Facial and ocular thermal mapping in black-and-gold howler monkey (Alouatta caraya) by infrared thermography: An ex situ study.

BACKGROUND: This study used infrared thermography (IRT) for mapping the facial and ocular temperatures of howler monkeys, to determine parameters for the diagnosis of febrile processes. There are no published IRT study in this species. METHODS: Were evaluated images of a group of monkeys kept under human care at Sorocaba Zoo (São Paulo, Brazil). The images were recorded during 1 year, in all seasons. Face and eye temperatures were evaluated. RESULTS: There are statistically significant differences in face and eye temperatures. Mean values and standard deviations for facial and ocular temperature were respectively: 33.0°C (2.1) and 36.5°C (1.9) in the summer; 31.5°C (4.5) and 35.3°C (3.6) in the autumn; 30.0°C (4.3) and 35.6°C (3.9) in the winter; 30.8°C (2.9) and 35.5°C (2.1) in the spring. CONCLUSIONS: The IRT was effective to establish a parameter for facial and ocular temperatures of black-and-gold howler monkeys kept under human care.

Measuring mantled howler monkey (Alouatta palliata) testes via parallel laser photogrammetry: Expanding the use of noninvasive methods.

Parallel laser photogrammetry (PLP), which consists of attaching two or three parallel laser beams at a known inter-beam distance to a camera, can be used to collect morphological measurements of organisms noninvasively. The lasers project onto the photo being taken, and because the inter-beam distance is known, they act as a scale for image analysis programs like ImageJ. Traditionally, this method has been used to measure larger morphological traits (e.g., limb length, crown-rump length) to serve as proxies for overall body size, whereas applications to smaller anatomical features remain limited. To that end, we used PLP to measure the testes of 18 free-living mantled howler monkeys (Alouatta palliata) at La Selva Biological Station, Costa Rica. We tested whether this method could reliably measure this relatively small and globular morphology, and whether it could detect differences among individuals. We tested reliability in three ways: within-photo (coefficient of variation [CV] = 4.7%), between-photo (CV = 5.5%), and interobserver (intraclass correlation = 0.92). We found an average volume of 36.2 cm3 and a range of 16.4-54.4 cm3, indicating variation in testes size between individuals. Furthermore, these sizes are consistent with a previous study that collected measurements by hand, suggesting that PLP is a useful method for making noninvasive measurements of testes.

Mantled howler monkey males assess their rivals through formant spacing of long-distance calls.

Formant frequency spacing of long-distance vocalizations is allometrically related to body size and could represent an honest signal of fighting potential. There is, however, only limited evidence that primates use formant spacing to assess the competitive potential of rivals during interactions with extragroup males, a risky context. We hypothesized that if formant spacing of long-distance calls is inversely related to the fighting potential of male mantled howler monkeys (Alouatta palliata), then males should: (1) be more likely and (2) faster to display vocal responses to calling rivals; (3) be more likely and (4) faster to approach calling rivals; and have higher fecal (5) glucocorticoid and (6) testosterone metabolite concentrations in response to rivals calling at intermediate and high formant spacing than to those with low formant spacing. We studied the behavioral responses of 11 adult males to playback experiments of long-distance calls from unknown individuals with low (i.e., emulating large individuals), intermediate, and high (i.e., small individuals) formant spacing (n = 36 experiments). We assayed fecal glucocorticoid and testosterone metabolite concentrations (n = 174). Playbacks always elicited vocal responses, but males responded quicker to intermediate than to low formant spacing playbacks. Low formant spacing calls were less likely to elicit approaches whereas high formant spacing calls resulted in quicker approaches. Males showed stronger hormonal responses to low than to both intermediate and high formant spacing calls. It is possible that males do not escalate conflicts with rivals with low formant spacing calls if these are perceived as large, and against whom winning probabilities should decrease and confrontation costs increase; but are willing to escalate conflicts with rivals of high formant spacing. Formant spacing may therefore be an important signal for rival assessment in this species.

Lead exposure and its relationship with fecal cortisol levels in black howler monkeys (Alouatta pigra).

