Influence of digestive morphology on resource partitioning in Amazonian ungulates

Summary Resource partitioning of diet and habitat use was studied in the entire Amazonian ungulate community of Northeastern Peru, which comprises the red brocket deer (Mazama americana), grey brocket deer (M. gouazoubira), collared peccary (Tayassu tajacu), white-lipped peccary (T. pecari), and lowland tapir (Tapirus terrestris). Each ungulate species partitioned at least one type of resource from every other species. Digestive morphology had a greater influence on resource partitioning of diet than body size. Neither digestive morphology nor body size were related to segregation of habitats. However, species with similar diets partitioned habitats, whereas species with different diets often used the same type of forest. Increases in habitat breadth of ungulates were positively correlated with increases in dietary breadth.     

Ungulate relative abundance in hunted and non-hunted sites in Calakmul Forest (Southern Mexico)

The relative abundance of an ungulate community in hunted and non-hunted areas was studied in Calakmul, a tropical forest in southern Mexico that includes a 723,815 ha Biosphere Reserve where no hunting is allowed and communal lands where hunting activity does take place. Tapir, white-lipped peccary (WLP), collared peccary, two species of brocket deer and white-tailed deer were the focal species. Relative abundance of ungulates was obtained by counting tracks along transects in three hunted sites and in the southern core area of the Calakmul Biosphere Reserve, a enforced part of the protected area of 350,000 ha where no hunting has been-permitted since 1989. A total of 1708 ungulates tracks were obtained along 206.1 km (n = 90 transects) divided among the four sites. Relative abundance of the species was estimated from the track encounter rate (TER) calculated as the number of tracks per species encountered per km of transect. No significant differences in brocket deer and collared peccary TER was found among hunted and non-hunted areas. TER for WLP was significant higher in the non-hunted area, while TER for white-tailed deer and tapir was significant higher in hunted sites. Hunting activity affected the ungulate species in Calakmul differently. Brocket deer, white-tailed deer and collared peccary are more resistant species toward the hunting pressure than the WLP, who needs an immediate protection plan throughout Calakmul. Hunting areas are important habitat refuges for tapir populations and they should be protected in these areas.

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Resumen La abundancia relativa de seis especies de ungulados fue obtenida a través de conteos de huellas a lo largo de transectos en tres sitios con cacería y en la parte sur de la Reserva de la Biosfera de Calakmul, una área de alrededor de 350,000 ha donde la cacería ha sido prohibida desde 1989. Tapir, pecarí de labios blancos, pecarí de collar, dos especies de venados temazates y el venado cola blanca fueron las especies estudiadas. Un total de 1708 huellas de ungulados fueron obtenidos a lo largo de 206.1 km (90 transectos) divididos en los cuatro sitios. La abundancia relativa de las especies fue estimada por medio de la tasa de encuentro de huellas (TEH) calculado como el número de huellas encontradas por km de transecto recorrido. La TEH de los venados temazates y el pecarí de collar fue similar entre las áreas con cacería y sin cacería. La TEH para el pecarí de labios blancos fue significativamente mayor en el área sin cacería, mientras que la TEH del venado cola blanca y el tapir fueron significativamente mayor en los sitios con cacería. La actividad de cacería afecta a las especies de ungulados diferente en la región de Calakmul. Los venados temazates, el venado cola blanca y el pecarí de collar están resistiendo mejor la presión de cacería que el pecarí de labios blancos quien necesita un plan inmediato de protección en toda el área. La evidencia sugiere que las áreas con cacería en este estudio son importantes para las poblaciones de tapir, y que se debe prestar atención a su protección en dichas áreas.

     

Ratite and ungulate preferences for woody New Zealand plants: influence of chemical and physical traits

