Resource use by howler monkeys (Alouatta palliata) in the rain forest of Los Tuxtlas,Veracruz, Mexico

The feeding preferences of howler monkeys at their northernmost distribution in the Neotropics are reported for an annual cycle. A remarkable selectivity for 27 species representing 15 families was observed. The Moraceae and Lauraceae plant families were the most important in the diet. The howlers spent an almost equal proportion of their feeding time eating leaves and fruit, and displayed a marked preference for young leaves and mature fruit. The consumption of different plant parts was markedly seasonal and the howlers’ ranging behavior was closely associated with the availability of young leaves and mature fruit. Their home range was unusually large (ca. 60 ha) for howlers and the food species exploited occur at very low densities (93%, ≤ 4 ind/ha). They chose food items richer in protein and energy. Alkaloid compounds, present in some of the leaves, play a secondary role in their dietary selectivity.     

Fruit eating and seed dispersal by howling monkeys (Alouatta palliata) in the tropical rain forest of Los Tuxtlas,Mexico

The frugivory and ranging habits of howling monkeys living in the rain forest of Los Tuxtlas, Veracruz, Mexico, were studied for 5 consecutive years with the aim of investigating the role of seed dispersal agents played by the howling monkeys. The howling monkeys' consumption of fruit occupied half of their feeding time, and 80% of this time was spent feeding on mature fruit. Observations confirmed use of 19 tree species and a preference for a few species of Moraceae and Lauraceae. Fruit exploitation was very seasonal; only two species provided fruit year-round. Home range size was about 60 ha, and animals in the troop (N = 16) showed a day range of 10–893 m. Monthly collection of fecal samples during 2 years indicated that 90% of the seeds (N = 7,928) in the samples belonged to eight tree species and to one liana; the rest belonged to 15 unidentified species of vines. Germination success for the seeds in the feces was about 60% and for control seeds was 35%. Howling monkeys created diverse seed shadows in the vicinity of their leaf and fruit sources, and while they dispersed the seeds of some plant species, they also produced a great deal of fruit and seed waste for others.     

Tropical rain forest fragmentation,howler monkeys (Alouatta palliata), and dung beetles at Los Tuxtlas,Mexico

In Neotropical rain forests, fresh mammal dung, especially that of howler monkeys, constitutes an important resource used by dung beetles as food and for oviposition and further feeding by their larvae. Tropical rain forest destruction, fragmentation, and subsequent isolation causing reductions in numbers of and the disappearance of howler moneys may result in decreasing numbers of dung beetles, but this has not been documented. In this study, we present information on the presence of howlers and dung beetles in 38 isolated forest fragments and 15 agricultural habitats. Howler monkeys were censused by visual means, while dung beetles were sampled with traps baited with a mixture of howler, cow, horse, and dog dung. Results indicated that loss of area and isolation of forest fragments result in significant decrements in howlers and dung beetles. However, dung beetle abundance was found to be closely related to the presence of howler monkeys at the sites and habitats investigated. Scenarios of land management designed to reduce isolation among forest fragments may help sustain populations of howler monkeys and dung beetles, which may have positive consequences for rain forest regeneration. Am. J. Primatol. 48:253–262, 1999. © 1999 Wiley‐Liss, Inc.     

Feeding and general activity patterns of a howler monkey (Alouatta palliata) troop living in a forest fragment at Los Tuxtlas,Mexico