Efficiently detecting early environmental threats to wildlife is vital for conservation. Beyond obvious dangers like habitat loss or deforestation, our study focuses on one of the most hazardous toxic metals for wildlife: lead (Pb). Pb is a widespread, cumulative, and insidious environmental pollutant that can trigger a wide range of physiological, biochemical, and behavioral disorders. In fact, Pb can cause permanent dysfunction of the major stress system, the hypothalamic pituitary adrenal (HPA) axis. We analyzed Pb and cortisol concentrations in fecal samples from Alouatta pigra in southern Mexico. Fecal samples were collected across six sites categorized as free-ranging (n = 65; conserved and disturbed) and from captive animals (n = 58). Additionally, we collected soil samples (n = 35). We found that Pb was present in 28% of fecal samples and 83% of soil samples. There was a positive relation between fecal and soil Pb levels, and fecal Pb concentration was negatively associated with cortisol levels. However, the claim of Pb being a direct interference with HPA axis requires further investigation. Given our findings, assessing wildlife exposure can be a valuable tool for understanding potential Pb exposure levels in the environment and its possible implications for human health. It can also serve as an early warning system of these consequences.

Noise intensity modulates the responses of mantled howler monkeys to anthropophony.

Anthropogenic noise is a major global pollutant but its effects on primates are poorly understood, limiting our ability to develop mitigation actions that favor their welfare and conservation. In this study, we used an experimental approach to determine the impact of variation in noise intensity on mantled howler monkeys (Alouatta palliata). We conducted the study at Los Tuxtlas (México), where we studied the physiological stress (proxied via fecal glucocorticoid metabolites, fGCM) and behavioral responses of 16 males. We played back chainsaw noise at two intensities (40 and 80 dB) and used days in which groups were not exposed to noise as matched controls. With increased noise intensity fGCM increased, vigilance and vocalizations were longer, and vigilance, vocalizations, and flight occurred quicker. Physiological and behavioral responses occurred even after low-intensity noise playbacks (i.e., 40 dB). Therefore, noise intensity is a significant factor explaining the responses of mantled howler monkeys to anthropogenic noise. These results imply that management actions aimed at eradicating anthropogenic noise are required for the conservation and welfare of mantled howler monkeys at Los Tuxtlas.

Behavioral responses of mantled howler monkeys to neighbor long-distance vocalizations.

OBJECTIVES: Long-distance vocalizations are used by primates in a variety of contexts and may have different functions. The long-distance vocalizations of howler monkeys (Alouatta spp.) underlie the spatial regulation of neighboring groups and could be associated with the defense of food resources. Here, we test the hypothesis that the behavioral responses of mantled howler monkeys (A. palliata) to neighbor long-distance vocalizations are influenced by the potential for range defensibility while accounting for location within the home range and food availability. METHODS: We studied two groups for 13 months and a total of 888 h at La Flor de Catemaco (Mexico). Group 1 had a 92-ha home range and Group 2 had a 24-ha home range. We recorded vocalizations (N = 178 calls) and movements (N = 74 movements) of focal groups following long-distance vocalizations produced by their neighbors. RESULTS: Movement responses, but not vocal responses, were predicted by range defensibility, location, and food availability. As predicted, the group living in the smaller and more defendable range showed stronger movement responses than the group in the larger home range. These movement responses had a shorter latency and longer duration in more valuable spatial and temporal contexts (i.e., the core area and during periods of low food availability). CONCLUSIONS: These results suggest that the tradeoff between the costs and benefits of range defense varies according to the interactions between home range size and both the spatial (core areas) and temporal (food availability) abundance of resources. Thus, the responses of mantled howler monkeys to neighbor long-distance vocalizations could be related to home range defensibility.

Hematological and serum biochemistry evaluation in howler monkeys (Alouatta caraya) and capuchin monkeys (Sapajus apella): A comparative study.

BACKGROUND: Evaluation of blood parameters in captive non-human primates (NHPs) is crucial for monitoring their health and ensuring that their environment meets their physiological requirements. METHODS: We performed hemogram, serum biochemistry, and parasitological exams in 20 howler monkeys and 21 capuchin monkeys. RESULTS: In both species, over 50% of the individuals presented at least one parasite. There was a negative effect of age on red blood cell (RBC), white blood cell, platelets, total protein, globulin, and alkaline phosphatase, and a positive effect on the A:G ratio, gamma-glutamyl transferase, and mean platelet volume (MPV). Capuchin monkeys presented the highest platelets and alanine aminotransferase (ALT) values and howler monkeys presented the highest MPV, aspartate aminotransferase, ALT, amylase, glucose, bilirubin, and triglycerides values. We observed an interaction between species and sex on RBC, Htc, mean corpuscular hemoglobin concentration, and cholesterol. CONCLUSIONS: Species differences found in blood parameters may reflect differences in physiological adaptations associated with ecological and morphological traits and are clinically relevant for evaluating animal health and the suitability of breeding programs.

