Dung survey bias and elephant population estimates in southern Mozambique

We used dung surveys to estimate population size and extracted an age structure from boli diameters for the elephants living in the Maputo Elephant Reserve. Our estimate was based on published defecation rates, dung decay rates, distance-sampling techniques and 1,672 dung piles encountered on 204 line-transects. The reserve had at least 311 (95% CI: 198–490) elephants at a density of 0.60 (95% CI: 0.38–0.94) per km². However, observer bias reduced effective strip widths and inflated estimates and their confidence limits. The age structure extrapolated from dung measurements indicated few newborn calves compared with other populations. To detect population changes of 2–5% at 80% power, dung surveys should be carried out every second year for the next 20 years using 100 transects of at least 500 m each. Comparison with a 1995 dung survey suggests that the population is stable and that previous fears of a major population decline during the civil war have no foundation.     

Seasonality in elephant dung decay and implications for censusing and population monitoring in south-western Cameroon

Dung decay rates have been used in the past to correct elephant dung counts to estimates of population numbers. These assume that the dung is in a ‘steady state’ where production rates are equal to disappearance rates. The inter‐seasonal variation in decay rates, however, has never been fully investigated for elephant dung. This study, in the Banyang‐Mbo Wildlife Sanctuary in SW Cameroon, investigated whether a single dung decay rate is really appropriate when converting dung counts to population density estimates. A total of 870 fresh elephant dung‐piles were monitored until they had completely decayed, with at least 60 fresh dung piles located each month of the year between July 1994 to June 1995. The elephant dung decay rate obtained was 55.6% to 66.7% lower than those obtained for the rainforests of Gabon. This has implications for using decay rates obtained elsewhere to compute elephant densities from dung counts. Statistically significant differences were found in the mean duration of dung‐piles between different months and different seasons. Environmental variables, with the exception of rainfall, did not aid in predicting the duration of elephant dung‐piles. The model of Plumptre & Harris was used to estimate the standing crop of dung at any particular time of the year. The results show that, if there was no emigration of elephants from the population, then there was only a three‐month period when the dung was in a ‘steady state’. If emigration occurred during the wet season as is thought to occur then there was no period of ‘steady state’.     

Primary Seed Dispersal by Red Howler Monkeys and the Effect of Defecation Patterns on the Fate of Dispersed Seeds1

The effectiveness of a seed disperser depends on the quantity and quality of dispersal. The quality of dispersal depends in large part on factors that affect the post–dispersal fate of seeds, and yet this aspect of dispersal quality is rarely assessed. In the particular case of seed dispersal through endozoochory, the defecation pattern produced has the potential of affecting the fate of dispersed seeds and consequently, dispersal quality and effectiveness. In this study, I assessed the effects of dung presence and dung/seed densities on seed predation by rodents and secondary dispersal by dung beetles. In particular, I compared seed fates in clumped defecation patterns, as those produced by howler monkeys, with seed fates in scattered defecation patterns, as those produced by other frugivores. I also determined the prevalence of red howler monkeys (Alouatta seniculus) as seed dispersers at the plant community level in Central Amazonia by determining the number of species they dispersed in a 25–month period. I found that dung presence and amount affected rodent and dung beetle behavior. Seed predation rates were higher when dung was present, and when it was in higher densities. The same number of seeds was buried by dung beedes, in dumped versus scattered defecation patterns, but more seeds were buried when they were inside large dung–piles versus small piles. Seed density had no effect on rodent or dung beetle behavior. Results indicate that caution should be taken when categorizing an animal as a high or low quality seed disperser before carefully examining the factors that affect the fate of dispersed seeds. Red howler monkeys dispersed the seeds of 137 species during the study period, which is the highest yet reported number for an Alouatta species, and should thus be considered highly prevalent seed dispersers at the plant community level in Central Amazonian terra firme rain forests.     