The feeding behavior and general activity patterns of a howler monkey troop living in a 3.6 ha forest fragment were studied at Los Tuxtlas, Mexico, for an annual cycle. Monthly samples of their feeding behavior indicated that they used 52 species of 24 plant families as sources of food. Of these, 67% were trees, which accounted for 96% of total feeding time recorded. Ten species of Moraceae, Cecropiaceae, Anacrdiaceae. and Sapotaceae contributed to 70% of the trees used and to almost 90% of feeding time. The number of plant species used per monthly record varied from 7 to 31 with an average of 19.9 species. Young leaves and ripe fruit were the principal items in the monthly diet of howlers and average percent of time spent consuming these plant parts was 46.7% and 34.8%, respectively. The use of tree species was found to be associated to their importance value and to their pattern of spatial dispersal in the study site. Availability of young leaves was fairly constant from month to month, but it presented a seasonal pattern, and there was a significantly lower number of tree species bearing ripe fruit through the year with brief pulses of production. The monthly activity pattern was found to be related to variations in the availability of young leaves and ripe fruit as well as to the values of the intermonthly overlap in plant species used. Resting and feeding presented a bimodal pattern of occurrence throughout the day that seemed to be related to variations in maximum ambient temperatures. Results are discussed in light of the small size and shape of the forest fragment inhabited by the howler troop. Am. J. Primatol. 48:167–183, 1999. © 1999 Wiley‐Liss, Inc.     

A preliminary study of resource overlap between howling monkeys (Alouatta palliata) and other arboreal mammals in the tropical rain forest of Los Tuxtlas,Mexico

Potential resource overlap between howling monkeys and other arboreal mammals was studied in the rain forest of Los Tuxtlas, Veracruz, Mexico. Eight species of mammals belonging to the orders Primates, Carnivora, Rodentia, and Marsupialia were found to share the canopy and to overlap trophically with howling monkeys. These mammals made up 77% and Alouatta 23% of the arboreal mammalian biomass under consideration. The arboreal porcupine and spider monkey were the only mammals that also fed on leaves. However, in this feeding niche, Alouatta is the only important mammalian folivore in Los Tuxtlas, and resource depression derived from leaf-eating insects is more important. The eight arboreal mammals may exert more pressure upon fruit resources, for they consumed 75% of the estimated total dry weight of fruit/ha/yr consumed by arboreal mammals.     

Use of leaf resources by howling monkeys (Alouatta palliata) and leaf-cutting ants (Atta cephalotes) in the tropical rain forest of Los Tuxtlas,Mexico

Use of leaf resources by a troop of howling monkeys and two colonies of leaf cutting ants was studied for an annual cycle in the rain forest of Los Tuxtlas, Mexico. Howling monkeys spent half their annual foraging time feeding on leaves; leaf-cutting ants spent at least 80% of their recorded foraging time harvesting leaves. Both herbivores preferred young leaves over nature ones, and chemical analysis showed that the protein: fibre ratio of the leaves used was correlated with these preferences. Howling monkeys used 34 tree species as leaf sources. Leaf-cutting ants used 40 plant species of which 38 were trees. Eighteen species used by Alouatta were also used by Atta; species of Moraceae and Lauraceae were among the most important in their foraging preferences. The plant species used by monkeys and ants occurred at low densities (? 4.0 ind/ha). The seasonal production of leaves, the high density of leaf-cutting ant colonies at the study site, and the high amounts of young foliage harvested by the ants from tree species, and individual trees used by howling monkeys as sources of young leaves suggest that the foraging activities of Atta may represent a significant pressure upon leaf resources available to Alouatta.     

Comparison of frugivory by howling monkeys (Alouatta palliata) and bats (Artibeus jamaicensis) in the tropical rain forest of Los Tuxtlas,Mexico

Information on the fruit diets of howling monkeys and fruit-eating bats in the tropical rain forest of Los Tuxtlas, Mexico was collected for a year to compare the plant species used. Howling monkeys used 19 plant species whereas bats used 32 plant species as fruit sources. Eleven species were common in the diet of both mammals. A rank analysis at the plant species level showed that the fruit diets of Alouatta and Artibeus were very different. In contrast to bats, howling monkeys displayed a seasonal pattern in fruit consumption. Diet overlap between the two mammals was highest during the monkeys' fruit-eating season. Measures of fruit production in eight trees (four species) indicated marked variations in fruit biomass produced and in length of fruiting from tree to tree and species to species. Peaks in fruit production were typical both at the species and the individual tree levels, demonstrating the very patchy nature of the fruit available to the monkeys and the bats.     

