Fruit Traits in Baboon Diet: A Comparison with Plant Species Characteristics in West Africa

Primate fruit choice among plant species has been attributed to different morphological plant and fruit characteristics. Despite a high abundance of animal-dispersed plant species in the savanna–forest mosaic of West Africa, few data are available on the interplay between morphological fruit traits and primate fruit consumers in this ecosystem. We tested whether olive baboons (Papio anubis) at Comoé National Park, north-eastern Ivory Coast, prefer fruit species with particular characteristics relative to the availability of these traits among the woody plant species at the study site. Specifically we were interested in the suites of traits that best predict fruit choice and seed handling by baboons. The baboons ate fruit/seeds from 74 identified plant species, representing 25 percent of the regional pool of woody plant species. They preferred trees to shrubs and lianas as fruit sources. Otherwise, baboons seemed to consume whatever fruit type, color, and size of fruit and seeds available, though they especially included larger fruit into their diet. Against expectations from the African bird–monkey fruit syndrome of brightly colored drupes and berries, baboons ate mostly species having large, dull-colored fruit. Fruit type and color best described whether baboons included a species into their diet, whereas fruit type and seed size best predicted whether baboons predated upon the seeds of their food plant species. As most plant species at the study site had medium-sized to large fruits and seeds, large frugivores like baboons might be particularly important for plant fitness and plant community dynamics in West African savanna–forest ecosystems.     

濒危树种闽楠种子和幼苗生态学研究

从1995至1998年,通过种子收集器布设、不同程度圈围的样方处理、实验室和野外发芽实验、相邻格子样方调查和土壤中种子的筛选等一系列方法,对福建罗卜岩闽楠种子质资源保护区内闽楠为优势的常绿阔叶林群落中闽楠种群的果实、种子库、种子散布、果实／种子捕良、种子萌发和幼苗存活等进行了研究。结果表明：1995至1996年果实量为116个／m^2,成熟高峰期在1996年1月中旬。1997年至1998年为103个/m^2,成熟高峰期在1997年12月中旬。1997年至1998年,绝大部分（90．1％）的果实直接从母树上掉落,低于9．5％的果实由鸟类传播。实验室的种子发芽率为93％,而在野外仅为12％。野外发芽率低多由种子霉烂引起。闽楠的种子寿命较短,其种子在次年4月初开始萌发,7月之后地面种子即丧夫活力。动物捕食不仅影响种子的存活,同时也影响幼苗的死亡率,未经圈围与样地中被取食的果实和幼苗数量有显著差异。在新形成的林窗中或远离母树均降低了果实的迁移和幼苗的死亡率。但各因素交互作用有所不同。密度相关效应发生在2－3个月的幼苗阶段,之后,密度高低对果实被取食和幼苗死亡没有显著的影响。这些结果表明,闽楠种子和幼苗经历较大的环境压力,即在高湿度生境中母树树冠下的种子不仅易受到土壤病原菌的感染,同样容易遭受动物所捕食,从而导致较低的野外种子发芽率和幼苗存活率。这是闽楠致濒的重要原因之一。     

Paternal transmission of a seed size reduction gene varies with age of a primary transformant and seed set is also influenced by gene expression in maternal tissues

We have constructed chimaeric genes that are expressed in embryo and endosperm compartments of the seed, induce dominant seed lethality and have potential to reduce seed size in 75% of seeds within a fruit such as Citrus [7]. The genes are not entirely seed-specific as a proportion of primary test tobacco transformants containing their gene were fully male-sterile [7]. Here we investigated why a proportion of apparently male-fertile transgenic plants showed segregation distortion from the 75% seed lethality expected for a single dominant gene. Reciprocal crosses were conducted between pollen fertile, primary tobacco transformants containing various copies of the CG1-400-RNase gene [7] and wild-type tobacco plants to examine the transmission of the gene through maternal and paternal gametes and also the effects of gene dosage in embryo and endosperm compartments on seed viability and phenotype. Pollen viability, seed set and seed phenotype were examined over a 16 month period to assess stability of gene expression in primary transformants because woody, fruit crops containing these genes will be vegetatively propagated from primary transformants. In male-fertile transformants, the gene was observed to be expressed to varying degrees post-meiotically in pollen over the time period examined resulting in lethality of transgenic pollen and reduced paternal transmission. A variable, low-level maternal expression component was also detected that resulted in seed lethality and influenced morphological variation in the seed lethal phenotype. The maternal and paternal expression components caused seed lethality to range from 50 to 75%. This study indicates the need to select for transformants with stable pollen transmission and high seed expression and raises questions in relation to possible environmental and epistatic effects on gene expression in primary, hemizygous transformants over long growth periods.     