Behavioral responses to riparian and anthropogenic edge effects in mantled howler monkeys (Alouatta palliata) in a disturbed riverine forest.

Fragmented forests contain natural edges, including riparian zones, and anthropogenic edges. Edges generally have lower plant density and fewer large trees than forest interior. Riparian edges, however, contain gap-specialist trees yielding leaves with high protein content, providing primates with important resources. We examined mantled howler monkeys' behavioral responses to riparian and anthropogenic edges at La Suerte Biological Research Station (LSBRS), Costa Rica. We predicted the monkeys would spend more time resting and feeding and less time traveling, and be less spatially cohesive, in both anthropogenic and riparian edges compared to forest interior due to lower resource abundance in edges, and in anthropogenic compared to riparian edge due to higher leaf quality in riparian zones. From 2017 to 2020, we collected data across forest zones on activity and spatial cohesion patterns via focal sampling, recording data every 2 min. Howler monkeys were significantly more likely to rest and significantly less likely to travel in both anthropogenic and riparian edges compared to forest interior; however, there were no differences between these edge types. There were significantly more monkeys within a 5-m radius of focal subjects in both anthropogenic and riparian edges compared to forest interior, but no differences between these edge types. While prior research found no differences across zones when only anthropogenic edge and forest interior were compared, results of this study demonstrate that howler monkeys at LSBRS modify their activity patterns in anthropogenic and riparian edge zones compared to forest interior, highlighting the importance of focusing on both natural and anthropogenic edge zones to fully understand primates' behavioral responses in fragmented landscapes.

Immunohematological features of free-living Alouatta belzebul (Linnaeus, 1766) red-handed howler monkeys in the Eastern Amazon.

The red-handed howler monkey (Alouatta belzebul) is one of the 35 threatened Brazilian primate species found in two highly endangered Brazilian biomes. Their Amazonian native populations have been declining due to exponential deforestation associated with human activities, especially the construction of dams. The studied population (n = 27) was located in the Belo Monte dam Area of Influence. For the first time, we presented hematological parameters and the basic profile of T (CD3) and B (BSAP PAX5) cells by immunocytochemistry. The results supported the hypothesis that the immuno-hematological profile is influenced by sex, age, and season. Eosinophils were significantly higher in females (p = 0.03), monocytes statistically greater in juveniles (p = 0.04), and total plasma protein increased significantly (p > 0.001) during the dry season. Furthermore, adults showed a statistically higher average absolute number of B lymphocytes than young individuals (p = 0.03), in contrast to T lymphocytes. Even without knowing the full history of antigenic exposure, these results not only contribute to elucidating the boundaries between health and disease but may help lay the groundwork for future research into the effects of anthropogenic stress on immune activation.

The faecal metabolome of black howler monkeys (Alouatta pigra) varies in response to seasonal dietary changes.

Mammals rely on the metabolic functions of their gut microbiota to meet their energetic needs and digest potentially toxic components in their diet. The gut microbiome plastically responds to shifts in host diet and may buffer variation in energy and nutrient availability. However, it is unclear how seasonal differences in the gut microbiome influence microbial metabolism and nutrients available to hosts. In this study, we examine seasonal variation in the gut metabolome of black howler monkeys (Alouatta pigra) to determine whether those variations are associated with differences in gut microbiome composition and nutrient intake, and if plasticity in the gut microbiome buffers shortfalls in energy or nutrient intake. We integrated data on the metabolome of 81 faecal samples from 16 individuals collected across three distinct seasons with gut microbiome, nutrient intake and plant metabolite consumption data from the same period. Faecal metabolite profiles differed significantly between seasons and were strongly associated with changes in plant metabolite consumption. However, microbial community composition and faecal metabolite composition were not strongly associated. Additionally, the connectivity and stability of faecal metabolome networks varied seasonally, with network connectivity being highest during the dry, fruit-dominated season when black howler monkey diets were calorically and nutritionally constrained. Network stability was highest during the dry, leaf-dominated season when most nutrients were being consumed at intermediate rates. Our results suggest that the gut microbiome buffers seasonal variation in dietary intake, and that the buffering effect is most limited when host diet becomes calorically or nutritionally restricted.

Effect of age and sex in renal function by ultrasound and serum chemistry in two primate species (Alouatta caraya and Sapajus apella).