Ratites (ostriches Struthio camelus) and ungulates (red deer Cervus elaphus scoticus and goats Capra hircus) were presented with 14 indigenous shrubs in cafeteria-style trials. The shrubs represented the spectrum of woody plant architecture, ranging from broadleaved monopodial species through to small-leaved highly branched divaricates. Trials were stopped when almost all shoots of the plant expected to be most preferred had been consumed. There were considerable differences between the herbivores in their selection of certain plant species. Aristotelia fruticosa was avoided by deer, neutrally selected by goats, and preferred by ostriches. All herbivores strongly avoided two species, Pseudopanax crassifolius and Coprosma rugosa. Analysis of relative offtake (proportion of biomass consumed from each species, relative to total biomass consumed) showed that all three herbivores ate less of species with small leaves. Consumption by all herbivores was reduced by structural plant traits (i.e. divarication and related attributes) more than by chemical plant traits. The immediate impact of browsing on the plants, measured as the rank of proportion of shoots eaten, was broadly similar across the herbivores. The broadleaved species (e.g. Myrsine australis, Aristotelia serrata) experienced the greatest browsing, while divaricate (e.g. Coprosma rugosa) and conifer species (e.g. Podocarpus hallii) were generally least browsed. Although cafeteria-style experiments have limitations, our results for deer broadly correspond to those of field-based diet preference studies.     

Frugivory in Amazonian Artiodactyla: evidence for the evolution of the ruminant stomach

Red and grey brocket deer and collared and white-lipped peccary of the Peruvian Amazon are frugivores and consume many types of seeds. The ruminant stomach of brocket deer functions as a mechanism to digest the abundant hard palm seeds of Iriartea sp., Euterpe sp. and Mauritia flexuosa . The sympatric peccaries also consume hard palm seeds; however, peccaries crack these palms by using their strong jaws, thick skull bones and interlocking canines. The ruminant stomach might have evolved as a means to digest structural components of seeds, similar to that employed by extant brocket deer, since ancestral ruminants appear to have evolved as smallbodied, forest-dwelling frugivores.     

Carrying Capacity and Potential Production of Ungulates for Human Use in a Mexican Tropical Dry Forest

Data are provided on the carrying capacity and potential production for sustainable human use of white-tailed deer ( Odocoileus virginianus ) and collared peccary ( Pecari tajacu ) in a protected tropical dry forest at Chamela on the Pacific coast of Mexico. In this paper, the carrying capacity was defined as the equilibrium density plus the number of animals removed by predators. The equilibrium point was estimated from the density dependent relationship between the finite population growth rate and the current density according to a logistic model. Annual density was estimated using the line transect method. Carrying capacity estimates were 16.5 to 17.2 deer/km² and 9.3–9.5 peccaries/km², representing a combined biomass of 841–874 kg/km². A potential production for human use of 2.1 deer/km² and 4.4 peccaries/km² was estimated employing the model of Robinson and Redford (1991) . The data suggest that, in the protected tropical dry forest of Chamela, the density and biomass of wild ungulates can maintain a similar or greater density and biomass than other Neotropical forests. To obtain an accurate estimation of the maximum sustainable yield ( MSY ), it is necessary to consider predation. From a management point of view, it is important to consider that carrying capacity varies as a function of the rainfall pattern.     

Distance-dependence in two Amazonian palms: effects of spatial and temporal variation in seed predator communities

Animals aid population growth and fitness in tropical forest communities through dispersal and negatively impact populations through seed predation. The interaction between dispersal and seed predation can produce distance- or density-dependence; powerful mechanisms for maintaining species diversity incorporated in the Janzen–Connell model. Large mammals, the highest biomass seed predators of intact Amazonian communities and at risk due to human disturbance, are potentially central to these interactions. This study tests the Janzen–Connell model and investigates the impact of mammalian seed predators on seedling recruitment and maintenance of tree diversity. Patterns of both vertebrate and invertebrate seed predation and seedling recruitment were studied in the two most abundant canopy tree species in western Amazonia (Arecaceae: Astrocaryum murumuru and Iriartea deltoidea). We specifically examined effects of both spatial and temporal variation of the highest biomass seed predator in southwest Amazonian forests, the white-lipped peccary (Tayassu pecari), on recruitment through disturbed and undisturbed sites and through a fortuitous 12 year natural extinction and recolonization event of T. pecari. Distance-dependent seedling recruitment was found in Astrocaryum and Iriartea at both sites. However, the median distance of seedlings was ~1.5× farther from reproductive adults in both palms at the undisturbed site. The number of Iriartea seeds escaping predation increased 6,000% in both space and time due to the decline of T. pecari abundance. The results demonstrate that Janzen–Connell effects are stronger in intact ecosystems and tie these mechanistically to changes in seed predator abundance. This study shows that anthropogenic changes in mammal communities decrease the magnitude of Janzen–Connell effects in Amazonian forests and may result in decreases in tree diversity.     