Primate seed dispersal: Coevolution and conservation implications

Early studies of primates have demonstrated that many species rely heavily on fruit, and that primates constitute a large component of the frugivore biomass in tropical forests. Consequently, primates have long been thought to be important seed dispersers. It is only recently that studies have been conducted that have illustrated the complex nature of the interactions between fruit-eating primates and their food trees. Such studies have raised questions as to the causes and conse-quences of the intriguing differences between primate communities, the importance of other animals in the interactions (such as dung beetles and rodents that secon-darily disperse seeds), how primate-plant interactions evolve, and the significance of primates in forest regeneration and conservation. Since subsistence and com-mercial hunting of primates has heavily impacted frugivore communities, but left the physical structure of the forest relatively unaltered, studies of primate seed dispersal have important implications for the future of forests where seed dispersers have declined or disappeared.     

Nutritional and Morphological Traits of Invasive and Exotic Fleshy‐fruits in South Africa

While fleshy‐fruited invasive alien plants are recognized as some of the worst invaders on a global scale, until recently, little consideration has been given to the frugivores that feed on these fruits and, more specifically, the fruit traits, which may influence this. We investigated a series of morphological and nutritive fruit traits for ca 30 species of fleshy‐fruited invasive alien and exotic species in South Africa. Invasive alien fruit traits were compared with comparable traits of a similar sample size of indigenous fleshy fruits, which occur in the same area. Finally, the similarity of traits for the same invasive alien species was compared with those fruits in Australia. Invasive alien fleshy fruits were similar in morphology, but greater in some nutritive aspects when compared with indigenous fruits. Furthermore, they were very similar in all aspects to their counterparts in Australia. Most seeds of invasive fleshy fruits were small and light, which may explain some of their invasive success, as benefits associated with small seededness may promote invasive potential. Nutritionally, most invasive alien fleshy‐fruits were hexose‐dominant, containing low lipid and nitrogen content. While frugivore preference trends remain to be formally investigated, this study provides insights into fruit traits, which may tentatively outline why invasive fruits are universally fed on and thus successfully spread.     

Aerial seed bank in a cold desert annual-ephemeral species: Role of anatomical structure of stem and delayed fruit dehiscence in timing of seed dispersal

Why the fruits are retained on dead upright herbaceous plants and how this relates to seed dispersal and timing of germination remain unclear. Stems of the annual Euclidium syriacum (Brassicaceae) with infructescences bearing indehiscent silicles remain upright after plants die in the spring. We investigated the effect of anatomical structures of stem and pedicle and delayed silicle dehiscence on seed dispersal phenology of this species. For comparison, sections were made of the stem of the annual Goldbachia laevigata (Brassicaceae), which has stems that fall over when plants die. Compared to G. laevigata, the stem of E. syriacum has vascular bundles that are closer together, a thicker xylem and phloem, more fibers, a thicker perimedullary zone and a smaller pith diameter:stem diameter ratio. The thickened pedicle did not form an abcission layer. By late October, 5–20% of seeds were dispersed, depending on the position of infructescences on the plant. Snow covered the plants in late autumn and when it melted in mid-April many of the plants had fallen over, with a high number of seeds germinating in attached silicles; seedlings became rooted in soil. After snowmelt, 14–15% of the silicles on the remaining upright plants contained seeds; all seeds were dispersed by early July. The anatomical structures of the stem and pedicle plus the delayed dehiscence of silicles explain the presence of an aerial seed bank in E. syriacum and delay of germination of many of seeds until spring. Further, pieces of upright plants are broken off and dispersed by wind, which helps to explain the wide distribution of E. syriacum in the cold desert.     