Fruit choice by red howler monkeys (Alouatta seniculus) in a tropical rain forest

This paper presents and discusses aspects of fruit selectivity by red howler monkeys (Alouatta seniculus) in relation with morphological characteristics of fruits. These data are used to provide an answer to the following questions: which are the fruit characteristics that lead fruit choice of howler monkeys and to what extent fruit characteristics play a role in seed dispersal by monkeys? The frugivorous diet of a troop of red howler monkeys was determined during a 2-year field study in French Guiana. The selection of fruit by howler monkeys was analyzed in relation to the fruit availability. Results showed that, although consumption followed availability, fruit species could be classified in three categories according to their selection ratio (percentage of consumption/percentage of abundance) as “high ranking,” “middle ranking,” and “low ranking” species. Also, the 97 species of fruit eaten by the monkeys were grouped according to the morphological characteristics thought to influence the monkeys' choice. This showed that howler monkeys consumed essentially fruits with juicy pulp, bright color, and a small number of well-protected seeds. Most of high ranking species had medium-sized fruits with yellow color, and low ranking species often had small fruits. However, howler monkeys are associated with the dispersal of seeds from fruit with a hard and indehiscent pericarp and/or large seeds, like those of the Sapotaceae family. Consequently, they can be considered as “specialized” frugivores for this fruit syndrome. © l996 Wiley-Liss, Inc.     

Risk factors connected to gastrointestinal parasites in mantled Alouatta palliata mexicana and black howler monkeys Alouatta pigra living in continuous and in fragmented rainforests in Mexico

In this study we document the prevalence of gastrointestinal parasites (helminths and protozoans) found in fecal samples of populations of Alouatta palliata mexicana and of A.pigra in Los Tuxtlas and in Palenque,southeast Mexico,and its relation to habitat condition,sex/age and season.Nineteen parasite morphotypes were detected in the fecal samples from populations of the two howler monkeys,of which 58％ were shared by both species.When considering all parasite species,populations of the two howler species were more likely to be parasitized in fragmented habitat compared to continuous habitat.Individuals of both howler monkey species that lived in fragmented habitat had a higher prevalence of Controrchis biliophilus.A.p.mexicana individuals had a higher prevalence of Trypanoxyuris minutus than A.pigra,probably the result of the larger group sizes found in the former species,and T.minutus was more likely to be found in A.palliata individuals that lived in fragmented habitat.Adult A.p.mexicana males had a higher risk of being parasitized compared to adult females,but these differences were not detected in A.pigra.Parasite species such as Entamoeba sp.,Nematoda sp.28.Nematoda sp.B and Parabronema sp.where only found during the wet season m both howler monkey species.Populations of both howler monkey species had a higher prevalence of Nematoda sp.A in the wet season and Ascaridae eggs were only detected during the wet season in A.pigra.Other parasites detected displayed no clear seasonal pattern.     

Structure and floristic composition of the lowland rain forest of Los Tuxtlas,Mexico