Estimating densities of large herbivores in tropical forests: Rigorous evaluation of a dung‐based method

When sighting‐based surveys to estimate population densities of large herbivores in tropical dense forests are not practical or affordable, surveys that rely on animal dung are sometimes used. This study tested one such dung‐based method by deriving population densities from observed dung densities of six large herbivores (chital, elephant, gaur, muntjac, sambar, and wild pig) in two habitats, dry deciduous forests (DDF) and moist deciduous forests (MDF), within Nagarahole National Park, southern India. Using the program DUNGSURV, dung pile counts, decay rates estimated from field experiments, and defecation rates derived from literature were analyzed together by a model that allows for random events affecting dung decay. Densities of chital were the highest, followed by sambar. Wild pig densities were similar in the two habitats, sambar densities were higher in DDF, and densities of the other species were higher in MDF than in DDF. We compared DUNGSURV estimates with densities estimated using distance sampling in the same season. DUNGSURV estimates were substantially higher for all species in both habitats. These differences highlight the challenges that researchers face in computing unbiased estimates of dung decay rates and in relying on defecation rates from literature. Besides the elephant, this study is the first to rigorously test the efficacy of using a dung‐based approach to estimate densities of large herbivore species in Asia, and based on this evaluation, we provide specific recommendations to address issues that require careful consideration before observed dung densities are used to derive animal densities. Our results underline the need for an experimental study of a known population in a fenced reserve to validate the true potential of using dung‐based approaches to estimate population densities.     

Factors Influencing the Survival of Sympatric Gorilla (<Emphasis Type="Italic">Gorilla gorilla gorilla</Emphasis>) and Chimpanzee (<Emphasis Type="Italic">Pan troglodytes troglodytes</Emphasis>) Nests

Accurate and precise surveys of primate abundance provide the basis for understanding species ecology and essential information for conservation assessments. Owing to the elusive nature of wild apes and the vast region of dense forest they inhabit, population estimates of central chimpanzees (Pan troglodytes troglodytes) and western lowland gorillas (Gorilla gorilla gorilla) have largely relied on surveys of their nests. Specific information about the nesting behavior of apes permits the estimation of the number of nests built (nest creation rate). Similarly, information on nest characteristics and environmental factors can be used to estimate the time it takes nests to decay (nest decay rate). Nest creation and decay rates are then used to convert nest density estimates to absolute ape densities. Population estimates that use site-specific estimates of nest creation and decay rates are more accurate and precise. However, it is common practice to generalize these conversion factors across sites because of the additional cost of studies required to gather the information to estimate them. Over a 9-mo study period, we detected and monitored the time to decay of gorilla nests (N = 514) and chimpanzee nests (N = 521) in northern Republic of Congo. We investigated the influence of nest characteristics and environmental factors on nest survivorship and estimated the mean time to nest decay (or equivalently survival) using MARK. Key factors influencing nest decay rate included ape species, forest type, nest height, mean rainfall, nest structure, nest type, and primary aspects of nest construction. Our findings highlight the synergistic effect of behavior and environment on great ape nest degradation, as well as providing practical insights for improving measures to monitor remaining populations of these endangered species.     

Are Dung Beetles Driving Dung-Fly Abundance in Traditional Agricultural Areas in the Amazon?

We evaluated the effects of different land-use systems on the ability of dung beetles to control the population of detritus-feeding flies. We tested the hypotheses that intensification of land use will reduce dung beetles richness, abundance and biomass and, consequently, their dung burial ability, affecting the interaction between dung beetles and flies and reducing its effectiveness as a natural biological control. In the Brazilian Amazon we sampled dung beetles, fly larvae and adults; and recorded the rate of dung removal by dung beetles across a gradient of land-use intensity from primary forest, secondary forest, agroforestry, agriculture to pasture. Our results provide evidence that land-use intensification results in a reduction of the richness, abundance and biomass of dung beetles, and this in turn results in lower rates of dung removal in the most simplified systems. We found no significant differences in the abundance of fly larvae between the different systems of land use. However, the number of adult flies differed significantly between land-use systems, presenting higher abundance in those sites with greater intensity of use (pasture and agriculture) and a lower abundance of adult flies in forested systems (primary and secondary forests, and agroforestry). Information-theoretic model selection based on AICc revealed strong support for the influence of land-use systems, dung removal rates and dung beetle abundance, biomass and richness on adult dung-fly abundance. Our results also reveal that dung beetles are not solely responsible for fly control and that other factors linked to land use are influencing the populations of these detritus-feeding insects.     