Frugivores and cheap fruits make fruiting fruitful

Animal seed dispersal provides an important ecosystem service by strongly benefiting plant communities. There are several theoretical studies on the ecology of plant–animal seed–disperser interactions, but few studies have explored the evolution of this mutualism. Moreover, these studies ignore plant life history and frugivore foraging behaviour. Thus, it remains an open question what the conditions for the diversification of fruit traits are, in spite of the multitude of empirical studies on fruit trait diversity. Here, we study the evolution of fruit traits using a spatially explicit individual‐based model, which considers the costs associated with adaptations inducing dispersal by frugivory, as well as frugivore foraging behaviour and abundance. Our model predicts that these costs are the main determinants of the evolution of fruit traits and that when the costs are not very high, the evolution of larger fruit traits (e.g. fleshy/colourful fruits) is controlled by the choosiness and response thresholds of the frugivores as well as their numerical abundance.     

Florivory indirectly decreases the plant reproductive output through changes in pollinator attraction

Species often interact indirectly with each other via their traits. There is increasing appreciation of trait‐mediated indirect effects linking multiple interactions. Flowers interact with both pollinators and floral herbivores, and the flower‐pollinator interaction may be modified by indirect effects of floral herbivores (i.e., florivores) on flower traits such as flower size attracting pollinators. To explore whether flower size affects the flower‐pollinator interaction, we used Eurya japonica flowers. We examined whether artificial florivory decreased fruit and seed production, and also whether flower size affected florivory and the number of floral visitors. The petal removal treatment (i.e., artificial florivory) showed approximately 50% reduction in both fruit and seed set in natural pollination but not in artificial pollination. Furthermore, flower size increased the number of floral visitors, although it did not affect the frequency of florivory. Our results demonstrate that petal removal indirectly decreased 75% of female reproductive output via decreased flower visits by pollinators and that flower size mediated indirect interactions between florivory and floral visitors.     

Fruit size preference in the New Zealand pigeon (Hemiphaga novaeseelandiae)

Abstract We investigated whether the New Zealand pigeon Hemiphaga novaeseelandiae (Columbidae) exhibits size‐based preferences for fruits. We tested the hypothesis that in small‐fruited species, pigeons would prefer larger fruits, but in larger‐fruited species, this preference would reverse as the pigeons become increasingly limited by their gape size. We collected undispersed fruits and bird‐dispersed seeds of 10 plant species, some over several sites or years (13 datasets in total). We estimated the fruit size of dispersed seeds by fitting regressions of fruit diameter to seed diameter in intact fruits. We were able to predict fruit diameter from seed diameter in 12 of the 13 populations, although the relationship was stronger in single‐seeded species than in multi‐seeded species. Seven of the 12 populations tested showed a significant difference in seed diameter among undispersed and dispersed seeds. However, our results showed no consistent pattern in fruit size preference by the New Zealand pigeon and did not support our hypothesis. The large‐bodied New Zealand pigeon is generally not gape limited and fruit size preferences appear to be independent of mean fruit size.     

被子植物果实和种子化石的获取和研究：介绍B.H.Tiffney的原理和方法

近年来繁殖器官化石的研究已成为国际被子植物化石研究领域的热门之一,其研究方法和技术经过多年的探索也已日趋成熟,美国古植物学家ＢｒｕｃｅＨ．Ｔｉｆｆｎｅｙ根据自己多年的研究积累并吸收同行的经验对获取和研究分散保存在岩石中的被子植物果实和种子化石的原理和方法进行了系统的总结和介绍（Ｔｉｆｆｎｅｙ,１９９）为方便国内同行门熟悉和运用这些方法对我国可能产出的繁殖器官化石进行研究,本文特将该文中涉及实验原理     

Reproductive biology in Acacia caven (Mol.) Mol. (Leguminosae) in the central region of Argentina

Studies on Acacia caven (Mol.) Mol. in central Argentina indicate that the species is polygamous (andromonoecious), some plants having a high proportion of staminate heads. Though pollen/ovule ratios of flowers, inflorescences and plants are at a level common for 'facultative xenogamic' systems, controlled pollination shows a marked xenogamy. The ratio between pollen grains in the polyad and the maximum number of seeds per pod is close to one. Observations support the idea of the inflorescence as a specialized reproductive unit with gynoecia functioning as fixed modules in which all or none of its ovules develop into seeds. Mass flowering prior to the growing season, the absence of other floral resources in the community, the high degree of fruiting limitations and scarce insect visitation suggest that, as in other Acacia , the reproductive system of Acacia caven involves a great sacrifice of floral resources.     