Interactive effects of insects and ungulates on root growth in a native grassland

Insects and ungulates co‐occur in grasslands, often feeding on the same plants at the same time and potentially having interactive effects on plant growth. Further, ungulate–insect interactions may differ between native ungulate guilds and domesticated cattle. Despite the prevalence of insects and ungulates in native grasslands, experiments simultaneously manipulating the densities of both these groups are rare. Using large, replicated paddocks, as well as insecticide application, we restricted access to vegetation by each group of herbivores. We also manipulated the species identity of the ungulate assemblage, allowing us to determine whether there are differential effects between native ungulate guilds (bison, elk and deer) and cattle on plant biomass. We found interactive effects of insect and ungulate herbivores on root growth. When insects were suppressed, both native ungulates and cattle caused an approximate doubling of root biomass. However, this stimulatory effect of ungulate grazing was eliminated when insects were also present. In contrast, neither insects nor ungulates had significant effects on shoot biomass at these densities. As a result, the dominant effects of above‐ground herbivory was on belowground plant growth. We suggest the effects of insect and ungulate assemblages on root biomass appear important in regulating primary production in this grassland and may account for some of the contradictory plant responses to ungulate herbivory in the literature.     

Reproductive biology of female Amazonian brocket deer in northeastern Peru

The aim of this study was to provide information on the reproductive biology of brocket deer. Hence, we analyzed female reproductive tracts collected by rural hunters from 1991 to 1998 in the Tamshiyacu-Tahuayo Communal Reserve, northeastern Peruvian Amazon. We characterized the basic reproductive biology of brocket deer, analyzed whether the distributions of conceptions and births are aseasonal, and compared their reproductive productivity in two different areas subject to heavy and slight hunting pressures, respectively. We found that: (1) red and gray brocket deer did not differ in ovulation, fertilization, and pregnancy rates; (2) average number of fetuses per birth was 1.2 for red brocket deer and one for gray brocket deer; (3) sex of fetuses suggests a male biased sex ratio for both species; (4) neither species shows reproductive seasonality; and (5) gross productivity does not differ between heavily and slightly hunted areas. Our results indicate that brocket deer exhibit reproductive characteristics similar to their conspecifics in other parts of their native distribution range.     

Spatial Niche Partitioning in Sub-Tropical Solitary Ungulates: Four-Horned Antelope and Barking Deer in Nepal

Differential resource use allows a diversity of species to co-exist in a particular area by specializing in individual ecological niches. Four-horned antelope Tetracerus quadricornis is endemic to the Indian subcontinent and has a restricted distribution in Nepal and India; however, the barking deer Muntiacus vaginalis is relatively common throughout its wide distribution range. We wanted a better understanding of their habitats and how these two similarly sized solitary ungulates manage to coexist in lowland Nepal. We used fecal pellet belt transect surveys in the Babai valley, Bardia National Park to study the habitat associations of both species. We found empirical evidence that four-horned antelope prefer hill sal forest and deciduous hill forest at higher elevations, whereas barking deer preferred riverine and sal forest in lower elevations. We found a clear niche differentiation of four-horned antelope and barking deer that made the coexistence of these similarly sized solitary ungulates possible. Hence, resource partitioning is the key to coexistence of these solitary ungulates, and the fine-grained habitat mosaic of different forest types in the study landscape appears to be the underlying feature. Therefore, maintaining the habitat mosaic and preserving valuable hill sal and deciduous hill forests will facilitate the coexistence of herbivores in sub-tropical regions.     

Effects of red deer on tree regeneration and growth in Aorangi Forest, Wairarapa

New Zealand forests have been substantially modified by introduced red deer over the past century. New Zealand’s indigenous forest managers need to know if regeneration of palatable tree species can be restored following control or eradication of browsing ungulates. Aorangi Forest, Wairarapa, suffered dramatic changes in forest understorey composition by the 1950s after more than seven decades of colonisation by red deer (Cervus elaphus), feral goats (Capra hircus) and pigs (Sus scrofa). Since then feral goats have been eradicated, but red deer and pigs persist throughout Aorangi Forest despite ongoing recreational hunting. This study uses data from paired fenced and unfenced plots established at seven sites in Aorangi Forest between 1981 and 1987, and re-measured in 2004, to show the effects of ungulates on tree (≥ 2 cm diameter at breast height) regeneration and growth. Our results show that browsing by red deer has prevented regeneration of kanono (Coprosma grandifolia), a highly palatable, fast-growing sub-canopy hardwood tree. Deer reduced the growth of mahoe (Melicytus ramiflorus) trees, probably by directly browsing epicormic shoots. The regeneration of other less palatable sub-canopy trees (e.g. porokaiwhiri, Hedycarya arborea), and slower-growing canopy species (e.g. hinau, Elaeocarpus dentatus and rewarewa, Knightia excelsa) appears to have been unaffected by deer browsing. Accordingly, tree species composition does not appear to have been greatly affected by browsing from deer populations in Aorangi Forest over the past two decades.     