Physiognomy, structure and floristic composition of one hectare of lowland tropical rain forest was studied in detail at Los Tuxtlas, Mexico. Physiognomically, the Los Tuxtlas forest should be classified as lowland tropical high evergreen rain forest. The forest showed a closed canopy at 30–35 m. Of all woody, non-climbing species with a DBH1.0 cm 89.4% (94.5% of all individuals) were evergreen, 25.4% (59.5% of the individuals) had compound leaves, and over 80% of species (and individuals) had leaves in the notophyll and mesophyll size classes. The forest structure was characterized by a low density (2976 individuals with a DBH1.0 cm, 346 individuals with a DBH10.0 cm, per ha, excluding vines) with an average basal area (38.1 m², DBH1.0 cm, 34.9 m², DBH10.0 cm, per ha, excluding vines). This was attributed to the relative maturity of the forest on the study plot. The study plot contained 234 species (11 208 individuals with a height 0.5 m), of which 55.1% (34.8% of individuals) were trees, 9.4% (6.8%) shrubs, 3.4% (44.3%) palms, 20.1% (5.2%) vines, 6.8% (8.7%) herbs and 5.1% (0.3%) of unknown lifeform. Furthermore, 58 species of epiphytes and hemi-epiphytes were found. Diversity of trees, shrubs and palms with a DBH1.0 cm was calculated as Shannon-Wiener index (4.65), Equitability index (0.65), and Simpson index (0.10). The dominance-diversity curve showed a lognormal form, characteristic for tropical rain forest. The community structure was characterized by a relative dominance of Astrocaryum mexicanum in the understorey, Pseudolmedia oxyphyllaria in the middle storeys, and Nectandra ambigens in the canopy. Species population structures of 31 species showed three characteristic patterns, differentiated by recruitment: continuously high, discontinuously high, and continuously low recruitment. Height/diameter and crown cover/diameter diagrams suggested a very gradual shift from height growth to crown growth during tree development. Forest turnover was calculated as 138 years. Compared to other tropical rain forests the Los Tuxtlas forest had 1. similar leaf physiognomical characteristics, 2. a lower diversity, 3. a lower density, 4. an average basal area, and 5. a slow canopy turnover.     

Bird species richness in vegetation fences and in strips of residual rain forest vegetation at Los Tuxtlas, Mexico

Fragmentation of the lowland tropical rain forest has resulted in loss of animal and plant species and isolation of remaining populations that puts them at risk. At Los Tuxtlas, Mexico, lowland rain forests are particularly diverse in the avian fauna they contain and while most of the forests have been fragmented by human activity, many of the fragments still harbor diverse assemblages of bird species. In these landscapes, linear strips of residual rain forest vegetation along streams as well as linear strips of vegetation fences (live fences) crossing the pastures might provide some connectivity to bird populations existed in forest fragments. We investigated bird species richness and relative abundance in one 6-km long section of live fences (LF) bordering a dirt road and in two 6-km long sections of residual forest vegetation along a river (MR) and one permanent stream (BS). We used point count procedures which resulted in the count of 2984 birds representing 133 species. At the LF site we detected 74% of the species, 72% at the BS site and 57% at the MR site. Only 38% of the species were common among sites. Neotropical migratory birds accounted for 34–41% of the species counted at all sites. While edge and open habitat birds accounted for 6–10% of the species and for 50% of the records at the three vegetation strips, about 90% of the species were forest birds. Distance to forest fragments and degree of disturbance of the vegetation seemed to negatively influence bird species presence at the BS and MR strips. Rarefaction analysis indicated that the LF strip was richer in species than the other two sites, but the occurrence of the three vegetation strips in the landscape seem to favor the presence of many more species. We discuss the value of these vegetation strips to birds as stepping stones in the fragmented landscape.     

Dung beetles in continuous forest, forest fragments and in an agricultural mosaic habitat island at Los Tuxtlas, Mexico

With the aim of determining what kind of landscape mosaics might sustainmaximum diversity and minimum species loss, dung beetles were sampled withbaited pitfall traps to compare species richness and species composition in atract of continuous forest, forest fragments and a habitat island consisting ofa mosaic of forest and arboreal crops in Los Tuxtlas, southern Mexico. Wecaptured 7332 dung beetles representing 33 species. Similar numbers of specieswere captured in the three habitats. However, 56% of individuals were capturedin the continuous forest, 29% in the mosaic habitat and 15% in the forestfragments. Eight species (Canthon femoralis,Copris laeviceps, Canthidium centrale,Onthophagus batesi, Deltochilumpseudoparile, O. rhinolophus, Canthonviridis vazquezae and Dichotomius satanus) accounted for 90% of thecaptures, but their relative dominance varied among habitats. A clear trend wasevident in the number of dung beetles captured in the dung processing guilds(rollers/tunnelers) as well as in the diurnal and nocturnal guilds, withcaptures decreasing from the continuous forest to the mosaic habitat to theforest fragments. A similar trend was detected in detection rates for medium andsmall size dung producing mammals. Species richness of forest fragments andmosaic habitat did not differ from that found in the continuous forest, butthese habitats differ significantly in species richness from isolated shaded andunshaded plantations, linear strips of vegetation, the forest–pasture edge andpastures according to rarefaction analysis. The co-occurrence of the continuousforest, the mosaic habitat and the cluster of forest fragments in closeproximity seems to be preserving a diverse assemblage of dung beetle species inthe local landscape.     