Predicting which tropical tree species are vulnerable to forest disturbances

Tropical forest management often focuses on a few high‐value timber species because they are thought to be the most vulnerable in logged forests. However, other tree species may be vulnerable to secondary effects of logging, like loss of vertebrate dispersers. We examined vulnerability of tree species to loss of vertebrate dispersers in Mabira, a heavily disturbed tropical rainforest in Uganda. Fruit characteristics and shade tolerance regimes of 269 tree species were compiled. Stem densities of tree species producing fruits of various sizes and having different shade tolerance regimes were computed for Mabira and compared with densities of conspecifics in Budongo, a less disturbed forest with similar floral composition. Seventy per cent of tree species in Mabira are animal‐dispersed, of which 10% are large‐fruited light demanders. These species are the most vulnerable because they rarely recruit beneath adult conspecifics and are exclusively dispersed by large vertebrates, also vulnerable in heavily disturbed forests. Comparison of densities between Mabira and Budongo showed that large‐fruited light demanders had a lower density in Mabira. Other categories of tree species had similar densities in both forests. It is plausible that the low density of large‐fruited light demanders is due to limited recruitment caused by dispersal limitations.     

Ecology of the fruit-colour polymorphism in Rubus spectabilis

Although some studies have focused on the colour polymorphisms of flowers and fruits, little is known of their ecological and evolutionary significance. We investigated the potential contribution of several factors to the maintenance of fruit-colour polymorphism in Rubus spectabilis, a common shrub in the temperate rainforests of southeast Alaska. Fruits occur in two colours (red and orange), whose frequencies vary geographically. The two colour morphs have similar size, weight, seed load and nutrient composition. Colour preferences of avian frugivores, in the aviary and in the field, varied among individuals, but the majority favoured red fruits. Seed predators (mostly rodents) did not discriminate between seeds from different morphs. The effect of seed passage through the digestive tract of frugivores (birds and bears) on germination was similar for both morphs, although there were significant differences among frugivores. The type of soil on which the seeds are deposited influenced their germination behaviour, suggesting that some soils could favour one morph over the other. Such differences may contribute to the regional differences in frequencies of the two morphs. This study emphasizes the need to investigate fruit and seed characteristics that correlate with fruit colour; the colour preferences of consumers is only one of several selection pressures that determine the frequency distribution of fruit colours. This revised version was published online in July 2006 with corrections to the Cover Date.     

Diet and Dietary Preferences of the Southern Cassowary (Casuarius casuarius) in North Queensland, Australia

We investigated the diet of the southern cassowary (Casuarius casuarius) by identifying the seeds and fruits in fecal droppings encountered on a set of transects over 2 yr in upland rain forest in the wet tropics of North Queensland. A total of 198 droppings containing 56 plant species were found. We surveyed fleshy fruit availability over the subsequent 68 mo on transects in the same area to ascertain fruiting patterns in the study area. The number of droppings found each month did not correspond to the pattern of available fruit biomass. There was no relationship between the fruit traits of moisture content, flesh to seed mass ratio, color, or crop size to contribution of a species to the diet. During the lean fruiting season (May–July) cassowaries relied more on species that fruited continuously throughout the year as they were significantly over‐represented in droppings, while annual fruiting species were under‐represented. During months of high fruit availability (October–December), continuously fruiting species were still over‐represented in the diet but became less important while annual and biennial species became more important. Significantly more species with large fruit and large seeds appeared in the diet than expected and we confirm that the cassowary contributes to the continued dispersal of these species over long distances and in large quantities.     

