Fallback foods and dietary partitioning among Pan and gorilla

Recent findings on the strong preference of gorillas for fruits and the large dietary overlap between sympatric gorillas and chimpanzees has led to a debate over the folivorous/frugivorous dichotomy and resource partitioning. To add insight to these arguments, we analyze the diets of sympatric gorillas and chimpanzees inhabiting the montane forest of Kahuzi-Biega National Park (DRC) using a new definition of fallback foods (Marshall and Wrangham: Int J Primatol 28 [2007] 1219–1235). We determined the preferred fruits of Kahuzi chimpanzees and gorillas from direct feeding observations and fecal analyses conducted over an 8-year period. Although there was extensive overlap in the preferred fruits of these two species, gorillas tended to consume fewer fruits with prolonged availability while chimpanzees consumed fruits with large seasonal fluctuations. Fig fruit was defined as a preferred food of chimpanzees, although it may also play a role as the staple fallback food. Animal foods, such as honey bees and ants, appear to constitute filler fallback foods of chimpanzees. Tool use allows chimpanzees to obtain such high-quality fallback foods during periods of fruit scarcity. Among filler fallback foods, terrestrial herbs may enable chimpanzees to live in small home ranges in the montane forest, whereas the availability of animal foods may permit them to expand their home range in arid areas. Staple fallback foods including barks enable gorillas to form cohesive groups with similar home range across habitats irrespective of fruit abundance. These differences in fallback strategies seem to have shaped different social features between sympatric gorillas and chimpanzees. Am J Phys Anthropol 140:739–750, 2009. © 2009 Wiley-Liss, Inc.     

Diet and seasonal changes in sympatric gorillas and chimpanzees at Kahuzi–Biega National Park

Based on 8 years of observations of a group of western lowland gorillas (Gorilla beringei graueri) and a unit-group of chimpanzees (Pan troglodytes schweinfurthii) living sympatrically in the montane forest at Kahuzi–Biega National Park, we compared their diet and analyzed dietary overlap between them in relation to fruit phenology. Data on fruit consumption were collected mainly from fecal samples, and phenology of preferred ape fruits was estimated by monitoring. Totals of 231 plant foods (116 species) and 137 plant foods (104 species) were recorded for gorillas and chimpanzees, respectively. Among these, 38% of gorilla foods and 64% of chimpanzee foods were eaten by both apes. Fruits accounted for the largest overlap between them (77% for gorillas and 59% for chimpanzees). Gorillas consumed more species of vegetative foods (especially bark) exclusively whereas chimpanzees consumed more species of fruits and animal foods exclusively. Although the number of fruit species available in the montane forest of Kahuzi is much lower than that in lowland forest, the number of fruit species per chimpanzee fecal sample (average 2.7 species) was similar to that for chimpanzees in the lowland habitats. By contrast, the number of fruit species per gorilla fecal sample (average 0.8 species) was much lower than that for gorillas in the lowland habitats. Fruit consumption by both apes tended to increase during the dry season when ripe fruits were more abundant in their habitat. However, the number of fruit species consumed by chimpanzees did not change according to ripe fruit abundance. The species differences in fruit consumption may be attributed to the wide ranging of gorillas and repeated usage of a small range by chimpanzees and/or to avoidance of inter-specific contact by chimpanzees. The different staple foods (leaves and bark for gorillas and fig fruits for chimpanzees) characterize the dietary divergence between them in the montane forest of Kahuzi, where fruit is usually scarce. Gorillas rarely fed on insects, but chimpanzees occasionally fed on bees with honey, which possibly compensate for fruit scarcity. A comparison of dietary overlap between gorillas and chimpanzees across habitats suggests that sympatry may not influence dietary overlap in fruit consumed but may stimulate behavioral divergence to reduce feeding competition between them.     

