Population density and use of space in howling monkeys (Alouatta villosa) in southwestern Panama

Eight troops ofAlouatta villosa (= A. palliata) were studied for 10 weeks in a natural forest in southwestern Panama. This is one of the few extended studies on the species to be conducted in a location other than Barro Colorado Island. The 151 animals belonging to 8 troops and 6 additional isolates utilized 37 acres of forest. This population density was 21 times greater than on Barro Colorado in 1932 and 12 times than on Barro Colorado in 1967. The average troop size was 18.9 animals; the average home range was 12.1 acres. Home ranges overlapped extensively and on the average troops had only 6% of their home range for exclusive use. There were no signs that the howling monkeys were exhausting their food supply or were experiencing decreased fertility. Several factors that could function in population control are evaluated.     

An example of “apparent” dominance-submission behavior between adult male howler monkeys (Alouatta palliata)

On Barro Colorado Island, Panama, two adult male howler monkeys (Alouatta palliata) belonging to the same troop were observed performing a sequence of behaviors typical of dominance-submission situations. The occurrence of these behaviors supports the hypothesis of an agegraded-male social organization for this species. There may also be a correlation between the appearance of these newly-reported behaviors and high population densities.     

Ocelots on Barro Colorado Island are infected with feline immunodeficiency virus but not other common feline and canine viruses

Transmission of pathogens from domestic animals to wildlife populations (spill-over) has precipitated local wildlife extinctions in multiple geographic locations. Identifying such events before they cause population declines requires differentiating spillover from endemic disease, a challenge complicated by a lack of baseline data from wildlife populations that are isolated from domestic animals. We tested sera collected from 12 ocelots (Leopardus pardalis) native to Barro Colorado Island, Panama, which is free of domestic animals, for antibodies to feline herpes virus, feline calicivirus, feline corona virus, feline panleukopenia virus, canine distemper virus, and feline immunodeficiency virus (FIV), typically a species-specific infection. Samples also were tested for feline leukemia virus antigens. Positive tests results were only observed for FIV; 50% of the ocelots were positive. We hypothesize that isolation of this population has prevented introduction of pathogens typically attributed to contact with domestic animals. The high density of ocelots on Barro Colorado Island may contribute to a high prevalence of FIV infection, as would be expected with increased contact rates among conspecifics in a geographically restricted population.     

Copulation-interrupting behavior between females within a howler monkey troop

On Barro Colorado Island, Panama, howler monkeys (Alouatta palliata) were observed copulating on numerous occasions. In one case, however, the copulation was interrupted by a female member of the same troop. This newly reported behavior may be a social mechanism related to high population densities. Supported by the Environmental Sciences Program, Smithsonian Institution; and the National Zoological Park, Washington, D.C.     

Isolation and characterization of microsatellite loci in the tropical tree Jacaranda copaia (Bignoniaceae)

We isolated 10 microsatellite loci for Jacaranda copaia for direct measurement of seed and pollen movement within the 50 ha Forest Dynamics Plot on Barro Colorado Island, Panama, and for studies of population genetic structure and gene flow across the Isthmus of Panama. Enriched repeat libraries were screened for microsatellite repeats and polymorphic primer pairs were developed. A total of 110 adult individuals were screened for allelic diversity and loci showed high levels of heterozygosity. These loci show both high maternal and paternal exclusion probabilities.     

Thief workers and variation in nestmate recognition behavior in a ponerine ant, Ectatomma ruidum

Our findings give new insight into the relationship between nestmate recognition and cleptobiosis, intraspecific thievery of newly collected food items, in a neotropical ponerine ant, Ectatomma ruidum. The expression of discrimination of nestmates from non-nestmates varied among local aggregations of E. ruidum in a population at Barro Colorado Island, Panama. This result is due to differences in the behavior of the guard ants among aggregations, rather than differences in the expression of recognition cues by ants. Baiting experiments show that E. ruidum colonies in Panama have a greater tendency to have overlapping home ranges than a similar population in Costa Rica. The pattern of cleptobiosis, however, is strikingly similar between the Panamanian and Costa Rican populations.     