Patterns of Niche Partitioning and Alternative Reproductive Strategies in an East African Dung Beetle Assemblage

Sympatric species of coprophagus dung beetles rely on essentially the same resource for provisioning broods, which raises the question of how local species diversity is maintained. Interspecific competition may be mitigated to some extent by large-scale spatial (e.g., habitat type) and temporal (e.g., seasonal) variation in activity. Niche partitioning also occurs at the scale of individual dung pads. We examined the extent to which inter- and intraspecific variation in adult morphology and behavior contribute to spatial and temporal partitioning above and below the dung pad in the dung beetle community of Kibale Forest, Uganda. Excavations in the vicinity of dung pads revealed significant heterogeneity among species and guilds in the lateral distance between tunnels and dung, and in tunnel depth. Patterns of succession and diel activity demonstrated strong temporal dissociations among guilds. In one species, Onthophagus multicornis, we found a bimodal distribution of male horn sizes and a tunneling pattern consistent with alternative reproductive tactics.     

The Effect of Dung and Dispersal on Postdispersal Seed Predation of Attalea phalerata (Arecaceae) by Bruchid Beetles1

Low postdispersal mortality of palm seeds in tapir dung is hypothesized to result from the mechanical barrier provided by dung against bruchid infestation and/or from the distance to adult palms at which seeds are dispersed. We tested these hypotheses by distributing endocarps of Attalea phalerata Mart. ex Spreng. in experimental dung piles in Beni, Bolivia. Predation rates were significantly lower for seeds covered by dung than for exposed or partially covered seeds, but did not differ between seeds placed below and 50 m away from palms. Thus, dung, not short‐distance dispersal, protects seeds against bruchid beetles, and may ultimately promote survival of palm seeds.     

Predation of experimental nests is linked to local population dynamics in a fragmented bird population

Artificial nest experiments (ANEs) are widely used to obtain proxies of natural nest predation for testing a variety of hypotheses, from those dealing with variation in life-history strategies to those assessing the effects of habitat fragmentation on the persistence of bird populations. However, their applicability to real-world scenarios has been criticized owing to the many potential biases in comparing predation rates of artificial and natural nests. Here, we aimed to test the validity of estimates of ANEs using a novel approach. We related predation rates on artificial nests to population viability analyses in a songbird metapopulation as a way of predicting the real impact of predation events on the local populations studied. Predation intensity on artificial nests was negatively related to the species' annual population growth rate in small local populations, whereas the viability of large local populations did not seem to be influenced, even by high nest predation rates. The potential of extrapolation from ANEs to real-world scenarios is discussed, as these results suggest that artificial nest predation estimates may predict demographic processes in small structured populations.     

Effects of Degradation and Replacement of Southern Brazilian Coastal Sandy Vegetation on the Dung Beetles (Coleoptera: Scarabaeidae)

In this paper we address the effects of anthropogenic disturbance and replacement of Brazilian Coastal sandy vegetation (restingas) on dung beetles communities. We sampled dung beetles in the four main vegetative physiognomies of Guriri Island, Espírito Santo State: forest restinga, restinga Clusia, disturbed restinga (from burning events), and pastures. We placed four sets of two pitfall traps (baited with horse and human dung) in four independent areas of each vegetation type, and collected 14,534 individuals of 13 dung beetle species. Neither log₁₀ of individuals nor log₁₀ of species richness were good predictors of restinga disturbance. However, a significant amount of variation in dung beetle abundance and richness could be explained by bait type. Ordination of these sites using hybrid multidimensional scaling revealed a gradient of habitat disturbance from undisturbed restinga samples to pasture. Dung beetle communities along this gradient demonstrated a complete turnover in species composition, from restinga‐specialists to invasive and generalists species respectively. This complete turnover signals the local extirpation of forest‐adapted species in disturbed and converted areas. Only a single dung beetle species in preserved restingas is protected by Brazilian law (Dichotomius schiffleri). Given the extent of the clearing of restinga habitat, the conservation status of dung beetles associated with restinga forest gives cause for concern.     

