White-lipped peccary (Tayassu pecari,Artiodactyla: Tayassuidae) diet and fruit availability in a Costa Rican rain forest

We studied fruit availability, diet and habitat use by white-lipped peccary (Tayassu pecari) in Corcovado National Park, southwest Costa Rica, from July 1996 to April 1997. The results show that the availability of important fruits for the white-lipped peccary differs between habitats and climatic seasons. Fruit availability was highest in the primary forest than secondary and coastal forest. There was a period of shortage of fruits to ends of the wet season, during which the consumption of not seasonal resources like leaves and shafts increased. The important fruits during this period of shortage were Ficus sp and Licania operculipetala. The several types of forest were used according to the fruit availability, and it was a direct relation between the consumption and the fruit availability.     

Spermatogenesis in white-lipped peccaries (Tayassu pecari)

We describe here morphological and functional analyses of the spermatogenic process in sexually mature white-lipped peccaries. Ten sexually mature male animals, weighing approximately 39 kg were studied. Characteristics investigated included the gonadosomatic index (GSI), relative frequency of stages of the cycle of seminiferous epithelium (CSE), cell populations present in the seminiferous epithelium in stage 1 of CSE, intrinsic rate of spermatogenesis, Sertoli cell index, height of seminiferous epithelium and diameter of seminiferous tubules, volumetric proportion of components of the testicular parenchyma and length of seminiferous tubules per testis and per gram of testis. The GSI was 0.19%, relative frequencies of pre-meiotic, meiotic and post-meiotic phases were, respectively 43.6%, 13.8% and 42.6%, general rate of spermatogenesis was 25.8, each Sertoli cell supported an average 18.4 germinative cells, height of seminiferous epithelium and diameter of seminiferous tubules were, respectively, 78.4 microm and 225.6 microm, testicular parenchyma was composed by 75.8% seminiferous tubules and 24.2% intertubular tissue, and length of seminiferous tubules per gram of testis was 15.8m. These results show that, except for overall rate of spermatogenesis, the spermatogenic process in white-lipped peccaries is very similar to that of collared peccaries, and that Sertoli cells have a greater capacity to support germinative cells than most domestic mammals.     

Nutritional values in the diet of white-lipped peccary Tayassu pecari (Artiodactyla: Tayassuidae) in Corcovado National Park, Costa Rica

We determined the potential nutritional levels in 25 species of plants, and in earthworms, that constitute part of the diet of white-lipped peccary (Tayassu pecari) in Corcovado National Park, Costa Rica, from January 1998 to March 1999. The highest content of fat and energy was found in seeds of the Myristicacea family. The highest content of calcium was found in vegetative parts of Dieffenbachia spp. Nutritious contents differed among plant parts (seeds, fruits, stems and leaves). Fat and energy content were larger in seeds and fruits, whereas the largest content of protein was found in fruits and leaves. Mineral content also differed among plant parts. Calcium, potassium and magnesium were higher in leaves whereas copper and zinc were higher in seeds. Differences of diet between white-lipped peccaries in Corcovado and in other tropical regions of Latin America could be partially explained by our results. We found several species with higher fat and energy content than palms, which can explain the low consumption of palm seeds in Corcovado. It is possible that the regular consumption of stems and leaves of some species is related to their high mineral content. Seasonality of reproduction in Corcovado seems to be related not only to fruit availability but also to the nutritional quality of food.     

Diet of peccaries (Tayassu tajacu and T. pecari) in a dry forest of Venezuela

The diets of Tayassu tajacu and T. pecari were assessed by means of direct observations, stomach content, and faecal analyses. The study was conducted in a dry forest located in the central Llanos region, Venezuela, where both species coexist. Peccaries were observed during one week of every month from August 1986 to May 1987. Fruits and seeds of Pithecellobium saman and Guazuma ulmifolia and roots and stems of Cissus rhombifolia comprised most of the diet of both peccary species. Proportions of these items differed significantly between species. There was no Acacia macracantha in stomachs nor in faeces of T. tajacu , but it was one of the most important items found in stomachs of T. pecari. Leaves appeared in all stomachs but in very low proportions (<1%). Insect pupae and larvae were found in small quantities. Results suggest that peccaries behave as frugivores and seed predators as most seed remains appeared cracked. Differences in the diet and behavioural traits described elsewhere may allow these similar species to coexist in the tropical dry forest of central Venezuela.     