Effect of trail pheromones and weather on the moving behaviour of the army ant <Emphasis Type="Italic">Eciton burchellii</Emphasis>

Most of what we know about the moving behaviour of the nomadic army ant Eciton burchellii comes from Barro Colorado Island (BCI) in Panama. Nomadic colonies raid roughly in straight line during the day and relocate their nests along this path in the evening. At BCI, nomadic colonies raid roughly in the same compass bearing of the previous day, presumably using their pheromone-marked raiding trails as cues to pick directions. Deviations from this direction occur when a nomadic colony fails to move, possibly due to environmental conditions. The generality of these results has been questioned. We studied nomadic colonies of E. burchellii at La Selva Biological Station, Costa Rica to evaluate the generality of the results obtained from BCI. We measured the angle between consecutive raids, manipulated the distribution of previous day’s raid pheromones around nests to evaluate the effect of raid pheromone on foraging direction, and evaluated the effect of rainfall on the probability of moving and on deviation from the previous day’s raid. Colonies did not follow the same compass bearing of the previous day and formed new raids on areas with previous day’s raid pheromones or without them. Rainfall can explain when nomadic colonies move, but did not explain deviation from the previous day’s raid direction. Our results suggest that caution should be taken when generalizing the insightful results obtained from the BCI population.     

Chasing behavior between males within a howler monkey troop

On Barro Colorado Island, Panama, a mature male howler monkey (Alouatta palliata) was observed chasing a young adult male howler, both members of the same troop. Detailed observations on births, deaths and solitary males before and after the chase suggest a strong correlation between an unstable social structure within the troop and overt aggression between different-aged males. Support for this field work was received from the Environmental Sciences Program, Smithsonian Institution; and the National Zoological Park, Washington, D.C.     

Eight microsatellite markers for the neotropical tree Luehea seemannii (Tiliaceae)

We isolated eight polymorphic microsatellites from the neotropical tree Luehea seemannii for gene flow and genetic structure studies. We used a streptavidin subtractive enrichment technique to develop a library of CA/GT repeats. Eight loci were screened for diversity from 96 individuals from Barro Colorado Island (BCI) and neighbouring Gigante peninsula in Panama. Luehea seemannii shows moderate levels of genetic diversity within these two populations. Allelic richness ranged from four to nine alleles and averaged 6.44 alleles per locus. Average expected heterozygosity was 0.65 on BCI and 0.60 on Gigante. Results are compared to microsatellite data from another wind‐dispersed gap colonizing species common in Panama.     

Negative density dependence of seed dispersal and seedling recruitment in a Neotropical palm

Negative density dependence (NDD) of recruitment is pervasive in tropical tree species. We tested the hypotheses that seed dispersal is NDD, due to intraspecific competition for dispersers, and that this contributes to NDD of recruitment. We compared dispersal in the palm Attalea butyracea across a wide range of population density on Barro Colorado Island in Panama and assessed its consequences for seed distributions. We found that frugivore visitation, seed removal and dispersal distance all declined with population density of A. butyracea, demonstrating NDD of seed dispersal due to competition for dispersers. Furthermore, as population density increased, the distances of seeds from the nearest adult decreased, conspecific seed crowding increased and seedling recruitment success decreased, all patterns expected under poorer dispersal. Unexpectedly, however, our analyses showed that NDD of dispersal did not contribute substantially to these changes in the quality of the seed distribution; patterns with population density were dominated by effects due solely to increasing adult and seed density.     

A mathematical model for the population dynamics of army ants

A stochastic cellular automata model for the population dynamics of the army antEciton burchelli on Barro Colorado Island in Panama is set up. It is simulated on the computer and shown to give good agreement with biological data. It is analysed using two approximations akin to the mean field approximation in statistical mechanics, and good agreement with the simulations is obtained. Finally, the role of distance between successive statary phase bivouacs is discussed with regard to the rate of colony growth. There are two aspects of the biological system studied here that make it of general importance. First, the population is structured, since the size of each colony of army ants is crucial. Second, the spatial behaviour of the population, as in many others, is not diffusion-like, although it is random. This has implications for the kind of model that is chosen.     