BACKGROUND: Comparative studies of kidney morphophysiology in nonhuman primates can help us investigate interspecies differences in growth and aging patterns. METHODS: We tested the effect of age and sex in renal morphophysiology in 21 Alouatta caraya and 21 Sapajus apella (age range = 0.5-26 years) by ultrasound, red blood cell (RBC) count, and kidney function tests. RESULTS: A. caraya had greater growth rate and absolute renal volume than S. apella, but the latter showed a greater relative renal volume and RBC count. There was a negative relationship between RBC and age, a positive relationship between creatinine and body mass, and an apparent negative relationship between creatinine and age only in S. apella. CONCLUSIONS: Our results indicate that A. caraya has a faster aging mechanism than S. apella, and the higher relative kidney volume in S. apella is suggestive of high metabolic demands in this species.

The influence of anthropogenic noise on the behavior of male mantled howler monkeys.

Anthropogenic noise is increasingly disturbing natural soundscapes and affecting the physiology, behavior, and fitness of wildlife. However, our knowledge about the impact of anthropogenic noise on wild primates is scant. Here, we assess the effects of anthropogenic noise on the behavior of male mantled howler monkeys (Alouatta palliata). Specifically, we describe the types, rates, and sound pressure level (SPL) of anthropogenic noise that occurs in areas inhabited by mantled howler monkeys and determine if the behavioral responses of males to anthropogenic noise are influenced by noise attributes. For 1 year (1753 h), we characterized anthropogenic noise in the Los Tuxtlas Biosphere Reserve (Veracruz, Mexico) and studied the behavior of males belonging to five groups. Anthropogenic noise was common, diverse, and varied among areas in terms of rate, type, and SPL. Males did not display behavioral responses toward most (60%) anthropogenic noises, but were more likely to respond to certain noise types (e.g., aerial traffic) and toward noise with high SPL. Group identity influenced the likelihood of displaying behavioral responses to noise. The most common behavioral responses were vocalizations and vigilance. Males vocalized in response to noise with high SPL, although this relationship depended on group identity. The effect of the number of noises on vocalizations also varied among groups. Males were more likely to display vigilance toward high SPL and infrequent noise, but, again, these relationships varied among groups. In sum, anthropogenic noise is pervasive in areas inhabited by mantled howler monkeys and influences male behavior. Experience and frequency of exposure may modulate the behavioral responses of male mantled howler monkeys to noise and explain the group differences.

Wild robust capuchin monkey interactions with sympatric primates.

Examining interactions among sympatric primate species can provide interesting information about competition, cooperation, and avoidance between those species. Those interactions can be neutral, positive, or negative for the species involved. Capuchin monkeys are medium-sized primates that can encounter both larger and smaller primates in their varied habitats. Gracile capuchins (Cebus) are reported to present different types of interactions with other primates. Interactions with howler monkeys frequently include physical aggression, while interactions with spider monkeys are mostly threats and chases. Moreover, interaction types are not consistent across populations. Among robust capuchins (Sapajus spp.), however, no reports have been published. Here we describe and classify encounters of Sapajus libidinosus and S. nigritus with Alouatta caraya, A. guariba, Brachyteles arachnoides, and Callithrix jacchus in three sites in the environments of Cerrado, Caatinga (savannah-like), and Atlantic forest, and compare the interaction patterns among sites and different group sizes. The latter is a factor that can influence the outcome, and we expected capuchins in larger groups to be more aggressive toward other primates. Our results of 8421 h of total contact with the capuchin groups show that, indeed, capuchins in sites with larger groups presented aggressive interactions with higher frequency. However, the other species' body size also seems important as smaller primates apparently avoided capuchins, and interactions with the larger muriquis were mostly neutral for the capuchin. Capuchins showed neutral or aggressive behaviors toward howler monkeys, with differences between the rainforest and savannah groups. We found that robust capuchins can present aggressive interactions even to primates larger than themselves and that aggressive behavior was the most common response in populations living in larger groups and drier environments.

Anthropogenic edges impact howler monkey (Alouatta palliata) feeding behaviour in a Costa Rican rainforest.

