 2 years to determine the abundance and richness of seeds dispersed by each carnivore species, through scat analysis. We assessed seed viability through optical densitometry using X‐rays, analyzed seed scarification by measuring seed coat thickness using a scanning electron microscope, and evaluated seed germination in an experiment as the percentage of seeds germinated per carnivore disperser, plant species, and forest type. In the temperate forest, four plant species (but mainly Arctostaphylos pungens) were dispersed by four mammal species. The gray fox dispersed the highest average number of seeds per scat (66.8 seeds). Bobcat dispersed seeds through diploendozoochory, which was inferred from rabbit (Sylvilagus floridanus) hair detected in their scats. The tropical dry forest presented higher abundance of seeds and richness of dispersed plant species (four species) than in the temperate forest, and the coati dispersed the highest number of seeds (8,639 seeds). Endozoochory and diploendozoochory did not affect viability in thick‐testa seeds (1,480 µm) in temperate forest and thin‐testa seeds (281 µm) in tropical dry forest. Endozoochory improved the selective germination of seeds. Nine plant species were dispersed by endozoochory, but only one species (Juniperus sp.) by diploendozoochory. These results suggest that carnivores can perform an important ecological function by dispersing a great abundance of seeds, scarifying these seeds causing the formation of holes and cracks in the testas without affecting viability, and promoting the selective germination of seeds.     

Seed dispersal and predation in the endemic Atlantic rainforest palm <Emphasis Type="Italic">Astrocaryum aculeatissimum</Emphasis> across a gradient of seed disperser abundance

Tropical forests have been subject to intense hunting of medium and large frugivores that are important in dispersing large-seeded species. It has been hypothesized that in areas with extinction or low abundance of medium and large-bodied animals the density of small rodents may increase. Therefore, this increment in the density of small rodents may compensate for the absence or low abundance of medium and large frugivores on seed removal and seed dispersal. Here, we fill up this gap in the literature by determining if seed removal, seed dispersal, and seed predation by small rodents (spiny rats, Trinomys inheringi and squirrels, Sciurus ingrami) are maintained in defaunated areas. We accessed seed removal, seed dispersal, seed predation, and seedling recruitment of an endemic Atlantic rainforest palm, Astrocaryum aculeatissimum, in a gradient of abundance of agoutis. We found that seed removal, scatter hoarding, and seed predation increase with the abundance of agoutis. In contrast, the proportion of dispersed but non-cached seeds decreased with the abundance of agoutis. We did not find any effect of the abundance of agoutis on seed dispersal distance, but we did find a positive trend on the density of seedlings. We concluded that small rodents do not compensate the low abundance of agoutis on seed removal, scatter hoarding, and seed predation of this palm tree. Moreover, areas in which agoutis are already extinct did not present any seed removal or scatter hoarding, not even by small rodents. This study emphasizes both the importance of agoutis in dispersing seeds of A. aculeatissimum and the collapse in seed dispersal of this palm in areas where agoutis are already extinct.     

Ignorant seed predators and factors affecting the seed survival of a tropical palm

In addition to acting as seed predators, some terrestrial mammals bury seeds via scatter hoarding. This study system used two permanent plots in examining the interaction between small rodents and the seeds of the palm Astrocaryum mexicanum. We tested how experimental burial, and fruiting status of the parent, distance to the parent, seed size, and microsite characteristics affect the survival of these seeds. Up to 34% of the buried seeds that were exposed only to ignorant rodent foragers (individuals not responsible for burial) survived. In comparison, less than 1% of seeds buried by scatter hoarding rodents survived in previous studies, a percentage that is comparable to the low survival of unburied seeds in this study (<2%). Although unburied seeds had very low survival, increasing distance and/or seed density positively affected survival of unburied seeds. Distance to parent had no effect on buried seed survival.
Buried seed survival was most strongly and significantly determined by the fruiting status of the trees under which they occurred. Seeds experienced significantly greater predation if buried under “parent” trees that fruited during the experiment. Buried seed survival was also negatively affected by germination, as germination may signal the presence of a seed to foraging rodents. There was some indication of a positive effect of tree density on seed survival between the two plots, whereas differences in rodent abundance appear to have no effect on seed survival. Seed size and microsite characteristics had no significant effect on buried seed survival, likely due to the greater proportional effects of other factors and the longevity of A. mexicanum seeds. The results of this study were used to generate a hypothetical causal network showing how comparatively low recovery of buried seeds by ignorant foragers – combined with processes determining the removal of scatter hoarding foragers from their scattered seed caches – may affect seedling recruitment in A. mexicanum.     

Seed predation,disease and germination on landslides in Neotropical lower montane wet forest

Abstract. Seed mortality (caused by predators and pathogens) and germination were compared between Puerto Rico and Costa Rica on landslides in lower montane wet forest. Seeds of six common species on five Puerto Rican landslides and four common species on two Costa Rican landslides were used with a Cecropia species and a Gonzalagunia species included at both sites. In the Puerto Rican experiments Cecropia schreberiana was the only species to show significant seed predation (which was due to insects), pathogens grew from all species and fewer seeds were lost to predators than pathogens. Also in Puerto Rico mean germination across all species was 57 % before dispersal (filled seeds collected while still on the tree) and 71 % after, with Phytolacca rivinoides seeds germinating most abundantly, followed in descending order by Ocotea leucoxylon, Cecropia spec, Miconia racemosa, Palicourea riparia and Gonzalagunia spicata. In the Costa Rican experiments three species had significant predation: Cecropia polyphlebia and Urera caracasana (both due to insects) and Witheringia coccoloboides (due to mammals); pathogenic disease caused more seed loss than predation, and germination was high (61 % pre-dispersal, 69 % post-dispersal). Similarities between these island and mainland sites included (1) percentage of seeds lost to predation and percentage lost to pathogens (all in the 5–15 % range), (2) generalist pathogens which claimed more seeds than predators and (3) majority germination with a general increase after dispersal. Finally sites were dissimilar only in the number of species with significant predation loss and whether it was by insects or mammals, casting doubt on the traditional island/mainland dichotomy.