Kinetics of spermatogenesis in the white-lipped peccary (Tayassu pecari)

This study aimed to characterize the stages of the seminiferous epithelium cycle by the tubular morphology method, and to determine the number of differentiated spermatogonia generations in the adult white-lipped peccary. Twenty adult white-lipped peccaries, obtained from commercial slaughterhouse, were used. Fragments of the testicular parenchyma were fixed in 3% glutaraldehyde and embedded into a methacrylate resin. The number of germ and Sertoli cells was estimated by the analysis of cell populations in 50 transversal sections of seminiferous tubules in different stages of the cycle. The tubular morphology method allowed the identification of cellular associations characteristic of the eight stages of the seminiferous epithelium cycle in white-lipped peccaries. The results showed the presence of six generations of differentiated spermatogonia in white-lipped peccaries, and that the cell composition of the eight stages of the seminiferous epithelium cycle in this species is very similar to that described for collared peccaries.     

Toxoplasma gondii antibodies in wild white-lipped peccary (Tayassu pecari) from Peru

In the Peruvian Amazon, the white-lipped peccary (Tayassu pecari) is a desirable game species and is important for the local rural economy. Blood samples from 101 white-lipped peccaries from Peru were collected from 3 different conservation areas located in the municipalities of Manu and Tambopata, southeastern region of the Peruvian Amazon. Antibodies were assayed using the modified agglutination test (MAT, cut of value of 25). Antibodies to Toxoplasma gondii were found in 89.1% (90 of 101) of animals, with titers of 1 ∶ 25 in 9, 1 ∶ 50 in 25, 1 ∶ 100 in 20, 1 ∶ 200 in 14, 1 ∶ 400 in 12, 1 ∶ 800 in 9, and 1 ∶ 3,200 in 1; 87.7% and 89.2% of males and females, respectively, tested positively, and no association (P ≥ 0.05) with gender and occurrence of antibodies was observed.     

Social interactions of juvenile Collared peccaries, Tayassu tajacu (Mammalia: Artiodactyla)

Is there a predictable relationship between species-typical behavioural development and social organization in mammals? Earlier work on canids and rodents suggested that adult sociality is associated with a developmental schedule in which juveniles showed, among themselves: (1) frequent social play; (2) frequent amicable contact; (3) frequent olfactory contact; (4) a delay in the emergence of agonistic behaviour. In the highly social Collared peccary (Tayassu tajacu) the behaviour of juveniles conformed only partially to this scheme. Juveniles engaged in social play with each other at an early age, but showed no other observable amicable or olfactory contact until eight to twelve weeks of age and showed no delay in the emergence of agonistic behaviour among themselves. Adults, in contrast, were conspicuously tolerant of juveniles, and 86% of juvenile social interactions in the first year took place with adults.     

Removal of palm fruits and ecosystem engineering in palm stands by white-lipped peccaries (Tayassu pecari) and other frugivores in an isolated Atlantic Forest fragment

Long-term studies in a 2,178 ha fragment of semideciduous Atlantic Forest demonstrated important interactions between white-lipped peccaries (Tayassu pecari) and the common palms, Syagrus romanzoffiana and Euterpe edulis. We conducted fruit removal and medium-to-large-sized mammalian exclusion experiments to: (1) quantify seasonal fruit consumption from high-density patches beneath parent trees by T. pecari and other consumers, and (2) measure impacts of T. pecari rooting and foraging activities on seedling dynamics in E. edulis stands. A diverse array of fauna consumed S. romanzoffiana fruits. During the dry season, when S. romanzoffiana palms provided 68% of fruit dry weight in the fragment, T. pecari consumed significantly greater amounts than other consumers, and along with Pecari tajacu and Tapirus terrestris, were potential seed dispersers. The rodents, Sciurus ingrami and Agouti paca, consumed most S. romanzoffiana fruits in the wet season, acting as both seed dispersers and predators. More than 95% of E. edulis fruit removal was due to seed predation by T. pecari. Intense removal during the dry season was closely linked with previously documented range shifts and habitat preferences by T. pecari. Exclusion plot experiments in E. edulis (palmito) stands showed that the number and proportion of nonpalmito (not E. edulis) seedlings increased dramatically in the absence of T. pecari rooting and foraging activities that disturbed soil and thinned seedlings. We discuss the importance of these ecosystem engineering activities and palm-peccary trophic interactions for long-term maintenance of E. edulis stands and T. pecari populations, as well as water balance, in the forest fragment.     