Anthropogenic forest fragmentation impacts many aspects of animal behaviour, including feeding ecology. With forests increasingly fragmented in tropical regions due to human development, the proportion of forest edge (≤ 100 m from clear-cut regions) is higher relative to forest interior. Forest edges differ in vegetation from interior, making it important to better understand how anthropogenic edges impact the feeding behaviour of primates such as mantled howler monkeys (Alouatta palliata). We predicted that howler monkeys would feed on higher-quality plant resources, from a larger number of tree families, and from larger trees in forest interior compared to anthropogenic forest edge. We surveyed howler monkey feeding behaviour across forest zones in a fragmented rainforest in Costa Rica, La Suerte Biological Research Station. We observed individual monkeys for 30-minute periods, collecting data on their feeding behaviour and tree use at 2-minute intervals. We measured feeding trees and recorded the plant parts and taxonomy of resources consumed. Monkeys consumed more leaves and fewer stems and fed from a smaller number of tree families in the forest interior, while they consumed fewer leaves and more stems and fed from a larger number of tree families in the forest edge. Monkeys also fed from larger, taller trees in the forest interior than the edge. The differences in howler monkey feeding behaviour between forest zones attest to the impact of human disturbance on howler monkey feeding ecology.

Maternal care according to offspring sex and maternal physical condition in mantled howler monkeys (Alouatta palliata).

The Trivers-Willard hypothesis (TWH) posits that maternal care will be biased in favor of the sex that provides the greatest fitness returns per unit of investment, depending on maternal physical condition. Our aim was to examine the TWH in mantled howler monkeys living at Los Tuxtlas (Veracruz, Mexico). The biological attributes of mantled howler monkeys (Alouatta palliata) meet the assumptions of TWH better than those of other explanations, so we expected that females in better physical condition should bias maternal care toward sons, whereas mothers in worse physical condition should bias care toward daughters. Between December 2017 and March 2019, we studied mother-infant interactions in 20 dyads with focal-animal sampling and continuous recording (N = 204 h). We performed genetic analysis to determine offspring sex (N = 7 daughters and 13 sons) and measured C-peptide in urine samples of mothers to assess their physical condition (N = 46 samples). Mothers in better physical condition spent less time in contact with their sons but more time in contact with their daughters. For proximity behavior, mothers in better physical condition spent more time near their sons and less time near their daughters. These results suggest a bias in maternal care towards daughters, contrary to our predictions. In light of current models of maternal investment, our results support that mothers obtain higher fitness returns through daughters.

Female reproductive energetics in mantled howler monkeys (Alouatta palliata): A follow-up study.

OBJECTIVE: Reproduction entails several challenges to primate females, among which energetic costs are remarkable at certain stages of the reproductive cycle. Still, females may use behavioral and physiological strategies to cope with those challenges. We had previously reported covariation between female energetic condition through the reproductive cycle and time-budget adjustments in mantled howler monkeys (Alouatta palliata). Accordingly, we suggested that behavioral flexibility allowed coping with the energetic challenges of reproduction. Subsequent evidence from the same population, however, suggested otherwise, so we performed a follow-up study on the variation in female reproductive energetics based on a larger sample of females. METHODS: We studied 48 free-ranging adult females at Los Tuxtlas (Mexico). We assessed energy balance via urinary C-peptide concentrations (2717 urine samples), behavioral energy intake and expenditure (5728 sampling hours), and physiological energy expenditure via fecal triiodothyronine metabolites (fTH3; 3138 fecal samples). RESULTS: We found that energy balance varied among reproductive states: (a) cycling was a period of low C-peptide concentrations; (b) the highest C-peptide concentrations occurred during gestation; and (c) the beginning of lactation marked a notable decrease in C-peptide concentrations, which then improved at mid-lactation to again decline at lactation offset. These peaks and valleys in energy balance did not seem to be associated with variation in energy acquisition but were rather mirrored by activity levels and fTH3 during lactation. DISCUSSION: Energy balance was not preserved through the reproductive cycle, supporting previous contentions that the reproductive performance of female mantled howler monkeys may be energetically constrained. The contrast between these and results that we have previously reported, highlights the importance of conducting follow-up studies to continually improve our understanding of the reproductive energetics of primate females.

Fibular and metatarsal osteosynthesis in a southern brown howler monkey (Alouatta guariba clamitans).

Here we describe a successful surgical management of a distal fibular fracture combined with a tarsocrural luxation and multiple metatarsal fractures in the left foot of a southern brown howler monkey (Alouatta guariba clamitans). We achieved satisfactory outcome by applying intramedullary pinning for each of the bone fractures and closed reduction of joint luxation-kept in place only by bone alignment, without further ligament reconstruction. Bone healing occurred uneventfully within eight weeks and the monkey's foot regained its normal function. Therefore, we could properly release the patient back into the wild.