Feeding Ecology of <Emphasis Type="Italic">Propithecus diadema</Emphasis> in Forest Fragments and Continuous Forest

Forest fragmentation is viewed as a serious threat to primates, yet whether or not it can disrupt food resources and cause energetic stress remains largely untested. I present the results of a 12-mo study of the feeding ecology of Propithecus diadema in fragmented and continuous forest at Tsinjoarivo, eastern Madagascar. Two continuous forest groups had higher dietary diversity and ate more fleshy fruit, but during the dry season, diversity was reduced and they relied heavily on mistletoe (Bakerella clavata). In contrast, 2 groups in fragments employed the lean season strategy of eating mistletoe year-round; the fruiting tree species that sustain continuous forest groups through the rainy season were largely absent. As expected, intersite dietary overlap was highest in the dry season. The level of specialization was high: fragment groups devoted 30–40% of feeding time to Bakerella clavata, compared to 28–30% in continuous forest. The major characteristic of Bakerella clavata enabling it to be an important fallback or staple resource, or both, is its extended phenology. The difference in resource utilization between sites may have important implications for nutritional status, as well as ranging and social behavior, largely owing to the small size and high abundance of feeding patches of Bakerella. Understanding resource shifts in fragments can shed light on socioecological questions by providing comparisons between continuous forest and fragment populations with differing diets and resource distributions. In addition, understanding dietary shifts in fragments can aid in species-specific conservation efforts, while contributing to a better understanding of the considerable interspecific variability of primates in responses to fragmentation.     

Transfer and Acclimatization Effects on the Behavior of Two Species of African Great Ape (<Emphasis Type="BoldItalic">Pan troglodytes</Emphasis> and <Emphasis Type="BoldItalic">Gorilla gorilla gorilla</Emphasis>) Moved to a Novel and Naturalistic Zoo Environment

Studying the effects of moving animals to new enclosures is of value to both captive managers and to scientists interested in the complex interplay between environment and behavior. Great apes represent some of the greatest challenges in this regard. Given the cognitive sophistication of these species and the substantial investments in new primate facilities, these investigations are particularly important. Using post-occupancy evaluation (POE) methodology, we compared behavior exhibited by chimpanzees (Pan troglodytes) and gorillas (Gorilla gorilla gorilla) in indoor hardscape-type exhibits to behavior of the same individuals in new naturalistic enclosures with outdoor access. In the new facility, chimpanzees showed decreases in the frequency of abnormal behaviors and visual monitoring of humans (attention behaviors) whereas gorillas exhibited reduced agonism as well as decreased attention behaviors. Both gorillas and chimpanzees demonstrated higher rates of inactivity after transfer to the new facility. All subjects in addition demonstrated transitory changes in behavior after the move to the new facility (higher rates of scratching in yr 1 than in subsequent years), indicating a period of acclimatization. Seasonal effects on feeding behavior and activity levels (both species were more active in the winter) were evident as well. The results indicate that behavioral adjustment to a new facility is an extended process for both species and that seasonal effects should be considered in longitudinal analyses of acclimatization. Behavioral patterns supported the benefits of naturalistic, functional exhibit spaces and the utility of post-occupancy evaluations in assessing captive animal welfare.     

Bed and bed-site reuse by western lowland gorillas (<Emphasis Type="Italic">Gorilla g. gorilla</Emphasis>) in Moukalaba-Doudou National Park,Gabon

In this paper we describe bed (nest) and bed-site reuse by western lowland gorillas (Gorilla g. gorilla) in Moukalaba-Doudou National Park, south-eastern Gabon. During an eight-month study 44 bed sites and 506 beds were found. Among these, 38.6% of bed sites and 4.1% of beds were reused. We analyzed the monthly frequency of bed-site reuse in relation to rainfall, fruit abundance, and fruit consumption by the gorillas. The different frequency of bed-site reuse in the rainy and dry seasons was not significant. More bed-site reuse was observed during the fruiting season than during the non-fruiting season. Results from fecal analysis suggested that gorillas ate more fruit in the fruiting season than in the non-fruiting season. The frugivorous diet of western gorillas may possibly cause gorillas to stay in some areas and, consequently, reuse their bed sites. Reuse of bed sites by gorillas suggests their frequent return to an area where preferred fruit is readily available. A higher percentage of arboreal beds may also affect bed-site reuse, because of the shortage of bed material.     