Opposing deer and caterpillar foraging preferences may prevent reductions in songbird prey biomass in historically overbrowsed forests

Overbrowsing by ungulates decimates plant populations and reduces diversity in a variety of ecosystems, but the mechanisms by which changes to plant community composition influence other trophic levels are poorly understood. In addition to removal of avian nesting habitat, browsing is hypothesized to reduce bird density and diversity through reduction of insect prey on browse‐tolerant hosts left behind by deer. In this study, we excluded birds from branches of six tree species to quantify differences in songbird prey removal across trees that vary in deer browse preference. Early in the breeding season, birds preyed on caterpillars at levels proportional to their abundance on each host. Combining these data with tree species composition data from stands exposed to experimentally controlled deer densities over 30 years ago, we tested whether overbrowsing by white‐tailed deer reduces prey biomass long after deer densities are reduced. Our analysis predicts total prey availability in the canopy of regenerating forests is fairly robust to historic exposure to high deer densities, though distribution of prey available from host species changes dramatically. This predicted compensatory effect was unexpected and is driven by high prey abundance on a single host tree species avoided by browsing deer, Prunus serotina. Thus, while we confirm that prey abundance on host trees can act as a reliable predictor for relative prey availability, this study shows that quantifying prey abundance across host trees is essential to understanding how changes in tree species composition interact with ungulate browse preference to determine prey availability for songbirds.     

The Value of Measuring Food Availability on the Ground for a Semiterrestrial Frugivore, the Tana River Mangabey (Cercocebus galeritus) of Kenya

Collecting phenological data, seasonal availability of plant resources that primates feed on, allows us to understand feeding ecology better. A number of primates are terrestrial or semiterrestrial frugivores, yet phenology is generally measured only in the canopy. I hypothesized that combining measurements of food availability on the ground with canopy measurements would more strongly correlate with diet than canopy measurements alone for a semiterrestrial frugivore, the Tana River mangabey (Cercocebus galeritus). From July 2005 until June 2006, I conducted monthly follows on a mangabey group. Phenology was measured in 105 individuals of their top seven food tree species. I measured canopy phenology on a 0–5 scale and counted fruits in three 1 m³ areas of the canopy, and measured ground phenology by counting fruits and seeds in four 1 m² quadrats under the canopy. I calculated each tree’s canopy volume and canopy shadow, and each species’ mean fruit weight, mean seed weight, and density. Monthly biomasses were calculated as kilograms per hectare. Spearman correlations were performed between diet contribution and canopy biomass, ground biomass, and total biomass. The hypothesis was not supported for seven species individually or combined. The hypothesis was supported for 3 of 12 diet items, although canopy biomass was also significant for 2 of those items. Two diet items correlated only with ground biomass. Studies of the Tana River mangabey may benefit from measuring ground phenology only for those items eaten exclusively on the ground. Primatologists studying terrestrial or semiterrestrial frugivores should consider feeding height when deciding on phenology methods.     

The role of forest structure and human occupation in structuring mammal assemblages in oligotrophic ecosystems of Central Amazonia

Large mammals are vulnerable to extinction, and respond directly to ecological gradients within the forest and to the intensity of forest product use by humans. In this study, we evaluated the effects of differences in forest structure and human occupation history on the composition of medium and large‐sized mammal assemblages of the terra‐firme forests of the Anavilhanas National Park, one of the most oligotrophic Amazonian ecosystems. Mammal surveys were conducted along 11 linear transects of 4 km, six of which were located in areas once inhabited by people and managed until the year that the park was created (over 30 years ago), and five in areas that were not inhabited at that time. We detected 469 individuals of 26 species during diurnal and nocturnal sampling, and 11 additional species outside transects. Human occupation history was strongly related to forest structure components. Fruit biomass, canopy cover and tree size were strongly associated with uninhabited areas, and influenced the structure of mammal assemblages. A direct relationship between diet category and species size was observed. Large frugivore‐herbivores and carnivores were more closely associated with areas with more fruit, larger trees and greater canopy cover. In contrast, small arboreal frugivore‐omnivores associated more closely with open canopy and smaller trees. Our study indicated that the effects of human occupation history on forest structure are still evident three decades after the removal of local people from the park. This long‐term effect can be explained by the low resilience of the Anavilhanas environments, demonstrating the fragility of mammal assemblages in face of anthropogenic variation in forest structure in the oligotrophic ecosystems of the Negro River basin.     

