Browsing by Subject "Alouatta"
Now showing 1 - 19 of 19
Results Per Page
Sort Options
Item Open Access A coprological survey of parasites of wild mantled howling monkeys, Alouatta palliata palliata.(J Wildl Dis, 1990-10) Stuart, MD; Greenspan, LL; Glander, KE; Clarke, MRFecal samples from 155 mantled howling monkeys (Alouatta palliata palliata) examined at Centro Ecologico La Pacifica, Guanacaste Province, Costa Rica, revealed 75 (48%) had parasitic infections. A sampling of nine howling monkeys from Santa Rosa National Park. Costa Rica indicated only one infected animal (11%). Only three of 19 (16%) spider monkeys (Ateles geoffroyi) also from Santa Rosa were infected. Controrchis biliophilus, Trypanoxyuris minutus, unidentified strongylid eggs and Isospora sp. oocysts were found. Three monkeys from La Pacifica died and were examined for adult helminths. They were infected with Ascaris lumbricoides, C. biliophilus and T. minutus.Item Open Access An assessment of skin temperature gradients in a tropical primate using infrared thermography and subcutaneous implants.(J Therm Biol, 2017-01) Thompson, Cynthia L; Scheidel, Caleb; Glander, Kenneth E; Williams, Susan H; Vinyard, Christopher JInfrared thermography has become a useful tool to assess surface temperatures of animals for thermoregulatory research. However, surface temperatures are an endpoint along the body's core-shell temperature gradient. Skin and fur are the peripheral tissues most exposed to ambient thermal conditions and are known to serve as thermosensors that initiate thermoregulatory responses. Yet relatively little is known about how surface temperatures of wild mammals measured by infrared thermography relate to subcutaneous temperatures. Moreover, this relationship may differ with the degree that fur covers the body. To assess the relationship between temperatures and temperature gradients in peripheral tissues between furred and bare areas, we collected data from wild mantled howling monkeys (Alouatta palliata) in Costa Rica. We used infrared thermography to measure surface temperatures of the furred dorsum and bare facial areas of the body, recorded concurrent subcutaneous temperatures in the dorsum, and measured ambient thermal conditions via a weather station. Temperature gradients through cutaneous tissues (subcutaneous-surface temperature) and surface temperature gradients (surface-ambient temperature) were calculated. Our results indicate that there are differences in temperatures and temperature gradients in furred versus bare areas of mantled howlers. Under natural thermal conditions experienced by wild animals, the bare facial areas were warmer than temperatures in the furred dorsum, and cutaneous temperature gradients in the face were more variable than the dorsum, consistent with these bare areas acting as thermal windows. Cutaneous temperature gradients in the dorsum were more closely linked to subcutaneous temperatures, while facial temperature gradients were more heavily influenced by ambient conditions. These findings indicate that despite the insulative properties of fur, for mantled howling monkeys surface temperatures of furred areas still demonstrate a relationship with subcutaneous temperatures. Given that most mammals possess dense fur, this provides insight for using infrared imaging in thermoregulatory studies of wild animals lacking bare skin.Item Open Access Body temperature and thermal environment in a generalized arboreal anthropoid, wild mantled howling monkeys (Alouatta palliata).(Am J Phys Anthropol, 2014-05) Thompson, Cynthia L; Williams, Susan H; Glander, Kenneth E; Teaford, Mark F; Vinyard, Christopher JFree-ranging primates are confronted with the challenge of maintaining an optimal range of body temperatures within a thermally dynamic environment that changes daily, seasonally, and annually. While many laboratory studies have been conducted on primate thermoregulation, we know comparatively little about the thermal pressures primates face in their natural, evolutionarily relevant environment. Such knowledge is critical to understanding the evolution of thermal adaptations in primates and for comparative evaluation of humans' unique thermal adaptations. We examined temperature and thermal environment in free-ranging, mantled howling monkeys (Alouatta palliata) in a tropical dry forest in Guanacaste, Costa Rica. We recorded subcutaneous (Tsc ) and near-animal ambient temperatures (Ta ) from 11 animals over 1586.5 sample hours during wet and dry seasons. Howlers displayed considerable variation in Tsc , which was largely attributable to circadian effects. Despite significant seasonal changes in the ambient thermal environment, howlers showed relatively little evidence for seasonal changes in Tsc . Howlers experienced warm thermal conditions which led to body cooling relative to the environment, and plateaus in Tsc at increasingly warm Ta . They also frequently faced cool thermal conditions (Ta < Tsc ) in which Tsc was markedly elevated compared with Ta . These data add to a growing body of evidence that non-human primates have more labile body temperatures than humans. Our data additionally support a hypothesis that, despite inhabiting a dry tropical environment, howling monkeys experience both warm and cool thermal pressures. This suggests that thermal challenges may be more prevalent for primates than previously thought, even for species living in nonextreme thermal environments.Item Open Access Dental microwear in live, wild-trapped Alouatta palliata from Costa Rica.