Tropical rain forest fragmentation and wild populations of primates at Los Tuxtlas,Mexico

In view of the extensive destruction, fragmentation, and conversion of primate habitats to anthropogenic vegetation, information on Neotropical primate ability to use a landscape consisting of forest fragments and agricultural habitats is necessary to understand the ecological flexibility of the species involved and it is of relevance to the design of conservation scenarios at the landscape level. We censused howlers and spider monkeys in 126 forest fragments and in 44 agricultural sites at Los Tuxtlas, southern Veracruz, Mexico, and used the IDRISI Geographic Information System to assess the extent of primate habitat remaining. We conducted economic surveys to assess the productivity of several systems of land management, including cattle ranching. Seventy- five percent of native habitat has been lost at Los Tuxtlas, 20% remains in the form of isolated fragments of vegetation, and only 5% consists of contiguous rain forest at high elevations (> 800 m). Howlers and spider monkeys were present in only 60 and 8% of the forest sites investigated, respectively, attesting to extensive local extinction. Populations of both species are small and found in a fragmented and isolated condition throughout the landscape. A large number of howlers were detected in artifactual habitats such as cacao, coffee and mixed (cacao and coffee) plantations shaded by rain forest trees. Residual rain forest vegetation along streams, rivers, and lakes facilitated the interfragment and interhabitat movement of howlers. Economic surveys showed that some of the agricultural habitats were more productive than cattle ranching. Conservation of spider monkeys requires large areas of contiguous forest, but only small areas are present at high elevations. Howlers are restricted to elevations < 800 m, where most of the forest has disappeared, but can apparently exist in a matrix of forest fragments, arboreal agricultural habitats, and pasture lands. Structural aspects of the vegetation and connectivity among habitat islands may be indispensable components of both landscape scenarios.     

Useful plants of the Los Tuxtlas rain forest (Veracruz, Mexico): Considerations of their market potential

The 640 hectare large Los Tuxtlas rain forest reserve was analyzed for the commercial potential of its 860 native flowering plant species. Excluding plants with medicinal or chemical use, 91 species (10.6%) are found in the market, and we consider an additional 72 species (8.4%) to have market potential. For each of the 163 species, the following information is given: scientific and common name, geographic distribution, growth form, use, and market status. Of the 163 species, 2.4% are endemic to the Los Tuxtlas region, 10.4% are restricted to Mexico, 42.3% extend further into Central America, and 39.3% occur also in South America, while 20.2% are found in the West Indies. The majority of species (68.7%) are trees. In terms of use categories, 35.0% are employed for their timber, 32.5% for their fuelwood, 24.5% for ornamental purposes, 22.7% for their edible fruits, leaves or flowers, 11.0% for plywood or paper, 6.1% for artwork and weaving, and 1.2% as fodder plants. The more than 160 marketable species would make it possible to develop species-diverse forests of high commercial value, as an alternative to current land use, which consists in converting forest into cattle pastures.     