绞股蓝果实的发育解剖学研究

应用石蜡切片和扫描电镜的方法,观察了绞股蓝果实与种子的发育过程,结果表明:花后12 d左右,绞股蓝合子开始进行第一次分裂,极核分裂在花后6～7 d,核型胚乳在胚的发育中被吸收,胚的发育为茄型,外珠被呈网格状花纹。假果,表面有蜡质,果壁可明显分为3层,内侧细胞后期解体,种子倒三角形,表面瘤状突起。脱落花果的结构表现为珠被萎缩和胚发育受阻。     

独行菜族8属(十字花科)植物果实及种子微形态研究

采用GMA(Glycol methacrylate,乙二醇甲基丙烯酸酯)半薄切片法,利用比较解剖学对独行菜族中代表植物8属25种果实及种子微形态结构进行观察分析。同时,以菥蓂属菥蓂为例详细介绍了假隔膜的形成过程。结果显示独行菜族果实及种子微形态特征明显,果实均为短角果,除厚壁荠属和沙芥属果实为背腹压扁外,其他属种均为两侧压扁;果实边缘有翅为独行菜族的典型特征,可分为周翅、微翅、宽翅及披针形翅;部分果皮细胞有纤维层,偶有木化,除菘蓝属、厚壁荠属、沙芥属无假隔膜结构外,其他属种均具有明显的假隔膜。种子大小为(0.8~10)mm×(0.5~2.8)mm,种皮纹饰丰富,遇水或潮湿环境可形成粘液种子。种皮通常由薄壁细胞构成,偶有木化,具内含物;除高河菜属和菥蓂属子叶与胚根的排列方式为子叶缘倚,其他属种均为子叶背倚。假隔膜为内果皮细胞向内延伸连接而成。本文完善了独行菜族果实及种子微形态结构信息,为分子系统学等其他相关研究提供基础资料。     

An analysis of diet specialization in frugivorous Pteropus poliocephalus (Megachiroptera) in Australian subtropical rainforest

Abstract The Grey-headed flying fox Pteropus poliocephalus Temminck 1825 is the only mammalian frugivore to occupy substantial areas of the subtropical rainforests of eastern Australia. The composition of the P. poliocephalus diet and diet specialization in the species are therefore pertinent to studies of trophic structure, seed dispersal and evolutionary processes in these forests. During a three-year diet study, P. poliocephalus used fruits from 44 species of canopy and edge plants. Their taxonomically diverse diet was dominated by the Myrtaceae and Moraceae. Dietary specialization by P. poliocephalus was examined using two criteria: the influence of fruit morphology on diet choice and dietary overlap with sympatric avian frugivores. There was no evidence from either approach that they were specialist feeders. Initial analyses comparing the morphological characters of diet fruits with fruits available to P. poliocephalus during the study period showed a preference for white fruits, berries, syconia and fruits with multiple seeds, and avoidance of black fruits and drupes. However, these significant results were not sustained when the confounding effects of correlations between fruit morphology and other traits were considered. All, except the response to berries, could be attributed to either avoidance by P. poliocephalus of secondary compounds in the Lauraceae or selection for the beneficial phenology of Ficus. Dietary overlap with frugivorous birds was notably high and the fruit diet of P. poliocephalus formed a subset of the avian diet. Associations between fruit colour, size and protective mechanisms have been documented in other rainforest areas and have been proposed as indicators of coadaptive relationships between vertebrate frugivores and their diet plants. However, these associations were not apparent in the morphological characters of fleshy fruits from Australian subtropical rainforest trees. An explanatory hypothesis of primarily avian influence on fruit traits is presented.     

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

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

Consequences of southern cassowary (Casuarius casuarius,L.) gut passage and deposition pattern on the germination of rainforest seeds

In Australia's tropical rainforests the endangered southern cassowary, Casuarius casuarius, L., is the largest native frugivore and many plant species, because of the size of their fruits or seeds, are thought to be largely dependent on cassowaries for their dispersal. In this study we asked whether the contribution of cassowaries to plant recruitment extends beyond removing seeds from the vicinity of the parent. To do this we conducted germination trials involving 17 rainforest plant species to test whether cassowary consumption and seed deposition pattern alter germination probability or time to germination. Twenty‐four per cent of species showed changed germination probabilities, with one species showing no germination without cassowary consumption, and 35% showed changed time to germination. However these differences did not translate into any significant effects when considered across all species. We examined gut scarification, fruit pulp removal (de‐inhibition) and deposition in faecal material as mechanisms for changing germination success; each contributed to the changed success of individual species. There was a negative effect of seed clumping on five species. We conclude that cassowary consumption can modify germination performance in a minority of rainforest plants and that the effect is generally positive. Although the effect on large seeded species was small it is most likely to be important as the cassowary is the only animal in Australia able to internally process large numbers of these seeds.