The Influence of Temporal Changes in Fruit Availability on Diet Composition and Seed Handling in Blue Monkeys (Cercopithecus mitis doggetti)1

We investigated the relation between temporally varying resources, diet composition, and seed-handling behaviors in a group of blue monkeys (Cercopithecus mitis doggetti) in a tropical montane forest of Rwanda. Changes in diet composition were related to concurrent phenological studies of fruit-producing trees, and density and abundance of tree resources within the monkey's home range. Fruit composed nearly 50 percent of the diet. Over 50 percent of the fruits eaten had juicy fleshy pulp. Observations of seed handling behavior provided insights into the role of these animals as potential seed dispersal agents. The monkeys moved the seeds of 29 species out of parent canopies by defecating seeds intact and by potentially carrying seeds in cheek pouches and dropping them later. Seeds of 18 species were found intact in fecal piles. Our study showed community-level phenology patterns did not indicate a decrease in fruit availability during the study period, but an analysis of the preferred fruits consumed by the monkeys showed distinct periods of low fruit availability. The study period included two dry seasons; only one of these produced a period of fruit scarcity for the animals. The animals employed different strategies during times of preferred fruit scarcity. They increased consumption of leaves and other fleshy fruits, and diet diversity increased, or became mainly seed predators and diet diversity decreased. The variable responses of these monkeys to changes in food availability highlights their dietary plasticity and imposes significant variations in their role as potential seed dispersers.     

Selecting replacements for invasive plants to support frugivores in highly modified sites: A case study focusing on Lantana camara

Summary Invasive plants are regarded as a major threat to biodiversity worldwide. Yet, in some cases, invasive plants now perform important ecological functions. For example, fleshy‐fruited invasive plants provide food that supports indigenous frugivore populations. How can the disparate goals of conservation versus invasive weed control be managed? We suggest using the fruit characteristics of the invasive plant to select replacement indigenous plants that are functionally similar from the perspective of frugivores. These could provide replacement food resources at sites where plants with these characteristics are part of the goal plant community and where such plants would not otherwise regenerate. Replacement plants could also redirect seed dispersal processes to favour indigenous, rather than invasive, plant species. We investigated the utility of this approach by ranking all indigenous fleshy‐fruited plant species from a region using a simple model that scored species based upon measures of fruit phenology, morphology, conspicuousness and accessibility relative to a target invasive species, Lantana (Lantana camara). The model successfully produced high scores for indigenous plant species that were used by more of the frugivores of Lantana than a random selection of plants, suggesting that this approach warrants further investigation.     

Tree Seed Fate in a Logged and Fragmented Forest Landscape, Northeastern Costa Rica1

We compared the seed fate of two animal‐dispersed, large‐seeded timber species (Dipteryx panamensis [Fabaceae] and Carapa guianensis [Meliaceae]) in logged and fragmented forests with that for continuous forest in northeastern Costa Rica. For both species, we quantified rates of seed removal (an index of vertebrate predation) and the fate of dispersed seeds (those carried away from their original location that either germinated or were not subsequently removed within three months). We predicted that (1) fewer seeds would be dispersed by vertebrates in fragmented forest than in continuous forest due to low population abundances after hunting and/or loss of suitable habitat, and (2) seed predation rates would be higher in forest fragments than in continuous forest due to high abundance of small‐bodied seed consumers. We compared three forest fragments currently managed for timber (140–350 ha) and a large reserve of continuous forest (La Selva, 1500 ha and connected to a national park). An exclusion experiment was performed (seeds placed in the open vs. seeds within semipermeable wire cages; 5 cm mesh size) to evaluate the relative roles of large and small animals on seed removal. Seed germination capacity did not differ among all four sites for both species. Removal of Dipteryx seeds was higher in forest fragments (50% removal within 10 days and related to the activity of small rodents) compared to La Selva (50% removal after 50 days). Also, more Dipteryx seeds were dispersed at La Selva than in fragmented forests. Contrary to our predictions, removal of Carapa seeds was equally high among all four sites, and there was a trend for more seeds of Carapa to be dispersed in fragments than in La Selva. Our results suggest that fragmentation effects on tree seed fate may be specific to species in question and contingent on the animal biota involved, and that management strategies for timber production based on regeneration from seed may differ between forest patches and extensive forests.     