Patterns of frugivore species richness and abundance in forest islands and in agricultural habitats at Los Tuxtlas,Mexico

Destruction and fragmentation of tropical rain forest result in a loss of species and of generating capacity of the ecosystem via animal vectors such as seed dispersal agents. To gather quantitative data regarding this ecological problem, birds and mammals were censused in 30 forest fragments, 15 agricultural islands representing five types of vegetation (coffee, cacao, citrus, pepper and mixed-crops) and in three pastures in Los Tuxtlas, southern Veracruz, Mexico. More than 6000 animals of 257 species were detected thus suggesting the existence of a rich species pool in the fragmented landscape. Frugivores accounted for 60% of species, for 72% of individuals censured and for 85% of the total animal biomass recorded. Clusters of small forest fragments (<100 ha) were richer in species and individuals than clusters of large area (>100 ha) forest islands. Pastures were especially poor in forest birds and mammals. While the agricultural islands studied contributed to only 1% of the total area of vegetation sampled, they contained 58% of all species detected and 34% of all individual birds and mammals censured. Recaptures indicated inter-island movements of forest birds and mammals. Forty percent of the species were detected in forest habitats only, the rest were detected in forest and in agricultural habitats. Seeds of forest interior plants dispersed by birds and bats were detected in the agricultural habitats. The value of agricultural islands as landscape features providing some degree of biotic connectivity among fragmented animal populations is discussed.     

Some observations on the present distribution and conservation of Alouatta and Ateles in Southern Mexico

Some information on the past and present distribution of Alouatta and Ateles in southern Mexico and on the impact of habitat destruction on their future existence is presented. Three surveys of primate populations and forest habitats were carried out between 1981 and 1983 in southeastern Mexico. Rapid destruction of the forest is reducing the number of forested areas where Alouatta and Ateles can exist and co-occur. Illegal hunting of the two primates also contributes to their rapid decline. Alouatta and Ateles live a precarious existence in a few large and small islands of forest, and efforts are needed to set aside some of these areas as reserves so that their future survival is ensured.     

Use of Fruiting Trees by Birds in Continuous Forest and Riparian Forest Remnants in Los Tuxtlas,Veracruz, Mexico1

The composition of plant species in fragmented landscapes may be influenced by the pattern of visitation by birds to fruiting trees and by the movement of seeds among and within fragments. We compared bird visitation patterns to two tree species (Dendropanax arboreus, Araliaceae; Bursera simaruba, Burseraceae) in continuous forest and remnants of riparian vegetation in a region dominated by pasture in Los Tuxtlas, Veracruz, Mexico. We quantified frequency of visitation, fruit consumption, consistency of visitation (percentage of total tree observation periods during which a given bird species was recorded), and species composition of birds at individuals of both tree species in continuous forest and riparian remnants. Bird visitation rate, species richness, and fruit consumption rates were similar within both tree species in the two habitats. Species assemblages at D. arboreus were different between continuous forest and remnants. Species assemblages at B. simaruba did not differ by habitat. Our results demonstrate that habitat disturbance may influence avian visitation patterns, which may in turn affect subsequent recruitment patterns in some tree species. Our results, however, were not consistent between the tree species, suggesting that it is difficult to generalize concerning the effects of forest disturbance on avian species assemblages in fruiting trees.     

Parasitic infections of three Mexican howler monkey groups (Alouatta palliata mexicana) living in forest fragments in Mexico

In order to better understand how patterns of parasitism in howler monkeys are affected by forest fragmentation, we carried out a 1 year survey of gastrointestinal parasites in fecal samples from three groups of Mexican howler monkeys inhabiting different forest fragments in the Los Tuxtlas Biosphere Reserve, Mexico. The study groups were chosen because the conditions in which they lived suggested a potentially negative gradient for parasite richness and a positive gradient for levels of parasitism. We report for the first time the presence of Entamoeba coli in Alouatta palliata mexicana and of hookworms (Family Ancylostomidae) in A. palliata. A reduction in home range size and an increase in disturbance was associated with a loss of parasite richness, which in general was high. Parasite prevalence and the proportion of contaminated samples in which each parasite taxon was present was also high in general and there were no differences between groups. A factor related to the generally high levels of parasitism in our study groups could be the high humidity in the study area, because this favors the survival of parasitic free forms and increases the chances of infection. This would also account for the tendency towards higher levels of parasitism observed in the rainy season. Finally we did not find a pattern relating sex and parasitism.