(Am J Phys Anthropol, 1991-07) Teaford, MF; Glander, KEOne problem with dental microwear analyses of museum material is that investigators can never be sure of the diets of the animals in question. An obvious solution to this problem is to work with live animals. Recent work with laboratory primates has shown that high resolution dental impressions can be obtained from live animals. The purpose of this study was to use similar methods to begin to document rates and patterns of dental microwear for primates in the wild. Thirty-three Alouatta palliata were captured during the wet season at Hacienda La Pacifica near Canas, Costa Rica. Dental impressions were taken and epoxy casts of the teeth were prepared using the methods of Teaford and Oyen (1989a). Scanning electron micrographs were taken of the left mandibular second molars at magnifications of 200x and 500x. Lower magnification images were used to calculate rates of wear, and higher magnification images were used to measure the size and shape of microwear features. Results indicate that, while basic patterns of dental microwear are similar in museum samples and samples of live, wild-trapped animals of the same species, ecological differences between collection locales may lead to significant intraspecific differences in dental microwear. More importantly, rates of microwear provide the first direct evidence of differences in molar use between monkeys and humans.Item Open Access Dental topography and molar wear in Alouatta palliata from Costa Rica.(Am J Phys Anthropol, 2004-10) Dennis, John C; Ungar, Peter S; Teaford, Mark F; Glander, Kenneth EPaleoprimatologists depend on relationships between form and function of teeth to reconstruct the diets of fossil species. Most of this work has been limited to studies of unworn teeth. A new approach, dental topographic analysis, allows the characterization and comparison of worn primate teeth. Variably worn museum specimens have been used to construct species-specific wear sequences so that measurements can be compared by wear stage among taxa with known differences in diet. This assumes that individuals in a species tend to wear their molar teeth in similar ways, a supposition that has yet to be tested. Here we evaluate this assumption with a longitudinal study of changes in tooth form over time in primates. Fourteen individual mantled howling monkeys (Alouatta palliata) were captured and then recaptured after 2, 4, and 7 years when possible at Hacienda La Pacifica in Costa Rica between 1989-1999. Dental impressions were taken each time, and molar casts were produced and analyzed using dental topographic analysis. Results showed consistent decreases in crown slope and occlusal relief. In contrast, crown angularity, a measure of surface jaggedness, remained fairly constant except with extreme wear. There were no evident differences between specimens collected in different microhabitats. These results suggest that different individual mantled howling monkeys wear their teeth down in similar ways, evidently following a species-specific wear sequence. Dental topographic analysis may therefore be used to compare morphology among similarly worn individuals from different species.Item Open Access Drinking from arboreal water sources by mantled howling monkeys (Alouatta palliata Gray).(Folia Primatol (Basel), 1978) Glander, KEDespite occasional trips to the ground and feeding in trees whose canopies touched the river, mantled howling monkeys were never seen to drink from any ground water. Drinking from arboreal cisterns was observed, but only during the wet season (meteorologically the less stressful season but phenologically the more stressful season). The lack of sufficient new leaves during the wet season forced the howlers to ingest more mature leaves which contained significantly less water. To compensate for the lowered amount of water in their food, the monkeys utilized arboreal water cisterns. The cisterns dried up during the dry season, but the howlers maintained their water balance by altering their time of actiivity and selecting a diet comprised largely of succulent new leaves. The effect of plant-produced secondary compounds on drinking also was discussed.Item Open Access Dust accumulation in the canopy: a potential cause of dental microwear in primates.