Removal of seeds from frugivore defecations by ants in a Costa Rican rain forest

At our Costa Rican field site, seeds defecated by frugivorous birds usually do not remain where they have been deposited. Many species of ants are attracted to frugivore defecations and remove seeds and/or pulp. Pheidole species selectively remove seeds, fungus-growing species (tribe Attini) remove both pulp and seeds. Seeds of many Melastomataceae have an appendage, which we hypothesized is an elaiosome. Indeed, preference trials demonstrated that two species of Pheidole selected seeds with the appendage over seeds of the same species in which the appendage had been removed. However, we found that these ants did not take the appendage when it was offered by itself. We conclude that the appendage is not an elaiosome. In further trials, different ant species preferentially selected different seed species. These ants consumed some seeds and deposited others unharmed in refuse piles. We conclude that because the composition of leaf-litter ant communities is highly variable between neighboring square meter plots, and the probability of seed predation depends upon the species of ant, the over-all effect of ants on seed shadows and seed banks is spatially unpredictable. Addendum: The names of the two Pheidole emphasized in this study. P. nebulosa and P. nigricula, are unpublished names from a generic revision being prepared by E. O. Wilson and W. L. Brown. Their use here is not intended to constitute taxonomic publication but is solely for more precise indentification in future ecological research of similar nature     

Root surface acid phosphatase activities of vascular epiphytes of a Costa Rican rain forest

The objective of this work was to compare root surface phosphatase activities of vascular epiphytes typical of a lowland tropical forest. Acid phophatase, measured at pH 5.0 with the substrate p-nitrophenyl phosphate, was detected in 22 species distributed within 10 plant families. Epiphytes were classified as trunk, canopy-mat or bare-limb species based upon their usual occurrence. Phosphatase activity was not significantly correlated with plant occurrence. However, phosphatase activity was generally highest in trunk occurring and canopy-mat epiphytes rooted in mosses and humus-like accumulations, and lowest in species restricted to bare limbs. Epiphyte shoot phosphorus and chlorophyll content were correlated with species occurrence, with phosphatase being positively correlated with plant P content. The observed changes in acid phosphatase production among habitats were consistent with predicted changes in the availability of organic P sources. However, observed changes also reflect accompanying shifts in root structure including: the occurrence of velamen or waxy layers, changes in root diameter, branching and root hair density.     

Variation in leaf litter nutrients of a Costa Rican rain forest is related to precipitation

By assessing current leaf litter nutrient dynamics, we may be able to predict responses of nutrient cycling in tropical ecosystems to future environmental change. The goal of this study was to assess whether nutrient cycling is related to seasonal variation in rainfall in a wet tropical forest. We examined leaf litter of an old-growth tropical rain forest in N.E. Costa Rica over a 4-year period to explore seasonal and inter-annual changes in leaf litter nutrient concentrations, and to evaluate potential short- and long-term drivers of variation in litter nutrient concentration, particularly that of phosphorus (P) and nitrogen (N). We also examined the temporal dynamics of calcium, potassium, and magnesium in the leaf litter. Leaf litter [P] and %N changed significantly with time, both seasonally and inter-annually. Seasonal changes in leaf litter [P] were strongly positively correlated with rainfall from the previous 2 weeks; cations, however, were inversely related to this measure of current rainfall, while %N was not related to rainfall. We propose that the positive relationship between current rainfall and leaf litter [P] is due to a response by the vegetation to an increase in nutrient availability and uptake. In contrast, given the negative relationship between current rainfall and cation concentrations, leaching from live leaf tissue is a more likely driver of short-term changes in cations. Should global climate change include altered rainfall patterns in this biome, one class of ecosystem-level responses could be significant changes in P and cation cycling.     

Microclimatic parameters influencing nitrogen fixation in the phyllosphere in a Costa Rican premontane rain forest

The acetylene reduction method was used to measure nitrogen fixation in the phyllosphere of attached leaves of different phorophytes under natural conditions in a premontane rain forest in Costa Rica. Maximum rates of nitrogen fixation (26 ng N · cm⁻² leaf area · h⁻¹) – mainly due to the activity of two species of Scytonema (Cyanobacteria) – were measured in the rainy season in bright sunlight. Rates of nitrogen fixation were correlated with the leaf area covered by Scytonema. In periods without precipitation the fixation activity decreased to zero within 2–3 days. As long as the epiphylls were sufficiently supplied with water, other microclimatic factors like temperature and light intensity also influenced nitrogen fixation rates, but to a lesser extent. Relative humidity and species of phorophyte showed no direct influence. It was concluded that the most important factor for nitrogen fixation in the phyllosphere was the availability of liquid water. Linking these results to meteorological data, the input of nitrogen by biological nitrogen fixation in the phyllosphere in the investigation area was estimated to be as much as 1.6 ± 0.8 kg N · ha⁻¹ · year⁻¹ per unit of leaf area index (LAI). For an LAI of 2 for the understory the nitrogen input would vary between 2 and 5 kg N · ha⁻¹ · year⁻¹. This work also demonstrates the importance of detailed knowledge of variation in microclimate throughout the year as a basis for extrapolation of the annual nitrogen input. Received: 21 December 1997 / Accepted: 14 June 1998     