Population- and sociogenetic structure of the leaf-cutter ant <Emphasis Type="Italic">Atta colombica</Emphasis> (Formicidae,Myrmicinae)

Relatedness and genetic variability in colonies of social insects are strongly influenced by the number of queens present and the number of matings per queen, but also by the genetic variability in the population. Thus, multiple paternity will enhance within-colony genetic variability more strongly when the males a queen mates with are unrelated. To study the kin-structure within colonies of the leaf-cutter ant Atta colombica and the population structure of this species around Barro Colorado Island, Panama, we developed five polymorphic microsatellite loci with a range of three to 17 alleles in At. colombica, all of which cross-amplify in other higher attines as well. The average effective mating frequency calculated from four-locus microsatellite genotypes was 1.89 ± 0.12 (harmonic mean ± SE) and thus slightly lower than the average observed mating frequency of 2.50 ± 0.11 (arithmetic mean ± SE) over the 55 colonies studied, confirming former studies that utilized fewer loci. The discrepancy between observed mating frequency and effective mating frequency is most probably due to paternity skew within colonies. The study population proved to be genetically diverse and in Hardy-Weinberg equilibrium, suggesting random mating within the study area. No population substructure was observed, neither considering nuclear (global F _ST = 0.011 ± 0.003 SE) nor mitochondrial markers (mean Φ_ST = 0.008). Consequently, gene flow is obviously promoted by both sexes across the range investigated here. Thus, multiple mating and long-distance dispersal appear to be two interconnected behavioural mechanisms to create and maintain genetic diversity in At. colombica. The advantages of this system are partly offset by paternity skew and the non-zero relatedness among colony fathers found in the study population. Received 18 March 2008; revised 14 July 2008; accepted 18 July 2008.     

Seasonal and age-related differential mortality by sex in the Mantled Howler monkey,Alouatta palliata

Age-specific, differential mortality occurs between male and female mantled howler monkeys, Alouatta palliata,on Barro Colorado Island, Panama. A peak in mortality between 5 and 7 years is seen in males but not infernales. Dates when howler cadavers have been found suggest a seasonal mortality, with peaks in July and perhaps also in December through January.     

Tree-rubbing behavior of a solitary male howler monkey

On Barro Colorado Island, Panama, a solitary male howler monkey (Alouatta palliata) was observed copulating with a female member of a troop. A unique tree-rubbing behavior was then performed by the male. Within two weeks he was the dominant male in that troop.     

Herbivory Rate of Leaf-Cutting Ants in a Tropical Moist Forest in Panama at the Population and Ecosystem Scales

Leaf-cutting ants are frequently characterized as the major herbivores in the Neotropics, but quantitative data to back up this assumption are scarce. In this study, the consumption and herbivory rates for the entire leaf-cutting ant ( Atta colombica , Formicidae) population in an old secondary forest on Barro Colorado Island (BCI) in Panama were determined over 15 mo (on average 49 colonies). The number of harvested leaf fragments was calculated from monthly refuse deposition rates of the colonies and the regression between refuse deposition and harvesting rates. The inclusion of fragment characteristics (proportion of leaf fragments in the harvest, average fragment weight, and area) allowed us to calculate consumption and herbivory rates at colony, population, and ecosystem levels. The A. colombica population harvested 13.2 tons of biomass/yr and 13.1 ha of leaf area/yr, and deposited 9.4 tons of refuse material/yr. Rates varied considerably among colonies. At the ecosystem level, i.e. , per forest area, herbivory rates were 132 kg biomass/ha/yr and 1310 m² foliage/ha/yr. For the area on BCI where A. colombica occurs (100 ha), this is equivalent to 2.1 percent of the foliage area in the forest or 1.7 percent of the annual leaf-area production. This value is considerably lower than previously published estimates of leaf-cutting ant herbivory rates in tropical forests.     