Factors Influencing the Survival of Sympatric Gorilla (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>) and Chimpanzee (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>) Nests

Accurate and precise surveys of primate abundance provide the basis for understanding species ecology and essential information for conservation assessments. Owing to the elusive nature of wild apes and the vast region of dense forest they inhabit, population estimates of central chimpanzees (Pan troglodytes troglodytes) and western lowland gorillas (Gorilla gorilla gorilla) have largely relied on surveys of their nests. Specific information about the nesting behavior of apes permits the estimation of the number of nests built (nest creation rate). Similarly, information on nest characteristics and environmental factors can be used to estimate the time it takes nests to decay (nest decay rate). Nest creation and decay rates are then used to convert nest density estimates to absolute ape densities. Population estimates that use site-specific estimates of nest creation and decay rates are more accurate and precise. However, it is common practice to generalize these conversion factors across sites because of the additional cost of studies required to gather the information to estimate them. Over a 9-mo study period, we detected and monitored the time to decay of gorilla nests (N = 514) and chimpanzee nests (N = 521) in northern Republic of Congo. We investigated the influence of nest characteristics and environmental factors on nest survivorship and estimated the mean time to nest decay (or equivalently survival) using MARK. Key factors influencing nest decay rate included ape species, forest type, nest height, mean rainfall, nest structure, nest type, and primary aspects of nest construction. Our findings highlight the synergistic effect of behavior and environment on great ape nest degradation, as well as providing practical insights for improving measures to monitor remaining populations of these endangered species.     

Seed Reingestion in Savannah Chimpanzees (<Emphasis Type="Italic">Pan troglodytes verus</Emphasis>) at Fongoli,Senegal

Coprophagy is common in captive primates but has also been reported in the wild. For example, wild apes extract and reingest items from faeces. We term this behavior seed reingestion because the dung matrix is not consumed. We assessed the importance of seed reingestion in a population of savannah chimpanzees at Fongoli, southeastern Senegal, one of the hottest and driest areas of the species’ range, where chimpanzees have a relatively narrow dietary repertoire. We observed habituated chimpanzees on 122 d during 8 mo in 2005 and 2006, employing both focal subject and scan sampling of identified individuals for 1278 h of data collection. Chimpanzees reingested seeds of 2 species: Parkia biglobosa and Adansonia digitata. Both seed species have a hard protective shell, and the embryos are rich in proteins and lipids. Chimpanzees initially ate the fruit matrix pulp and swallowed intact seeds before reingesting and chewing/destroying seeds. Seed reingestion accounted for almost 2% of feeding time. We suggest that at Fongoli this behavior may be an adaptive strategy to maximize food intake, by softening the seed’s shell and making the seed’s content more accessible.     

Comparative Feeding Ecology of Sympatric <Emphasis Type="Italic">Microcebus berthae and M. murinus</Emphasis>

Most Malagasy primate communities harbor a diverse assemblage of omnivorous species. The mechanisms allowing the coexistence of closely related species are poorly understood, partly because only preliminary data on the feeding ecology of most species are available. We provide an exemplary feeding ecology data set to illuminate coexistence mechanisms between sympatric gray and Madame Berthe’s mouse lemurs (Microcebus murinus, M. berthae). We studied their feeding ecology in Kirindy Forest/CFPF, a highly seasonal dry deciduous forest in western Madagascar. Between August 2002 and December 2007, we regularly (re-)captured, marked, and radiotracked females of both species. A combination of direct behavioral observations and fecal analyses revealed that both Microcebus species used fruit, arthropods, gum, insect secretions, and small vertebrates as food sources. However, Microcebus berthae and M. murinus differed in both composition and seasonal variation of their diets. Whereas the diet of Microcebus murinus varied seasonally and was generally more diverse, M. berthae relied mainly on insect secretions supplemented by animal matter. The differences were also reflected in a very narrow feeding niche of Microcebus berthae and a comparatively broad feeding niche of M. murinus. Resource use patterns of Madame Berthe’s and more so of opportunistic gray mouse lemurs broadly followed resource availability within the strongly seasonal dry forest. Feeding niche overlap between the 2 sympatric species was high, indicating that food resource usage patterns did not reflect niche partitioning, but can instead be explained by constraints due to food availability.     