Rodent pollination of Blakea (Melastomataceae) in a Costa Rican cloud forest

Several species of rodents have been observed visiting the flowers of an undescribed species ofBlakea (Melastomataceae) in a cloud forest in Costa Rica. The rodents drink the copious nectar secreted at the base of the stamens and are dusted with pollen. The first report of pollination by non-flying mammals in the neotropics suggests that this type of pollination may be important where there is little competition with bats.     

Contrasting patterns of vertical stratification in two moth families in a Costa Rican lowland rain forest

The vertical stratification of two species-rich moth families (Lepidoptera: Arctiidae, Geometridae) was analysed in a lowland rain forest in Costa Rica. Moths were trapped with automatic 8 W ultraviolet light traps at three sites in the understorey and canopy of a primary forest at the La Selva Biological Station (10.4° N, 84.0° W). A total of 846 arctiid moths (148 species) and 946 geometrid moths (140 species) were analysed. Species richness and diversity of arctiid moths was significantly lower in the understorey than in the canopy (Fisher's alpha: 24 vs. 49) whereas geometrid moths showed an inverse pattern (Fisher's alpha: 44 vs. 27). This resulted in an overall increase in the proportion of conspicuously coloured species towards the canopy. Moth ensembles were clearly separated in multidimensional scaling ordinations, and differed significantly in their faunal composition and dominance between the strata. The available host plant data suggest that the flight height of moths was determined by larval resource availability. Examples include understorey flyers such as the geometrid genus Eois feeding on Piper, and canopy flyers such as the arctiid moth genera Aclytia, Macrocneme and Poliopastea which feed on lianas.     

Evaluation of genetic variability in the collared peccary Pecari tajacu and the white-lipped peccary Tayassu pecari by microsatellite markers

In this study, the microsatellite technique was used to evaluate the genetic variability in populations of collared and white-lipped peccaries kept in captivity. Six primers developed for domestic pigs were used and amplified in both species. They revealed the presence of five polymorphic loci and one monomorphic locus. The polymorphic loci included 4 of the 16 alleles in collared peccaries, and 3 of the 10 alleles in the white-lipped peccaries. Polymorphic information content (PIC) in both species and all the loci was highly informative. The probability of paternity exclusion (PEC), if one of the parents is known, was almost as high in white-lipped peccaries (95.53%) as in the collared (99,48%). The Fst values for collared (0.042) and white-lipped (0.1387) peccaries showed that both populations are not structured. The Fis values for all loci, except ACTG2 in white-lipped peccaries (-0.0275) and in both species (0.1985 to 0.9284 in collared peccaries and 0.3621 to 0.4754 in the white-lipped), revealed a high level of homozygosis, probably caused by inbreeding. Data on heterologous amplification and genetic variability in collared and white-lipped peccaries are presented for the first time.     

Influence of land crabs Gecarcinus quadratus (Gecarcinidae) on distributions of organic carbon and roots in a Costa Rican rain forest

In Costa Rica's Corcovado National Park, the fossorial land crab, Gecarcinus quadratus (Gecarcinidae), densely populates (1 - 6 m(-2)) a region of forest extending from the Park's Pacific coastline inland to ca. 600 m. Throughout this coastal forest ('crabzone'), crabs selectively forage for fallen leaves and relocate them to subterranean burrow chambers. Comparisons between surface soils (0 - 15 cm) sampled from the crabzone and forest lying immediately inland that is naturally devoid of crabs ('crabless zone') suggest that crabzone top soils contained less organic carbon and fewer fine and very fine roots. In contrast, soils sampled from 70 - 100 cm depths in the crabzone contained twice the carbon of the crabless zone during the dry season but similar values during the wet season. Two years of experimental crab exclusion from 25 m2 replicates established in the crabzone resulted in 16% more organic carbon content in surface soils relative to baseline conditions (n.s.) and 22% more carbon than final control values (P < 0.05). Excavations of burrow chambers, with an average (+/- SD) depth of 48 +/- 12 cm, indicated localized, subterranean pockets of elevated (+ 60 %) organic carbon and increased densities of fine and very fine roots relative to same-depth samples from the crabzone unassociated with burrows chambers.     