(Am J Phys Anthropol, 1995-06) Ungar, PS; Teaford, MF; Glander, KE; Pastor, RFDental microwear researchers consider exogenous grit or dust to be an important cause of microscopic wear on primate teeth. No study to date has examined the accumulation of such abrasives on foods eaten by primates in the forest. This investigation introduces a method to collect dust at various heights in the canopy. Results from dust collection studies conducted at the primate research stations at Ketambe in Indonesia, and Hacienda La Pacifica in Costa Rica indicate that 1) grit collects throughout the canopy in both open country and tropical rain forest environments; and 2) the sizes and concentrations of dust particles accumulated over a fixed period of time differ depending on site location and season of investigation. These results may hold important implications for the interpretation of microwear on primate teeth.Item Open Access Dust in the wind: How climate variables and volcanic dust affect rates of tooth wear in Central American howling monkeys.(Am J Phys Anthropol, 2016-02) Spradley, Jackson P; Glander, Kenneth E; Kay, Richard FOBJECTIVES: Two factors have been considered important contributors to tooth wear: dietary abrasives in plant foods themselves and mineral particles adhering to ingested food. Each factor limits the functional life of teeth. Cross-population studies of wear rates in a single species living in different habitats may point to the relative contributions of each factor. MATERIALS AND METHODS: We examine macroscopic dental wear in populations of Alouatta palliata (Gray, 1849) from Costa Rica (115 specimens), Panama (19), and Nicaragua (56). The sites differ in mean annual precipitation, with the Panamanian sites receiving more than twice the precipitation of those in Costa Rica or Nicaragua (∼3,500 mm vs. ∼1,500 mm). Additionally, many of the Nicaraguan specimens were collected downwind of active plinian volcanoes. Molar wear is expressed as the ratio of exposed dentin area to tooth area; premolar wear was scored using a ranking system. RESULTS: Despite substantial variation in environmental variables and the added presence of ash in some environments, molar wear rates do not differ significantly among the populations. Premolar wear, however, is greater in individuals collected downwind from active volcanoes compared with those living in environments that did not experience ash-fall. DISCUSSION: Volcanic ash seems to be an important contributor to anterior tooth wear but less so in molar wear. That wear is not found uniformly across the tooth row may be related to malformation in the premolars due to fluorosis. A surge of fluoride accompanying the volcanic ash may differentially affect the premolars as the molars fully mineralize early in the life of Alouatta.Item Open Access Field methods for capture and measurement of three monkey species in Costa Rica.(Folia Primatol (Basel), 1991) Glander, KE; Fedigan, LM; Fedigan, L; Chapman, CA total of 54 free-ranging monkeys were captured and marked in Santa Rosa National Park, Costa Rica, during May 1985, and an additional 17 were captured during March 1986. The animals were darted using a blowpipe or a CO2 gun. The drugs used were Ketaset, Sernylan and Telazol. Ketaset was effective for Cebus capucinus but unsuccessful for Alouatta palliata and Ateles geoffroyi. Sernylan was successful for A. geoffroyi and A. palliata but is no longer commercially available. Telazol proved to be an excellent alternative capture drug for both A. palliata and A. geoffroyi.Item Open Access Getting Humans Off Monkeys' Backs: Using Primate Acclimation as a Guide for Habitat Management Efforts.(Integrative and comparative biology, 2020-05-29) Thompson, Cynthia L; Williams, Susan H; Glander, Kenneth E; Teaford, Mark F; Vinyard, Christopher JWild primates face grave conservation challenges, with habitat loss and climate change projected to cause mass extinctions in the coming decades. As large-bodied Neotropical primates, mantled howling monkeys (Alouatta palliata) are predicted to fare poorly under climate change, yet are also known for their resilience in a variety of environments, including highly disturbed habitats. We utilized ecophysiology research on this species to determine the morphological, physiological, and behavioral mechanisms howlers employ to overcome ecological challenges. Our data show that howlers at La Pacifica, Costa Rica are capable of modifying body size. Howlers displayed reduced mass in warmer, drier habitats, seasonal weight changes, frequent within-lifetime weight fluctuations, and gradual increases in body mass over the past four decades. These within-lifetime changes indicate a capacity to modify morphology in a way that can impact animals' energetics and thermodynamics. Howlers are also able to consume foods with a wide variety of food material properties by altering oral processing during feeding. While this capability suggests some capacity to cope with the phenological shifts expected from climate change and increased habitat fragmentation, data on rates of dental microwear warns that these acclimations may also cost dental longevity. Lastly, we found that howlers are able to acclimate to changing thermal pressures. On shorter-term daily scales, howlers use behavioral mechanisms to thermoregulate, including timing activities to avoid heat stress and utilizing cool microhabitats. At the