Larger seeds in tropical floras: consistent patterns independent of growth form and dispersal mode

Many species of tropical moist forests have large seeds compared to those found in temperate floras. This could be attributed to a prevalence of woody growth forms, or adaptation to dispersal by vertebrates, or the dense shade of a closed canopy, rather than to an intrinsic tropical v. temperate difference. We compared tropical v. temperate seed mass data at two geographic scales; firstly within Australia between tropical, subtropical and temperate open woodlands, then between six tropical and temperate datasets from five continents. Within Australia we found that seed mass increased with decreasing latitude in ten out of eleven growth form-dispersal mode combinations: only wind-dispersed graminoids showed no significant effect. While the pattern of generally larger seeds in the tropical flora was associated with tropical families tending to have larger seeds than temperate families, we also found that tropical species had larger seeds than temperate cofamilials. As a preliminary test of the generality of these findings we included a further four tropical and temperate data sets in a second analysis. Average tropical seed masses were larger than temperate seed masses in all testable growth form-dispersal mode combinations, and statistically larger in five out of seven cases: unassisted, vertebrate- and wind-dispersed woody species, and unassisted and vertebrate-dispersed forbs. No difference was found for wind-dispersed forbs or for graminoids lacking dispersal structures. That an effect was found in these combinations rules out explanations based solely on characteristics of tropical v. temperate dispersers, although this may be a contributing factor. Instead we suggest that some aspect of tropical systems, for example higher metabolic costs of seedling production, has selected for a general increase in seed mass among tropical taxa.     

Fruit size in a tropical tree species: variation,preference by birds,and heritability

Ocotea tenera (Lauraceae), an understory bird-dispersed tree, produces single-seeded fruits that vary in diameter from 1.4 to 2.4 cm. Much of the variation within a population at Monteverde, Costa Rica occurred within individual trees. The relative size of fruits produced by different trees remained generally constant over an 11-year period despite slight differences between years in the average size of fruits produced by a given tree.Fruit-eating birds could thus express their preferences for particular fruit size characteristics by choosing among trees that have distinct distributions of fruit diameters, and between individually variable fruits within trees. In a field study of individually marked fruits, birds removed 46.2% of fruits; the rest of the fruits were destroyed by invertebrate (25.3%) and vertebrate (4.3%) pulp-feeders or aborted by the plant after remaining ripe but uneaten for as long as 100 days (24.2%). The four major avian seed dispersers of O. tenera each have gape widths exceeding all but the largest fruits. Birds preferred plants with greater-than-average-sized fruits; within trees, they favored larger fruits, apparently because net pulp mass increases with fruit diameter. Fruits that ripened early in the season were more likely to be removed and were removed more quickly than late-ripening fruits.Based on mother-offspring regressions of mean fruit size, the phenotypic variation in fruit diameter in O. tenera is highly heritable, indicating the potential for an evolutionary response to selection by birds. Nonetheless, directional selection on fruit size or shape is likely to be inconsistent, constrained by genetic correlations, and weak compared to selection on traits like fecundity or phenology.     

Early embryogenesis in discoglossoid frogs: a study of heterochrony at different taxonomic levels

We studied the timing of the early development of the cranial neural crest, neural tube differentiation, somite formation and body elongation in the closely related frog species Discoglossus montalenti , Discoglossus galganoi , Discoglossus pictus , Bombina orientalis , Bombina variegata and Bombina maxima . Despite their different egg and embryo sizes, event pairing shows that there are only very few heterochronies among the three Bombina species and the three Discoglossus species in early developmental events. Instead, differences become apparent at the genus level (interpreted as phylogenetic signal). For example, the separation into a mandibular and a hyoid portion of cranial neural crest could be seen earlier in the Discoglossus than in the Bombina species when compared with the development of the neural tube. The development of the cranial neural crest streams also shifts (relatively) forward in time when compared with body elongation and somite formation in the Discoglossus species in contrast to the Bombina species. Comparisons with other species show that attempts to correlate heterochronies in early development with life history changes should take into account the possibility of a rather high evolvability, i.e. a high probability of convergence in the timing of these early events.     

Differences in caste dimorphism among three hornet species (Hymenoptera: Vespidae): forewing size, shape and allometry

Caste shape dimorphism (CShD) has previously been studied in wasps through comparison of different body parts, originating from different imaginal discs. Using geometric morphometrics with a new protocol for measuring wings of pinned specimens from natural history collections, we tested CShD of three hornet species in an organ developed from a single imaginal disc: the forewing. Gaussian mixture models retrieved most castes and species levels, confirming that caste is an important component of wing variations in females of these hornets. Size and allometry - the influence of size on shape - contribution to wing dimorphism between castes was major, but failed to explain the entire shape dimorphism. This deviation from simple allometric scaling was not similar in the three species: in Vespa tropica, allometric directions in the shape space differed between castes, whereas in V. crabro and V. velutina, they were similar but a significant part of CShD resulted from lateral transpositions. These results clearly indicate that queens are not just enlarged workers. They also support that the different patterns of CShD may result from different developmental mechanisms. Finally, they highlight that even in a highly social group like hornets, there is still variation in caste dimorphism among species.     