Assessing Local Knowledge of Game Abundance and Persistence of Hunting Livelihoods in the Bolivian Amazon Using Consensus Analysis

We used cultural consensus models to test whether hunters shared perceptions of wildlife abundance and the relative importance of hunting and fishing in a Guarayo indigenous community in the Bolivian Amazon. Results show that highly prized animals that were considered rare are either those with lower reproductive rates and more sensitivity to land use changes and harvest (white-lipped peccary and spider monkey) or those with marked seasonal distribution patterns (barred sorubim and tiger-fish). Rapidly reproducing and resilient species (agouti and armored catfish) were perceived as abundant. More tapirs and red brocket deer were present than predicted by scientific models possibly because hunters were harvesting these species in new forest management areas. Residents identified hunting and fishing among their most important livelihood activities, recognized bush meat and fish as basic food resources, and expected wildlife harvests to be part of their future livelihoods, although market-based livelihoods and domestic replacements for bush meat were reported.     

Patterns of wolfCanis lupus predation on wild and domestic ungulates in the Western Carpathian Mountains (S Poland)

We studied the predator-prey relationships among wolvesCanis lupus Linnaeus, 1758, wild ungulates, and livestock in the managed mountain forests of the Western Carpathians (S Poland). Though roe deerCapreolus capreolus dominated in the community of wild ungulates and livestock was abundant within the study area, the three wolf packs preyed mainly on red deerCervus elaphus (42% of food biomass), and next on the roe deer (33%). In both species of deer, wolves preferred killing females and juveniles more frequently than expected from their respective shares in the populations. Wild boarSus scrofa made up 4% of the food biomass, in accordance with its low share in the ungulates community. Despite the easy access of wolves to numerous unprotected sheep flocks pastured on meadows among woods, livestock constituted only 3% of the wolf food biomass. Wolves preyed mostly on sheep (88%), killing on average 34 per year. Most cases of livestock depredation occurred in August –September, on pastures located most often >50 m apart from buildings. Usually, lack of proper guarding was conducive to wolf attacks.     

The Seedling Ratio Method for determining ungulate impacts on forest understoreys: Utility in an Australian ecosystem

The Seedling Ratio Method was devised to assess the impacts of introduced ungulates on plant species richness in forest understoreys. The method has successfully assessed ungulate impacts on species richness in forests in New Zealand and Hawaii, which do not have native herbivores. We tested the three critical assumptions that underlie this method to investigate its potential for use in an Australian ecosystem. This study was conducted in the Yarra Ranges National Park (YRNP), Victoria, which had a high‐density population of an introduced deer species, Sambar (Cervus unicolor), in addition to several native herbivore species. One of the three key assumptions of the Seedling Ratio Method was not supported, whereas conventional differential exclosures showed clear and separate impacts by Sambar and native herbivores. We conclude that the Seedling Ratio Method could not provide a clear indication of browsing impact on forest understoreys in YRNP.     

An experimental study of the impacts of understorey forest vegetation and herbivory by red deer and rodents on seedling establishment and species composition in Waitutu Forest, New Zealand