seasonal scale, animals employ hormonal pathways to influence heat production. These lines of evidence cumulatively indicate that howlers possess morphological, physiological, and behavioral mechanisms to acclimate to environmental challenges. As such, howlers' plasticity may facilitate their resilience to climate change and habitat loss. While habitat loss in the tropics is unlikely to abate, our results point to a potential benefit of active management and selective cultivation to yield large, interconnected forest fragments with targeted phenology that provides both a complex physical structure and a diversity of food sources. These steps could assist howlers in using their natural acclimation potential to survive future conservation threats.Item Open Access Mechanical defenses in leaves eaten by Costa Rican howling monkeys (Alouatta palliata).(Am J Phys Anthropol, 2006-01) Teaford, MF; Lucas, PW; Ungar, PS; Glander, KEPrimate species often eat foods of different physical properties. This may have implications for tooth structure and wear in those species. The purpose of this study was to examine the mechanical defenses of leaves eaten by Alouatta palliata from different social groups at Hacienda La Pacifica in Costa Rica. Leaves were sampled from the home-ranges of groups living in different microhabitats. Specimens were collected during the wet and dry seasons from the same tree, same plant part, and same degree of development as those eaten by the monkeys. The toughness of over 300 leaves was estimated using a scissors test on a Darvell mechanical tester. Toughness values were compared between social groups, seasons, and locations on the leaves using ANOVA. Representative samples of leaves were also sun-dried for subsequent scanning electron microscopy and energy dispersive x-ray (EDX) analyses in an attempt to locate silica on the leaves. Both forms of mechanical defense (toughness and silica) were found to be at work in the plants at La Pacifica. Fracture toughness varied significantly by location within single leaves, indicating that measures of fracture toughness must be standardized by location on food items. Monkeys made some food choices based on fracture toughness by avoiding the toughest parts of leaves and consuming the least tough portions. Intergroup and seasonal differences in the toughness of foods suggest that subtle differences in resource availability can have a significant impact on diet and feeding in Alouatta palliata. Intergroup differences in the incidence of silica on leaves raise the possibility of matching differences in the rates and patterns of tooth wear.Item Open Access Mechanical defenses in leaves eaten by Costa Rican howling monkeys (Alouatta palliata).(Am J Phys Anthropol, 2006-01) Glander, Kenneth Earl; Lucas, PW; Teaford, Mark F; Ungar, PSPrimate species often eat foods of different physical properties. This may have implications for tooth structure and wear in those species. The purpose of this study was to examine the mechanical defenses of leaves eaten by Alouatta palliata from different social groups at Hacienda La Pacifica in Costa Rica. Leaves were sampled from the home-ranges of groups living in different microhabitats. Specimens were collected during the wet and dry seasons from the same tree, same plant part, and same degree of development as those eaten by the monkeys. The toughness of over 300 leaves was estimated using a scissors test on a Darvell mechanical tester. Toughness values were compared between social groups, seasons, and locations on the leaves using ANOVA. Representative samples of leaves were also sun-dried for subsequent scanning electron microscopy and energy dispersive x-ray (EDX) analyses in an attempt to locate silica on the leaves. Both forms of mechanical defense (toughness and silica) were found to be at work in the plants at La Pacifica. Fracture toughness varied significantly by location within single leaves, indicating that measures of fracture toughness must be standardized by location on food items. Monkeys made some food choices based on fracture toughness by avoiding the toughest parts of leaves and consuming the least tough portions. Intergroup and seasonal differences in the toughness of foods suggest that subtle differences in resource availability can have a significant impact on diet and feeding in Alouatta palliata. Intergroup differences in the incidence of silica on leaves raise the possibility of matching differences in the rates and patterns of tooth wear.Item Open Access Natal emigration by both sexes in the La Pacifica population of mantled howlers: when do some stay?