Influence of Fruit Availability on Fruit Consumption in a Generalist Primate,the Rhesus Macaque <Emphasis Type="Italic">Macaca mulatta</Emphasis>

Fluctuations in resource availability occur in all ecosystems. To survive, species must alter their foraging strategies according to the quantity, quality, and distribution of available food. The rhesus macaque (Macaca mulatta), a commensal primate, is considered a generalist omnivore and very few studies have addressed how its feeding strategies change with respect to resource availability. We examined dietary diversity and frugivory levels in a group of rhesus macaques at the Buxa Tiger Reserve in northern India across one year. Using behavioural observations of diet and phenological monitoring, we found that although rhesus macaques fed on 107 food items including leaves, flowers, fruits, seeds, and insects, fruits made up ca. 74% of their diet. Fruit consumption correlated positively with fruit availability, but fruit preference appeared to play an important role; 16% of all the fruit species they fed on accounted for >50% of all fruit feeding observations. We suggest that afforestation programs involving preferred fruit species at the agricultural land–forest interface would prevent forest groups of rhesus macaques from gravitating toward human habitations and reduce conflict over anthropogenic resources. We further propose that the movement of certain primates in the direction of human habitations may be contingent on resource availability and food preference rather than an inherent propensity to gravitate to anthropogenic areas.     

Decreased frugivory and seed germination rate do not reduce seedling recruitment rates of <Emphasis Type="Italic">Aristotelia chilensis</Emphasis> in a fragmented forest

Habitat fragmentation reduces frugivorous bird abundance. Such a reduction may lead to a reduction in seed dispersal, thereby compromising seedling recruitment rate with far reaching consequences for plant population persistence. We assessed frugivory, seed germination, and seedling recruitment rates in a fragmented forest of central Chile by comparing a continuous forest with four forest fragments surrounded by pine plantations. Frugivory was 2.4 times higher in continuous forest than in forest fragments. Seeds eaten by birds germinated 1.7 and 3.7 times higher than non-eaten seeds from continuous forest and fragments respectively. Non-eaten seeds from continuous forest germinated 2.2 times higher than those from forest fragments, suggesting inbreeding depression. However, seedling recruitment rates at forest fragments were far higher than in continuous forest where no seedling recruited in the five years analysed. Therefore, despite forest fragmentation negatively affected frugivory, it did not translate into a decreased fitness of plants, thus highlighting the importance of considering the overall processes leading the reproductive success of plants following anthropogenic disturbances.     

Feeding selectivity of introduced hedgehogs <Emphasis Type="Italic">Erinaceus europaeus</Emphasis> in a dryland habitat,South Island,New Zealand

We describe the diet of introduced European hedgehogs Erinaceus europaeus (Linnaeus, 1758) in a New Zealand dryland system and provide the first quantitative analysis of food selectivity for this species. We also describe and compare the diets of nine hedgehogs and measure dietary overlap between these individuals. The most commonly eaten foods were beetles, including rare native species (in 94% of droppings), earwigs (92%), spiders (25%) and native skinks (14%). Remains of at least three skinks were found in one dropping. Earwigs and darkling beetles (Tenebrionidae) were the most preferred food types, and Hymenoptera and cylindrical bark beetles (Colydiidae) were least preferred. Consumption of most foods mirrored their availability. Most individuals’ diets conformed to the pattern seen at the population scale, with 21–54% of dry faecal mass derived from beetles and 13–39% from earwigs. One animal frequently consumed large amounts of fruit. Dietary overlap between pairs of individuals was high (mean Horn’s index, 0.84). This may be the result of limited opportunities to diversify in a very moisture-limited and low diversity habitat.     