Impact of Atta colombica Colonies on Understory Vegetation and Light Availability in a Neotropical Forest1

We quantified patterns of vegetation removal and light availability above Atta colombica nests on Barro Colorado Island, Panama. Ants cleared vegetation less than 1 cm in diameter from an area of 77 m², and up to 3 m above ground level. Overall light availability 1.5 m above ground level was 49 percent greater at ant nest sites than at sites in undisturbed understory. These higher light levels fell within the range known to enhance growth of both shade tolerant and pioneer species.     

Predation-mediated Mortality of Early Life Stages: A Field Experiment with Nymphs of an Herbivorous Stick Insect (Metriophasma diocles)

We quantified predation pressure on first instar nymphs of a stick insect in predator‐exclusion experiments in the forest of Barro Colorado Island, Panamá. After considering intrinsic mortality (19%) and potential emigration (negligible), we estimated that 54 percent of the nymphs died due to predation in a two‐week period. Predation on nymphs was highest at night and may explain the low abundance of Metriophasma diodes in the understory.     

Density and distance-to-adult effects of a canker disease of trees in a moist tropical forest

We compared the spatial distribution of stem cankers on the canopy tree Ocotea whitei (Lauraceae) in a 20-ha plot on Barro Colorado Island, Panama, with spatial and temporal patterns of mortality in this host over the previous decade. The cankers occur both on adult and juvenile individuals, aothough juveniles are much more likely the adults to show symptoms. Disease incidence is host-density dependent, and both the presence of the disease and host mortality are more likely close to than far from a conspecific adult, which resulted in a net spatial shift of the juvenile population away from conspecific adults through time. Disease incidence is lower than expected among juveniles of O. whitei growing near to adults of the non-susceptible canopy tree Beilschmiedia pendula. The coincidence of spatial patterns of canker incidence and host mortality suggest a role for the disease in regulating host spatial distribution, in agreement with predictions of the Janzen-Connell hypothesis.     

Skeletal Pathologies in a Population of Alouatta palliata: Behavioral, Ecological, and Evolutionary Implications

We examined the pathological conditions in a sample of 200 skulls of Alouatta palliata recovered from Barro Colorado Island (BCI), Panama, to investigate the prevalence of disease and injury in a free-ranging monkey population. Forty-five individuals (22.5%) have some type of nondental pathological condition, indicating that illness and injury are relatively common in this population. Ten individuals, all older adult males, exhibit clear signs of antemortem trauma in the form of healed fractures. We attribute the trauma primarily to fighting, and its frequency (16.4% of adult males) contradicts previous assertions that BCI howlers are nonaggressive. Nine of the 37 scorable immature individuals (24%) have signs of a larval infestation, supporting the suggestion that ectoparasites play an important role in BCI howler mortality. Other pathological conditions in the sample include shifted muscle attachments, periostitis, arthritis, neoplasm, periapical abscesses, and facial deformity, as well as various dental abnormalities.     

Beyond species area curves: application of a scale‐free measure for spatial variability of species richness

We report a novel pattern in species richness, complementary to the well‐known species–area relationship. We show that, as sample area increases, the variation in relative richness decreases among otherwise comparable spatial units. This pattern holds for southern African birds, French birds, Cape Proteaceae and the trees of Barro Colorado Island. We propose a scale‐free method for quantifying this pattern by measuring the multifractal intensity of species richness, which is the multi‐scale tendency of adjacent patches with the same area to differ in richness. By this measure, spatial variability is strongest for Cape Proteaceae and weakest for Barro Colorado Island trees. Our results have implications for area‐dependent estimates of species‐richness, for example in reserve planning and in simulation‐based studies. They imply that such estimates are most accurate for large areas, and will be subject to substantial uncertainty when the multifractal intensity is high and the area is small. For comparative purposes, multifractal intensity may be used as a supplement or as an alternative to mean richness, as well as for other ecological densities, such as biomass distribution and local abundance.