(Am J Primatol, 2008-02) Clarke, Margaret R; Glander, Kenneth EWe have reported previously that all male and female mantled howlers emigrate from natal groups at Hacienda La Pacifica, Costa Rica. In the years since that report, a small number of juveniles have stayed in the natal group without experiencing a solitary phase. Here, we present a post hoc analysis on juvenile emigration in six groups of howlers under observation for varying amounts of time between 1972 and 2005. Our records revealed 139 juveniles for whom emigration status was certain, and 125 of these did emigrate. There was a significant association between presence of mother and emigration: juveniles without mothers were more likely to remain in their natal group (chi(1)(2) = 53.1, P<.0001). The mean age of emigration for all juveniles (n = 125) was 2.47 years (SD = 0.9, range = 1.5-6.5). There was no difference in age of emigration by adult male composition (one-male, multi-male, both), but juveniles of unknown sex emigrated younger than either known males or females (F(2,116) = 4.4, P<.02). For emigrating juveniles of known sex (n = 99), both males and females without mothers left at a later age than those with mothers (F(1,95) = 6.5, P<.02). Although philopatry or delayed emigration occurs in a few motherless animals, most males and females do emigrate from their natal groups at ages consistent with those reported for other species of howlers.Item Open Access Plasma lipoproteins of free-ranging howling monkeys (Alouatta palliata).(Comp Biochem Physiol B, 1987) Clark, SB; Tercyak, AM; Glander, KE1. Plasma lipids and lipoproteins of free-ranging howling monkeys from Costa Rica (Alouatta palliata), aged 5 months to 23 years, were characterized. 2. High density lipoproteins were lipid-rich, similar to HDL2 of human plasma. 3. Fatty acid compositions of major lipid classes of very low, low and high density lipoproteins differed among social groups, possibly due to both dietary and genetic factors. 4. Low and high density lipoprotein phospholipids were enriched in phosphatidylethanolamine. 5. Howler plasma cross reacted with antihuman apoA-I antibodies but not with antihuman LDL antibodies. 6. No dimeric form of apoA-II was present, unlike human apoA-II.Item Open Access Reproduction and population growth in free-ranging mantled howling monkeys.(Am J Phys Anthropol, 1980-07) Glander, KEFree-ranging mantled howling monkey (Alouatta palliata Gray) females experienced a regular estrus cycle averaging 16.3 days, demonstrated sexual skin changes, and participated in multiple matings before becoming pregnant. Gestation averaged 186 days. The average interval between births was 22.5 months. Sexual maturity occurred at approximately 36 and 42 months for females and males, respectively. Female age at first birth was about 3 1/2 years. Births were scattered during some years and clustered during others. The age, rank, and parity of the females affected infant survival. More female than male infants survived to one year of age. Increased population size was the result of immigration rather than births.Item Open Access Secondary transfer of adult mantled howlers (Alouatta palliata) on Hacienda La Pacifica, Costa Rica: 1975-2009.(Primates, 2010-07) Clarke, Margaret R; Glander, Kenneth ENatal emigration by male and female mantled howlers (Alouatta palliata), and subsequent immigration into breeding groups, is well documented for the free-ranging population on Hacienda La Pacifica, Costa Rica, but secondary transfer was considered rare (Glander in Int J Primatol 3:415-436, 1992). Population surveys in 1998 and 2006 caused us to question our assumptions and to re-evaluate our long-term data set from a post hoc perspective. We first identified all animals observed or captured as adults in more than one non-natal group anywhere in the population. We then systematically analyzed joining or leaving by adults in seven groups tracked for various times from 1975 to 2005 for patterns suggesting secondary transfer. Fourteen adults (nine females, five males) were found in two different non-natal groups as adults. In addition, one male and one female that became dominant and reproduced in their natal group later transferred to a second group, and one female was known to be a tertiary transfer. Data from the seven tracked social groups indicate that 35% of all the males and 29% of all the females were potential secondary transfers. In these groups, males leaving or joining was not associated with group size or absolute number of females. Females leaving or joining was not associated with group size or absolute number of males, but females left groups with more females and joined groups with fewer females. Both sexes left groups with unfavorable sex ratios for their sex and joined groups with sex ratios more favorable for their sex. Since a favorable sex ratio is associated with reproductive success in other howler populations, this suggests secondary transfer as a reproductive strategy. Other factors could also influence secondary transfer.Item Open Access Testing parallel laser image scaling for remotely measuring body dimensions on mantled howling monkeys (Alouatta palliata).