Food web overlap among native axolotl (<Emphasis Type="Italic">Ambystoma mexicanum</Emphasis>) and two exotic fishes: carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) and tilapia (<Emphasis Type="Italic">Oreochromis niloticus</Emphasis>) in Xochimilco,Mexico City

Two exotic fishes, common carp (Cyprinus carpio) and tilapia (Oreochromis niloticus), were introduced more than 20 years ago into Xochimilco for aquaculture, and now dominate the system in terms of biomass and numbers. Over this same period, wild populations of the microendemic axolotl salamander (Ambystoma mexicanum) have been dramatically reduced, and it currently persists in isolated areas of this aquatic system, which is situated at the edge of Mexico City. In this study, we examine potential trophic interactions and niche overlap among two exotic fishes: carp and tilapia, and the native axolotl. Axolotl had more diverse diets and a higher trophic position compared to the exotics. Stable isotope analysis revealed substantial trophic niche overlap among axolotl and the exotics. The two exotics occupied a larger niche area than the axolotl, suggesting higher levels of omnivory and diet specialization. Current exotic fish removal efforts will further our understanding of interactions between the axolotl and exotic species, and are expected to benefit axolotl recovery efforts.     

Coprophagy-related interspecific nocturnal interactions between Japanese macaques (<Emphasis Type="Italic">Macaca fuscata yakui</Emphasis>) and sika deer (<Emphasis Type="Italic">Cervus nippon yakushimae</Emphasis>)

The influence of sympatric large animals on the sleeping behavior of primates in the wild is still largely unknown. In this study, we observed behaviors of wild Japanese macaques (Macaca fuscata yakui) at their sleeping sites, using a highly sensitive video camera. We found evidence of nocturnal interspecific interactions, such as agonistic interactions, between Japanese macaques and sika deer (Cervus nippon yakushimae). Deer approached sleeping clusters of macaques, which slept on the ground, to eat their feces or unidentified materials near the sleeping clusters, and as a result, the macaques were often quickly displaced from their sleeping site. There was a significant difference in the occurrence of macaque–deer agonistic interactions between seasons. Our results suggested that the size of the sleeping cluster, the number of adult macaques in the cluster, and the existence of adult males in the cluster did not influence the occurrence of the agonistic interactions. Finally, we discuss the influence of this interaction on macaques and speculate on the influential factors leading to nocturnal coprophagy of macaques’ feces by deer.     

A Multi-Forest Comparison of Dietary Preferences and Seed Dispersal by <Emphasis Type="BoldItalic">Ateles</Emphasis> spp

Investigations of coevolutionary relationships between plants and the animals that disperse their seeds suggest that disperser-plant interactions are likely shaped by diffuse, rather than species-to-species, coevolution. We studied the role of dietary plasticity in shaping the potential for diffuse coevolution by comparing dietary fruit preferences and seed dispersal by 3 species of spider monkeys (Ateles spp.) in 4 moist forests in Colombia, Ecuador, Panama, and Surinam. In all forests, spider monkeys were highly frugivorous and preyed upon seeds of few species. We estimated dietary use of fruiting taxa based on absolute consumption and preference, which accounts for resource availability. Of the 59 genera that comprised the 20 most frequently consumed genera summed in each forest, only 3—Brosimum (Moraceae), Cecropia (Cecropiaceae) and Virola (Myristicaceae)—ranked within the top 20 at every forest. Most genera were within the 20 most frequently consumed at only 1 or 2 forests. Based on preferences, only 4 genera ranked in the 20 most-preferred in all 4 forests: Brosimum, Cecropia, Ficus (Moracae), and Virola. Patterns in fruit consumption and preference at the familial level were similar in that only 2 families—Myristicaceae and Moraceae—were in the 10 most-consumed or most-preferred in all 4 forests. Interforest variation in plant specific composition and abundances and supra-annual fruiting phenologies, combined with dietary flexibility of Ateles spp., may partly explain these patterns. Our results suggest that variation in plant community structure strongly influences dietary preferences, and hence, seed dispersal by spider monkeys. Thus, diffuse coevolution in spider monkey-plant relationships may be limited to few taxa at the generic and familial levels.     