Lone males: Solitary and group-living male howler monkey (Alouatta palliata) behavioral ecology in a Costa Rican rainforest.

OBJECTIVES: Many group-living primate species have evolved the capacity for some individuals to live alone for part of their lives, but this solitary life stage has rarely been the subject of focused research. The mantled howler monkey (Alouatta palliata) is a social primate species with bisexual dispersal that lives in mixed-sex groups with low male-to-female sex ratios. Consequently, males often spend a long period of their lives as solitary individuals. This study compares the tree use, feeding, and long-distance vocalization behavior of solitary and group-living mantled howler monkey males living within a fragmented rainforest in Costa Rica, La Suerte Biological Research Station. Based on differences in competitive ability between solitary and group-living males, we predicted that lone males would be found in significantly smaller feeding and resting trees, consume more low-quality foods, and produce shorter howling bouts made at lower rates than group-living males. MATERIALS AND METHODS: We collected data on tree use and feeding during 30-min focal samples on male focal animals, recording data at 2-min intervals. We measured the trees in which the monkeys fed and rested for two or more intervals, and recorded the plant parts consumed. We recorded howling behavior using all-occurrences sampling. RESULTS: Lone males used significantly smaller feeding and resting trees, consumed more low-quality foods, and howled at lower rates but had longer howling bouts triggered by anthropogenic noise more than group-living males. DISCUSSION: Our results demonstrate that lone males differ in their behavioral ecology compared to group-living males, thus improving understanding of the solitary male life stage in primates.

Mantled Howler Monkeys (Alouatta palliata) in a Costa Rican Forest Fragment Do Not Modify Activity Budgets or Spatial Cohesion in Response to Anthropogenic Edges.

Forest fragmentation increases forest edge relative to forest interior, with lower vegetation quality common for primates in edge zones. Because most primates live in human-modified tropical forests within 1 km of their edges, it is critical to understand how primates cope with edge effects. Few studies have investigated how primates inhabiting a fragment alter their behaviour across forest edge and interior zones. Here we investigate how anthropogenic edges affect the activity and spatial cohesion of mantled howler monkeys (Alouatta palliata) at the La Suerte Biological Research Station (LSBRS), a Costa Rican forest fragment. We predicted the monkeys would spend greater proportions of their activity budget feeding and resting and a lower proportion travelling in edge compared to forest interior to compensate for lower resource availability in the edge. We also predicted that spatial cohesion would be lower in the edge to mitigate feeding competition. We collected data on activity and spatial cohesion (nearest neighbour distance; number of individuals within 5 m) in forest edge and interior zones via instantaneous sampling of focal animals. Contrary to predictions, the monkeys spent equal proportions of time feeding, resting and travelling in forest edge and interior. Similarly, there were no biologically meaningful differences in the number of individuals or the distance between nearest neighbours in the edge (1.0 individuals; 1.56 m) versus the interior (0.8 individuals; 1.73 m). Our results indicate that A. palliata at LSBRS do not adjust their activity or spatial cohesion patterns in response to anthropogenic edge effects, suggesting that the monkeys here exhibit less behavioural flexibility than A. palliata at some other sites. To develop effective primate conservation plans, it is therefore crucial to study primate species' responses to fragmentation across their geographic range.

The occurrence of the red-handed howler monkey (Alouatta belzebul) in amazonian savannas is related to forest patch area and density of flooded area palms.

All Neotropical primates are arboreal and thus depend on forests for their survival. Arboreality puts many Neotropical primates at risk of extinction due to the high rates of deforestation in the tropics. We assessed the influence of vegetation structure and forest patch attributes on the occurrence of the threatened red-handed howler monkey (Alouatta belzebul) in an Amazonian savanna. Using a sample of 38 forest patches in a region of approximately 2000 km2 in the state of Amapá, northern Brazil, we used logistic regression to find the best predictors of the occurrence of A. belzebul. We assessed patch area, patch isolation, the proportion of seasonally flooded forest in the patch, the density of flooded area palms, forest height, canopy cover, and diameter at breast height of trees. Patch area and palm density were the best predictors of the occurrence of A. belzebul in forest patches, both having a positive effect on the probability of occurrence. Our results indicate that areas of flooded forest in forest patches may be keystone habitats for A. belzebul living in Amazonian savannas. The observed effect of palm density on A. belzebul suggests that this variable is useful for planning conservation actions, including the selection of areas for protection and management strategies for areas inhabited by this primate.