(Am J Primatol, 2015-08) Barrickman, Nancy L; Schreier, Amy L; Glander, Kenneth EBody size is a fundamental variable for many studies in primate biology. However, obtaining body dimensions of wild primates through live capture is difficult and costly, so developing an alternative inexpensive and non-invasive method is crucial. Parallel laser image scaling for remotely measuring body size has been used with some success in marine and terrestrial animals, but only one arboreal primate. We further tested the efficacy of this method on the arboreal mantled howling monkey (Alouatta palliata) in La Pacifica, Costa Rica. We calculated interobserver error, as well as the method's repeatability when measuring the same animal on different occasions. We also compared measurements obtained physically through live capture with measurements obtained remotely using parallel laser image scaling. Our results show that the different types of error for the remote technique are minimal and comparable with the error rates observed in physical methods, with the exception of some dimensions that vary depending on the animals' posture. We conclude that parallel laser image scaling can be used to remotely obtain body dimensions if careful consideration is given to factors such as species-specific morphology and postural habits.Item Open Access Thyroid hormone fluctuations indicate a thermoregulatory function in both a tropical (Alouatta palliata) and seasonally cold-habitat (Macaca fuscata) primate.(Am J Primatol, 2017-11) Thompson, Cynthia L; Powell, Brianna L; Williams, Susan H; Hanya, Goro; Glander, Kenneth E; Vinyard, Christopher JThyroid hormones boost animals' basal metabolic rate and represent an important thermoregulatory pathway for mammals that face cold temperatures. Whereas the cold thermal pressures experienced by primates in seasonal habitats at high latitudes and elevations are often apparent, tropical habitats also display distinct wet and dry seasons with modest changes in thermal environment. We assessed seasonal and temperature-related changes in thyroid hormone levels for two primate species in disparate thermal environments, tropical mantled howlers (Alouatta palliata), and seasonally cold-habitat Japanese macaques (Macaca fuscata). We collected urine and feces from animals and used ELISA to quantify levels of the thyroid hormone triiodothyronine (fT3). For both species, fT3levels were significantly higher during the cooler season (wet/winter), consistent with a thermoregulatory role. Likewise, both species displayed greater temperature deficits (i.e., the degree to which animals warm their body temperature relative to ambient) during the cooler season, indicating greater thermoregulatory pressures during this time. Independently of season, Japanese macaques displayed increasing fT3levels with decreasing recently experienced maximum temperatures, but no relationship between fT3and recently experienced minimum temperatures. Howlers increased fT3levels as recently experienced minimum temperatures decreased, although demonstrated the opposite relationship with maximum temperatures. This may reflect natural thermal variation in howlers' habitat: wet seasons had cooler minimum and mean temperatures than the dry season, but similar maximum temperatures. Overall, our findings support the hypothesis that both tropical howlers and seasonally cold-habitat Japanese macaques utilize thyroid hormones as a mechanism to boost metabolism in response to thermoregulatory pressures. This implies that cool thermal pressures faced by tropical primates are sufficient to invoke an energetically costly and relatively longer-term thermoregulatory pathway. The well-established relationship between thyroid hormones and energetics suggests that the seasonal hormonal changes we observed could influence many commonly studied behaviors including food choice, range use, and activity patterns.Item Open Access When top-down becomes bottom up: behaviour of hyperdense howler monkeys (Alouatta seniculus) trapped on a 0.6 ha island.(PLoS One, 2014) Orihuela, Gabriela; Terborgh, John; Ceballos, Natalia; Glander, KennethPredators are a ubiquitous presence in most natural environments. Opportunities to contrast the behaviour of a species in the presence and absence of predators are thus rare. Here we report on the behaviour of howler monkey groups living under radically different conditions on two land-bridge islands in Lago Guri, Venezuela. One group of 6 adults inhabited a 190-ha island (Danto) where they were exposed to multiple potential predators. This group, the control, occupied a home range of 23 ha and contested access to food resources with neighbouring groups in typical fashion. The second group, containing 6 adults, was isolated on a remote, predator-free 0.6 ha islet (Iguana) offering limited food resources. Howlers living on the large island moved, fed and rested in a coherent group, frequently engaged in affiliative activities, rarely displayed agonistic behaviour and maintained intergroup spacing through howling. In contrast, the howlers on Iguana showed repulsion, as individuals spent most of their time spaced widely around the perimeter of the island. Iguana howlers rarely engaged in affiliative behaviour, often chased or fought with one another and were not observed to howl. These behaviors are interpreted as adjustments to the unrelenting deprivation associated with bottom-up limitation in a predator-free environment.