Jack of all trades masters novel host plants: positive genetic correlations in specialist and generalist insect herbivores expanding their diets to novel hosts

One explanation for the widespread host specialization of insect herbivores is the 'Jack of all trades-master of none' principle, which states that genotypes with high performance on one host will perform poorly on other hosts. This principle predicts that cross-host correlation in performance of genotypes will be negative. In this study, we experimentally explored cross-host correlations and performance among families in four species (two generalist and two specialist) of leaf beetles (Cephaloleia spp.) that are currently expanding their diets from native to exotic plants. All four species displayed similar responses in body size, developmental rates and mortality rates to experimentally controlled diets. When raised on novel hosts, body size of larvae, pupae and adults were reduced. Development times were longer, and larval mortality was higher on novel hosts. Genotype × host-plant interactions were not detected for most traits. All significant cross-host correlations were positive. These results indicate very different ecological and evolutionary dynamics than those predicted by the 'Jack of all trades-master of none' principle.     

Factors Limiting Tropical Rain Forest Regeneration in Abandoned Pasture: Seed Rain,Seed Germination,Microclimate, and Soil1

Vegetation, seed rain, seed germination, microclimate, and soil physical and chemical parameters were measured in a recently abandoned pasture and adjacent primary rain forest in southern Costa Rica. The goal of this study was to assess the importance of these factors in limiting forest regeneration in abandoned pastures. Seed rain of animal dispersed species decreased dramatically in the pasture >5 m from the forest/pasture edge; fewer wind dispersed seeds fell in the pasture than in the forest, but the difference was much less than for animal dispersed seeds. Percent seed germination of most species studied was similar in the forest and in pasture with grasses; seed germination was lower during the dry season in areas of pasture cleared of grasses. Air temperature, vapor pressure deficit (VPD), and photon flux density (PFD) were much higher in the pasture than in the forest at 1 m above the ground. VPD and PFD at ground level and soil temperature were similar in the pasture and the forest, indicating that pasture grasses strongly modify microclimatic conditions near the soil surface. The lowest gravimetric water content recorded in the pasture during the dry season was 0.5 and leaf relative water contents of the two species measured in the forest and pasture were identical, suggesting that plants in the pasture were not water stressed. Levels of most soil nutrients were lower in the pasture as compared to the forest; however, aboveground and root biomass for seedlings grown in pasture and forest soils did not differ significantly. Although a number of factors impede forest recovery in abandoned pastures, these results suggest that the most imporrant limitation is lack of seed dispersal.     

The Importance of Where to Dump the Refuse: Seed Banks and Fine Roots in Nests of the Leaf‐Cutting Ants Atta cephalotes and A. colombica1

The location of the nutrient‐rich organic refuse produced by a leaf‐cutting ant colony varies among ant species. Atta cephalotes locate their organic refuse in subterranean chambers, whereas A. colombica place their organic refuse on the soil surface near the nest. We studied the effect of the absence or presence of external organic refuse on the abundance of fine roots and seed bank composition in the superficial horizons of ant nests. We sampled soils from ant nests or dumps and adjacent areas of 15 adult nests of A. cephalotes at La Selva (LS), Costa Rica, and of 15 of A. colombica nests on Barro Colorado Island (BCI), Panama. Soils from A. cephalotes nests did not differ from adjacent soils in abundance of fine‐root and seed diversity. In contrast, organic refuse from A. colombica nests was less diverse in seed composition (due to the great abundance of Miconia argentea) and had a greater abundance of fine roots than adjacent areas. Thus the external location of the ant‐nest organic refuse is potentially important in determining the different types of plant recolonization in abandoned or dead ant nests. The relative abundance of these Atta species may influence the structure and/or composition of tropical forests.     