Dung beetle activity affects rain forest seed bank dynamics and seedling establishment

Dung beetles relocate vertebrate feces under the soil surface, and this behavior has many ecological consequences. In tropical forests, for example, seeds defecated by mammals that are subsequently buried by dung beetles are less likely to suffer predation. While the effects of dung beetles on the fate of defecated seeds have been relatively well studied, their effect on seeds already buried in the soil has not. To contribute to fill this gap, we designed a study with three objectives: (a) Describe the vertical re‐distribution of soil seeds that occurs due to dung beetle activity; (b) Determine if beetle activity favors establishment of seedlings from the soil seed bank; and (c) Determine if the effect of dung beetles is stronger in sites of recurrent mammal defecation. We carried out three complementary field experiments, one with artificial seeds (plastic beads) of three sizes buried at known depths, one with two species of seeds buried at those same depths, and one with the natural soil seed bank in sites of single vs. recurrent defecation. Buried beads were moved by dung beetles along the vertical axis, both upwards (9.5%) and downwards (11.5%); smaller beads were more frequently moved downwards while the contrary occurred for larger beads. Dung beetle activity caused an increase in seedling establishment, both from experimentally buried seeds and from the natural seed bank. Defecation recurrence had no effect on seedling establishment. We conclude that dung beetle activity affects seed bank dynamics with important consequences for seedling establishment in tropical forests.Abstract in Spanish is available with online material.     

Fragmentation and Alteration of Seed Dispersal Processes: An Initial Evaluation of Dung Beetles, Seed Fate, and Seedling Diversity1

Given current accelerated trends of tropical land conversion, forest fragments are being incorporated into many conservation programs. For investing in fragments to be a viable conservation strategy, forest fragments must maintain their ecological integrity over the long term. Based on fieldwork in 22 forest fragments in the crater lakes region of western Uganda and in the continuous forest of Kibale National Park, we examined (1) seed predation on experimentally dispersed seeds, (2) abundance and composition of the dung beetle community that may play a major role in removing seeds from sites of high seed predation, and (3) compared the fragments’ seedling community composition to adult tree community composition and the seedling community in continuous forest. First, the rate of seed removal at experimental stations was lower in forest fragments (85% remaining after 1 day) than at stations in the continuous forest (79% remaining) and the probability of stations being discovered by seed predators was lower in fragments (23%) than in the intact forest (41%). Second, there was a 62 percent decline in fragment dung beetle abundance. The magnitude of this decline varied among dung beetle guilds that process dung and seeds in different fashions. The abundance of large rollers that move large seeds away from sites of defecation did not differ, but medium and smaller rollers and burying beetles that process small and medium‐sized seeds were less common in the fragments than in the intact forest. Finally, we compared the seedling community composition relative to adult tree community composition by identifying all adult trees in each fragment and by sampling the composition of the seedling community. We found some evidence to suggest that there was movement of seeds among forest fragments by large‐bodied dispersers, particularly chimpanzees (Pan troglodytes) and hornbills (Ceratogymna subcylindricus).     

Ape Abundance and Habitat Use in the Goualougo Triangle, Republic of Congo

Chimpanzee (Pan troglodytes troglodytes) and western gorilla (Gorilla gorilla gorilla) populations in central Africa are rapidly declining as a result of disease epidemics, commercial bushmeat hunting, and habitat destruction. Our main objective was to estimate the absolute abundance and habitat utilization of chimpanzees and gorillas in the intact forests of the Goualougo Triangle in the Republic of Congo, and in an adjacent area in which selective logging will take place in the near future. The estimates provide a unique baseline for apes inhabiting an undisturbed environment. A second objective was to compare estimates of abundance and patterns of habitat utilization generated by different techniques: 1) distance sampling of individual ape nests and nest sites along line transects, 2) direct observations of apes during reconnaissance surveys, and 3) observations of ape traces during reconnaissance surveys. We completed a total of 222 km of line transect surveys in 4 sampling areas, resulting in overall density estimates of 1.53 chimpanzees/km² and 2.34 gorillas/km² from nest sites. We generated a density estimate of 2.23 chimpanzees/km² from direct observations during reconnaissance surveys of a semihabituated community in 1 of the 4 sampling areas. Habitat use profiles that nest surveys depicted on transects differed from those of direct observations and traces we encountered on reconnaissance surveys. We found the highest overall abundance of chimpanzee nests in monodominant Gilbertiodendron forest, whereas our direct observations showed that chimpanzees preferred mixed species forest. Transects that traversed the core area of the community range had the highest encounter rates of chimpanzee nests and nest sites. Gorilla nests on transects showed a preference only for mixed species forest with an open canopy, but direct observations and traces on reconnaissance surveys clearly indicated that gorillas use several habitat types. We conclude by evaluating the precision of these nest surveys and our ability to detect future trends in ape densities in the Goualougo Triangle.