Survey of gorillas (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>) and chimpanzees (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>) in Southwestern Cameroon

A study on the distribution and population density of the western lowland gorilla (Gorilla g. gorilla) and the central chimpanzee (Pan t. troglodytes) was undertaken between December 1997 and August 2000 in the Campo and Maan Forests in southwestern Cameroon. The aim of this survey was to estimate the densities of the apes in different parts of the area, to assess the importance of the region for the conservation of these endangered species and to determine the influence of human activities such as logging and hunting. The survey was based on night nest counts on a total of 665.5 km of line transects. The overall density in the Campo Forest was estimated at 0.2 gorillas/km² and at 0.63–0.78 chimpanzees/km². The overall density of chimpanzees in the Maan Forest was estimated at 0.8–1 individuals/km². Gorilla density in this area was too low to allow an estimation. The highest gorilla nest density was found in secondary forest. The gorilla density in unlogged forest was significantly lower. Chimpanzees showed a clear preference for less disturbed areas. In unlogged forest, old secondary forests (logging more than 23 years ago) and areas of recent logging with large remaining patches of primary forest, significantly higher densities were calculated than inside the more heavily exploited logging concession. In areas with both logging and high hunting pressure both species were rare or even absent. The Campo Maan area is considered a very important area for the conservation of gorillas and chimpanzees. Conservation measures are urgently required to reduce the impact of logging and hunting. The creation of the Campo Maan National Park in January 2000 is an important measure to preserve the unique biodiversity in this so far hardly protected area.     

Ontogenetic shift in dietary preference and low dietary overlap in rohu (<Emphasis Type="Italic">Labeo rohita</Emphasis>) and common carp (<Emphasis Type="Italic">Cyprinus carpio</Emphasis>) in semi-intensive polyculture ponds

In order to investigate ontogenetic changes in diet and diet overlap between rohu (Labeo rohita) and common carp (Cyprinus carpio) in polyculture ponds, food preferences of different size classes of these fishes were quantified. Rohu diet consisted of both phytoplankton and zooplankton, and there was a distinct ontogenetic shift in the relative importance of these food items. Zooplankton was the dominant food for rohu up to 20.6 cm total length (TL) and then gradually decreased in importance as fish grew. Phytoplankton was initially a minor component of rohu diet but gradually increased in importance and became the dominant food for rohu at 24.2 cm TL. Phytoplankton biovolume in rohu guts was positively correlated with fish size (TL). Chesson’s α indicated that rohu of all sizes preferentially selected Cladocera and avoided Cyanophyceae and Euglenophyceae. Young rohu initially preferred Rotifera and Copepoda but gradually switched to Bacillariophyceae and Chlorophyceae. Common carp diet consisted of phytoplankton, zooplankton, and benthic macroinvertebrates, but was dominated by benthic macroinvertebrates (63–92% of total diet). As common carp grew, the proportion of zooplankton ingested decreased and the proportion of benthic macroinvertebrates increased. Benthic macroinvertebrate biovolume in common carp guts was positively correlated with fish size. Common carp of up to 15.4 cm TL preferentially selected zooplankton, but common carp larger than 18.9 cm TL avoided this food item. Common carp of all sizes avoided phytoplankton. A low dietary overlap was found between rohu and common carp (Schoener overlap index: 0.08–0.35), probably due to ingestion of smaller quantities of zooplankton by the latter. Dietary overlap also decreased with increasing rohu and common carp size because of divergent ontogenetic shifts in dietary preferences of the two species.     

Behavioral Ecology of Sympatric Chimpanzees and Gorillas in Bwindi Impenetrable National Park,Uganda: Diet

Via a field study of chimpanzees (Pan troglodytes schweinfurthii) and gorillas (Gorilla gorilla beringei) in Bwindi Impenetrable National Park, Uganda, we found that their diets are seasonally similar, but diverge during lean seasons. Bwindi chimpanzees fed heavily on fruits of Ficus sp., which were largely ignored by the gorillas. Bwindi gorilla diet was overall more folivorous than chimpanzee diet, but was markedly more frugivorous than that of gorillas in the nearby Virunga Volcanoes. During 4 mo of the year Bwindi gorilla diet included more food species than that of the chimpanzees. Three factors in particular—seasonal consumption of fibrous foods by gorillas, interspecific differences in preferred fruit species, and meat consumption by chimpanzees—contributed to dietary divergence between the two species. When feeding on fruits, gorillas ate Myrianthus holstii more frequently than chimpanzees did, while chimpanzees included more figs in their annual diet. Chimpanzee diet included meat of duikers and monkeys; gorilla frequently consumed decaying wood.     