The Structure of Ground Beetle Assemblages (Coleoptera: Carabidae) at Fruit Falls of Melastomataceae Trees in a Brazilian Terra Firme Rain Forest1

Although most Carabidae are recognized as predacious, recent studies have shown that some tropical rain forest Carabidae use the small seeds of fallen fruit from some tree species as a key food resource (particularly figs [Moraceae] but also those from other families). Assemblages of carabid beetles (Coleoptera: Carabidae) at fruit falls of the Melastomataceae tree genera, Bellucia, Loreya, and Miconia, were studied in a terra firme rain forest near Manaus (Amazonia, Brazil) from February 1992 to August 1996. Of 5483 specimens of 14 carabid species collected at 53 fruit falls, 98.2 percent of the collected beetles represented the seed-feeding genus Notiobia. Four of the 7 Notiobia species were found to reproduce at Melastomataceae fruit falls. The most abundant species, N. glabrata, was found to be a Melastomataceae seed generalise while N. maxima seemed to prefer the seeds of Bellucia, and N. aulica and Af. umbrifera the seeds of Miconia. The presence of larvae indicated reproduction of N. glabrata at fruit falls of all three tree genera and reproduction of N. maxima at those of Bellucia. Fruit falls of different species and genera of Melastomataceae appeared to occur at different times of die year and not continuously throughout the year. The seed-feeding Notiobia species, which specialize on Melastomataceae, used fruit falls of figs as migratory “stepping stones” during periods when there were no Melastomataceae fruit falls. The comparable suite of fig seed-feeding specialist species of Notiobia did not appear to do the reverse and seemed to avoid Melastomataceae fruit falls. The carabid assemblage at fig fruit falls was more diverse than the assemblage at those of Melastomataceae. Among the Melastomataceae, carabid species richness was highest at Miconia fruit falls and lowest at Loreya fruit falls. The three most common predatory species caught at Ficus fruit falls were also caught at those of Melastomataceae and appeared to reproduce at both fruit fall types. The proportion of the carabid assemblage accounted by predatory species was much lower at Melastomataceae (1.8%) than at fig fruit falls (8%).     

Comparative palynology and seed morphology in annual pansies (Viola sect. Melanium,Violaceae): implications for species delimitation

Abstract

This integrated study provides new insights into pollen and seed morphology and pollen heteromorphism of four closely related annual taxa of Viola sect. Melanium. The plant material, both fresh and dried, was collected in Italy and studied using light and scanning electron microscope. Palynological data for V. hymettia together with a detailed comparative analysis of seed morphology and micromorphology of the four species are reported for the first time. Results of this work highlight some pollen and seed features as useful diagnostic characters. The pollen size proves to be of diagnostic value to easily separate V. kitaibeliana, having the smallest pollen grains, from the others, especially from V. arvensis with the largest ones. Exine ornamentation is microreticulate, showing no relevant differences among species. We can partially confirm the diagnostic value of the prevailing pollen morph as it can be useful only for V. arvensis (five-aperturate) versus V. tricolor and V. hymettia (four-aperturate). The macro- and micromorphology of seeds provide additional useful distinguishing characters. Particularly, seed size was found to be a good delimitating character, especially to distinguish V. kitaibeliana (with the smallest seeds) from V. arvensis, and easy to be measured with no need of particular equipment.     

Light requirements for regeneration in tropical forest plants: Taxon-level and ecological attribute effects

Abstract Understanding co-adapted traits and strategies in organisms has been increasingly shown to require a broader view of structure, function, taxonomic affiliation (phylogeny) and their possible interactions. In order to understand mechanisms of recruitment dynamics in tropical forest, nine juvenile (fruit, seed and seedling) and seven adult ecological and morphological traits were examined in 89 woody plant species, from 75 genera, 35 families and 21 orders. The species were from tropical rainforest in northern Queensland and varied widely in light requirements and hence regeneration niches. The importance of taxonomic (phylogenetic) constraints was considered using various techniques. Data were collected mostly from herbarium specimens, augmented by observations in the field. The main source of variation in most traits was at the species level, although fruit size and light requirement varied most at the genus level, and flower size at the level of order. Covariation among attributes increased in strength with increasing taxonomic level. Cluster analyses of the species arising either from five or three groups reflected juvenile rather than adult attributes. However, these groups were not morphologically distinct. Sets of juvenile traits poorly predict adult functions (and vice-versa) at species, genus and family levels, but do so more strongly at the order level. Analyses of variance and multivariate analyses, with and without the effects of phylogeny, indicated that fruit size, seed size, hardness of seed coat, seedling cotyledon function, fruit maturation period and, to a lesser extent, seed number per fruit and wood density could predict light requirement, and hence regeneration niche. Flower size showed a strong phylogenetic effect, and thus is conserved along taxonomic lines. It is concluded that as different traits may vary at different taxonomic levels, the influence of phylogeny on interspecific variation needs to be routinely recognized by ecologists involved in comparative analyses of plant life-history functions.