Factors Affecting Daily Nest Survival of Burrowing Owls Within Black-Tailed Prairie Dog Colonies

ABSTRACT Identifying environmental parameters that influence probability of nest predation is important for developing and implementing effective management strategies for species of conservation concern. We estimated daily nest survival for a migratory population of burrowing owls (Athene cunicularia) breeding in black-tailed prairie dog (Cynomys ludovicianus) colonies in Wyoming, USA. We compared estimates based on 3 common approaches: apparent nesting success, Mayfield estimates, and a model-based logistic-exposure approach. We also examined whether 8 intrinsic and extrinsic factors affected daily nest survival in burrowing owls. Positive biases in apparent nest survival were low (3–6%), probably because prior knowledge of nest locations and colonial behavior among nesting pairs facilitated discovery of most nests early in the nesting cycle. Daily nest survival increased as the breeding season progressed, was negatively correlated with ambient temperature, was positively correlated with nest-burrow tunnel length, and decreased as the nesting cycle progressed. Environmental features were similar between failed and successful nests based on 95% confidence intervals, but the seasonal midpoint was earlier for failed nests (31 May) compared to successful nests (15 Jun). The large annual variation in nest survival (a 15.3% increase between 2003 and 2004) accentuates the importance of multiyear studies when estimating reproductive parameters and when examining the factors that affect those parameters. Failure to locate and monitor nests throughout the breeding season may yield biased estimates of nesting success in burrowing owls (and possibly other species), and some of the variation in nesting success among years and across study sites may be explained by annual and spatial variation in ambient temperature. Any management actions that result in fewer prairie dogs, shorter burrow lengths, or earlier nesting may adversely affect reproductive success of burrowing owls.     

Estimating brocket deer (<Emphasis Type="Italic">Mazama gouazoubira</Emphasis> and <Emphasis Type="Italic">M. americana</Emphasis>) abundance by dung pellet counts and other indices in seasonal Chiquitano forest habitats of Santa Cruz,Bolivia

We assessed dung and track counts for indexing brocket deer abundance in seasonal habitats on a ranch where hunting is not practiced in the Bolivian lowlands. Surveys were replicated along four 10-km trails (totaling 180 km in the wet season and 90 km in the dry season) through four semideciduous forest habitats. Dung pellets and tracks were identified as belonging to Mazama gouazoubira or M. americana by size and shape. Pellet groups lasted more than 1 month during the dry season, but decayed within 1–2 weeks in the wet season. Mean density estimates based on dung counts varied widely between seasons for gray brockets (from 6.77±4.0 to 30.57±23.5 ind/km²; mean±SD) but not for reds (from 3.52±4.6 to 6.98±7.2 ind/km²). These values were probably too high due to underestimation of daily deposition rate, and were reduced during the wet season because of dung decay. We found consistently more dung in the dry season and more tracks in the wet season. Sightings of red brockets were too few for line-transect analysis (n=6), but those of gray brockets (n=42) produced an overall estimate of 5.6 ind/km² (95% CI=3.5–9.0 ind/km²). Different estimates indicated that gray brockets were more abundant than reds in all situations, except perhaps in the riverine forest. Environmental factors affected these indices of abundance differently, and while we recommend the use of dung counts in dry-season scenarios, we think that index reliability should be assessed locally before conducting population comparisons.     

Trap Spacing and Transect Design for Dung Beetle Biodiversity Studies1

Standardized sampling methods are essential for comparing species diversity and abundance patterns across different studies and sites. Although dung beetles are widely used as a focal taxon in biodiversity studies, nothing appears to be known about the effective sampling area of dung‐baited traps. Mark‐recapture experiments using Canthon acutus showed that at least 50 m between traps should minimize trap interference, and that wind affects trap detectability. Consequently, we propose a standardized dung beetle sampling design.     