Species specificity in settling-plant selection by foundress ant queens in <Emphasis Type="Italic">Macaranga</Emphasis>–<Emphasis Type="Italic">Crematogaster</Emphasis> myrmecophytism in a Bornean dipterocarp forest

Previous studies have demonstrated that the obligate myrmecophytism between Macaranga ant-plants and Crematogaster plant-ants is highly species specific, although multiple Macaranga species can coexist in a microhabitat. However, the species specificity has been described based on the study of trees with established plant-ant colonies. We studied how the process of settling into the partner Macaranga seedlings by single foundress Crematogaster queens contributes to species specificity. By sampling seedlings of three sympatric Macaranga myrmecophytes species in the field, we tested two hypotheses. The first is that foundresses correctly select their specific partner plant species when they settle into seedlings. The second hypothesis is that the seasons in which seedlings available for settling by foundresses appear are segregated among the Macaranga species, and the seasons in which foundress queens settle are synchronized to the appearance of seedlings of specific partner species; thus species specificity is consequently generated. Our results support the former hypothesis but not the latter: we always observed foundresses settling species-specific host plants, and seedlings suitable for settling were always available in each Macaranga species. Electronic Publication     

Determinants of fruit removal in <Emphasis Type="Italic">Geonoma pauciflora</Emphasis>, an understory palm of neotropical forests

Competition for seed dispersers is supposedly a selective force that drives the evolution of plant reproductive traits. In the understory of tropical forests, such competition should be especially severe among bird-dispersed plant species because (i) the production of copious fruit crops is limited by low light availability; (ii) there usually is a high density of fleshy fruited plants, and (iii) understory frugivorous birds are not abundant. In this paper, we took advantage of a high-density population of Geonoma pauciflora, a bird-dispersed palm species growing in the understory of the Brazilian Atlantic forest, to investigate the influence of plant traits and its immediate neighborhood on fruit removal. Intrinsic (crop and fruit sizes) and extrinsic traits (related to light availability) affected the fruit removal of G. pauciflora. Crop size had a greater influence than fruit size on the absolute number of fruits removed, whereas none of the investigated traits influenced decisively fruit removal efficiency (i.e., the proportion of an individual’s crop removed). The influence of light availability was mostly indirect, through its positive influence on fruit production. A significant positive spatial autocorrelation in removal efficiency occurred among neighboring plants within a 7-m radius, which is indicative of facilitation among neighboring individuals. The consequence of such positive spatial autocorrelation in removal efficiency for clonal plants such as G. pauciflora is that by attracting a frugivore a given ramet may promote the removal of fruits of other ramets, thus enhancing the reproductive output of the genet as a whole.     

Crop size is more important than neighborhood fruit availability for fruit removal of <Emphasis Type="Italic">Eugenia uniflora</Emphasis> (Myrtaceae) by bird seed dispersers

For a plant with bird-dispersed seeds, the effectiveness of seed dispersal can change with fruit availability at scales ranging from individual plants to neighborhoods, and the scale at which frugivory patterns emerge may be specific for frugivorous species differing in their life-history and behavior. The authors explore the influence of multispecies fruit availability at two local spatial scales on fruit consumption of Eugenia uniflora trees for two functional groups of birds. The authors related visitation and fruit removal by fruit gulpers and pulp mashers to crop size and conspecific and heterospecific fruit abundance to assess the potential roles that facilitative or competitive interactions play on seed dispersal. The same fruiting scenario influenced fruit gulpers (legitimate seed dispersers) and pulp mashers (inefficient dispersers) in different ways. Visits and fruit removal by legitimate seed dispersers were positively related to crop size and slightly related to conspecific, but not to heterospecific fruit neighborhoods. Visits and fruit consumption by pulp mashers was not related to crop size and decreased with heterospecific fruit availability in neighborhoods; however, this might not result in competition for dispersers. The weak evidence for facilitative or competitive processes suggest that interaction of E. uniflora with seed dispersers may depend primarily on crop size or other plant’s attributes susceptible to selection. The results give limited support to the hypothesis that spatial patterns of fruit availability influence fruit consumption by birds, and highlight the importance of considering separately legitimate and inefficient dispersers to explain the mechanisms that lie behind spatial patterns of seed dispersal.