No short-term change in soil properties following four-fold litter addition in a Costa Rican rain forest

We experimentally manipulated forest floor litter to investigate the influence of litter quality and quantity on soil properties over the short-term (weeks to months) in a wet tropical forest in NE Costa Rica. The study included old growth forest on high fertility soils, old growth forest on low fertility soils, and secondary forest on intermediate fertility soils. Forest floor litter was removed from a 16 m² area and added to an adjacent 4 m² area in March 2003, resulting in a one to four-fold increase in the annual litter input to the forest floor. We created three addition, three removal and three control plots per forest type. We measured treatment effects on variation in soil moisture, temperature, pH, and Bray-1 P (plant available) over a 5-month period that captured the dry-wet season transition. Litter manipulation had no effect on any of the soil properties measured during the 5-month study period. Significant variability through time and a similar temporal pattern across the three forest stands suggest that climatic variability is driving short-term patterns in these soil properties rather than seasonal inputs of litter. In general, soils were warmer, drier and more basic with higher available P during dry season months. Even in wet tropical forests, small variability in climate can play an important role in soil dynamics over periods of weeks to months. Although litter manipulation did not influence soil properties over the 5-month study period, a longer lag may exist between the timing of litter inputs and the influence of that litter on soil properties, especially plant available P.     

Ovarian folliculogenesis in collared peccary, Pecari tajacu (Artiodactyla: Tayassuidae)

The sustainability and production of collared peccary (Pecari tajacu) has been studied in the last few years; however, further information on its reproduction is necessary for breeding systems success. Understanding folliculogenesis aspects will contribute to effective reproductive biotechniques, which are useful in the preservation and production of wildlife. The aim of this study was-to evaluate the ovarian folliculogenesis in collared peccary. Ovaries from six adult females of collared peccary were obtained through ovariectomy and analyzed. These were fixed in aqueous Bouin's solution and sectioned into 7 microm slices, stained with hematoxilin-eosin and analyzed by light microscopy. The number of pre-antral and antral follicles per ovary was estimated using the Fractionator Method. The follicles, oocytes and oocyte nuclei were measured using an ocular micrometer. Results showed that the length, width, thickness, weight, and the gross anatomy of the right and left ovaries were not significantly different. However, the mean number of corpora lutea was different between the phases of the estrous cycle (p<0.05), with the highest mean in the luteal phase. Primordial follicles were found in the cortex; the oocytes were enveloped by a single layer of flattened follicular cells. In the primary follicles, proliferation of the follicular cells gave rise to cuboidal cells (granulosa cells). The secondary follicle was characterized by two or more concentric layers of cuboidal cells (granulosa), beginning of antrum formation, and the presence of pellucid zone and theca cells. Antral follicles were characterized by a central cavity (antrum), the presence of cumulus oophorus and theca layers (interna and externa). In the right ovary, the values of the primordial and primary follicles were similar, but significantly different from the secondary ones (p<0.05). In the left ovary, significant differences were observed between all follicles in the follicular phase (p<0.05); the mean number of primordial and primary follicles was similar in the luteal phase. The mean number of pre-antral follicles and antral follicles in the follicular phase was higher in the left ovary (p<0.05). The mean number of antral follicles in the luteal phase was similar in both ovaries. We also found significant differences in mean diameter of preantral follicles, oocyte, granulosa layer and oocyte nucleus during the estrous cycle. In the antral follicles a significant difference was observed only in follicular diameter (p<0.05). The predominance of active primordial and primary follicles was found in both phases; otherwise the secondary follicles and antral follicles showed a high degree of degeneration. The results obtained in the present work will strengthen the development of biotechnology programs to improve the productive potential and conservation of the collared peccary.     

Phylogenetic Systematics of Peccaries (Tayassuidae: Artiodactyla) and a Classification of South American Tayassuids

Tayassuidae is a family of pig-like Artiodactyla restricted to the New World. Despite its rich fossil history, they have received less attention from a taxonomic and phylogenetic perspective when compared to the Old World pigs, Suidae. In this study, we performed a computer assisted phylogenetic analysis using morphological and molecular data including fossil and extant Tayassuidae, using parsimony and Bayesian approaches. We recovered the monophyly of the family Tayassuidae, confirming previous proposals, as well as the monophyly of the subfamilies Hesperhyinae and Tayassuinae, and the genus Platygonus, which we placed in a new taxon of tribe level. The three living peccaries and a number of fossil species belong to a new, tribe level, monophyletic group. The genus Catagonus comes out as paraphyletic, leading us to propose to restrict the generic name to the type species, C. metropolitanus, and a new taxonomic arrangement for the remaining species previously included in it, revalidating the genera Brasiliochoerus and Parachoerus, and describing a new genus, Protherohyus, gen. nov.