Introduced mammalian herbivores are changing the structure and composition of New Zealand’s forest ecosystems and may modify forest succession after natural disturbances. We studied how introduced ungulates (red deer and feral pigs) and rodents (rats and house mice) affected the rate of recovery (i.e. the engineering resilience) of the forest understorey following artificial disturbance. We imposed disturbances by clearing understorey vegetation dominated by Blechnum ferns in forests on relatively fertile alluvium and elevated infertile marine terraces, and recorded recovery of vegetation (seedling establishment, species composition, cover and volume) in herbivore exclosures and controls. Seedlings quickly established on cleared plots: after 2 years, numbers of woody seedlings and ground cover of vascular plants relative to initial values were similar on cleared and uncleared treatments. Volume of plant biomass <2 m remained low on cleared subplots. Ungulates significantly reduced the re-establishment of woody seedlings ≥ 10 cm tall: only one seedling reached this height outside exclosures, compared with 29 seedlings inside. The number of seedlings <10 cm tall, expressed relative to numbers present pre-clearing, was not significantly affected by ungulates. The species composition of regenerating vegetation was more similar (Jaccard index) to pre-clearing understorey vegetation inside ungulate exclosures than outside. No consistent effect of rodents (primarily house mice) on seedling establishment or species composition was detected after 2 years, and rodent exclosures did not significantly affect survival of seedlings (Griselinia littoralis and Aristotelia serrata) planted as an index of rodent herbivory pressure. No significant differences in vegetation recovery were apparent between forest types. Rapid seedling recruitment in the absence of understorey vegetation and the presence of herbivores provided evidence that understorey vegetation competes with seedlings for light. Ungulate effects were consistent with other experiments that showed herbivores reduced the rate and altered the trajectory of vegetation regrowth after disturbance.     

Ungulate browsers promote herbaceous layer diversity in logged temperate forests

Ungulates are leading drivers of plant communities worldwide, with impacts linked to animal density, disturbance and vegetation structure, and site productivity. Many ecosystems have more than one ungulate species; however, few studies have specifically examined the combined effects of two or more species on plant communities. We examined the extent to which two ungulate browsers (moose [Alces americanus]) and white‐tailed deer [Odocoileus virginianus]) have additive (compounding) or compensatory (opposing) effects on herbaceous layer composition and diversity, 5–6 years after timber harvest in Massachusetts, USA. We established three combinations of ungulates using two types of fenced exclosures – none (full exclosure), deer (partial exclosure), and deer + moose (control) in six replicated blocks. Species composition diverged among browser treatments, and changes were generally additive. Plant assemblages characteristic of closed canopy forests were less abundant and assemblages characteristic of open/disturbed habitats were more abundant in deer + moose plots compared with ungulate excluded areas. Browsing by deer + moose resulted in greater herbaceous species richness at the plot scale (169 m²) and greater woody species richness at the subplot scale (1 m²) than ungulate exclusion and deer alone. Browsing by deer + moose resulted in strong changes to the composition, structure, and diversity of forest herbaceous layers, relative to areas free of ungulates and areas browed by white‐tailed deer alone. Our results provide evidence that moderate browsing in forest openings can promote both herbaceous and woody plant diversity. These results are consistent with the classic grazing‐species richness curve, but have rarely been documented in forests.     

Habitat associations of four ungulates in mountain forests of southwest China, based on camera trapping and dung counts data

The present study aimed to assess abundance indices and habitat associations of four sympatric ungulate species (alpine musk deer Moschus chrysogaster, tufted deer Elaphodus cephalophus, Chinese serow Capricornis milneedwardsii, and Chinese goral Naemorhedus griseus) in Baima Xueshan Nature Reserve of southwest China, using camera trapping and dung counts data. Camera traps were set along six dung transects in different habitats and explored habitat use of the sympatric ungulates using trapping rates. The results obtained revealed that Chinese serow showed a negative association with open canopy cover and low canopy cover. Alpine musk deer were associated with oak shrubs, oak forests and open canopy cover, while tufted deer avoided oak shrubs. Goral showed no significant associations with habitat variables. Alpine musk deer and tufted deer had considerable habitat overlap with Chinese serow. By finding a high correlation between indices, the study indicates that camera trapping may represent a valid index of relative abundance, matching results from other studies.     

Insect herbivory on mangrove leaves in North Queensland*

Estimates of leaf damage by insect herbivores are presented for 25 species of mangrove plants, comprising canopy and understorey species. Leaf area loss was highly variable among the species sampled, with means ranging from 0.3 to 35.0% of expanded leaf area. There was also great variability amongst leaves within species, and the mean coefficient of variation for leaf loss from the 25 species was 266%. Of the 12 species sampled at more than one site in North Queensland, eight exhibited small, but significant, between-site differences in herbivory. In general, it did not appear that height in the canopy influenced herbivory. For the dominant mangrove forest community type at Missionary Bay, an estimated mean of 2.1% of leaf production, or 11 g m^-2 per year, entered the direct grazing pathway. This very low figure is compared with estimates from other studies on mangrove forests and estimates from a variety of Australian terrestrial forests.