Feral horses dung piles as potential invasion windows for alien plant species in natural grasslands

Small scale disturbances could act as patches that provide sites for the colonization of competitively inferior species, promoting the establishment of non-native species in some cases. We analyzed the vegetation associated with feral horse dung piles in montane pampas grasslands in Mid-East Argentina and described the changes following their abandonment, evaluating whether dung piles act as invasion windows, allowing the entrance of alien plant species. We estimated the portion of the study area directly covered by horse manure and dung height was used to estimate the time elapsed after the abandonment of each pile. Vegetation replacement on dung piles of different ages was assessed and compared with grassland controls using discriminant analysis. We used regression analysis to look for changes in vegetation cover, species richness, species diversity and evenness in response to height (age) of the dung piles, and principal component analyses (PCA) to identify groups of plants associated with different successional stages. We compared cover of alien plant species on dung piles with grassland controls using one-way ANOVA. On average, 2.5% of the study area was covered by horse dung. Total vegetation cover, species richness, diversity and evenness increased after the piles were abandoned. Characteristic plant groups were associated with initial, middle and last phases of the studied succession. Vegetation on the dung piles significantly differed from that in grassland controls and two species were consistently associated with dung piles: the invasive Red Star Thistle, Centaurea calcitrapa, and a native grazing-intolerant grass, Nassella clarazii. Non-native species cover was also higher in dung piles than in control plots. Dung piles cover a significant portion of grassland area in our study site, produce significant changes in the vegetation and are associated with some invasive alien plants that could eventually colonize more pristine areas in the vicinity. On the other hand, they might represent refuges for palatable species, since horses seem to avoid them for grazing.     

The Old and Young Amazon: Dung Beetle Biomass, Abundance, and Species Diversity

The Amazon Basin can be divided into two geomorphological regions based on the age of its soils: young (< 30 mya) and old (> 300 mya). We tested the effects of soil age on dung beetle communities by comparing biomass, abundance, and species between reserves in Ecuador on young soils and reserves in Brazil on old soils. Beetle biomass in the old Amazon was one-third that in the young Amazon, and beetle abundance in the old Amazon was one-fourth that in the young Amazon. Species richness, rarefied to equal sample sizes, was not significantly different between old and young soils. These data suggest young soils of the Amazon support a significantly greater biomass and abundance of dung beetles than old soils, but that species richness across the Basin is similar. As dung beetles are bio-indicators of mammals, our data support previous studies indicating a greater biomass of mammals on young versus old Amazon soils.     

Seasonality in the Ecology and Life Histories of Mountain Gorillas (Gorilla gorilla beringei)

The abundance of food, especially that of fruit and often that of young leaves, varies considerably over time for most primates. This variation can depend on or be independent of seasonality in rainfall. Mountain gorillas (Gorilla gorilla beringei) in the Virungas are exceptional: their habitat contains almost no edible fruit, and they mostly eat perennially available herbs and vines that are densely and evenly distributed in much of their habitat. Earlier studies documented little consistent temporal variation in mountain gorilla diets and habitat use, except for seasonal use of bamboo by some groups, and documented no birth seasonality. Long-term data ( 7 years) on 6 mountain gorilla groups confirm these results for habitat use, except that they show some seasonality in use of the upper altitudinal extremes of the gorillas' home ranges for unclear reasons. Relatively low and inconsistent variation in habitat quality over time should lower the costs of grouping for gorillas compared to other apes. Long-term data also confirm the absence of seasonality in births and conceptions. However, they show that mortality rates and risk of respiratory infections vary directly with rainfall. These relationships are probably causal and may be mediated through thermoregulatory stress.     

Dung fungi as a proxy for megaherbivores: opportunities and limitations for archaeological applications

The use of spores of coprophilous fungi from sedimentary sequences as proxy evidence for large herbivore abundance has garnered pronounced attention and scrutiny over the past three decades. In response to the rapid rate at which new information is being discovered on this topic, this paper presents a brief review of the archaeological applications so far, and outlines opportunities and limitations of using Sporormiella as a proxy for herbivore abundance. Specific archaeological uses of this proxy include understanding megaherbivore extinctions and human land use patterns such as pastoralism and agriculture. We analyse how dung fungal records are formed and review the mycological literature to outline factors affecting spore reproduction and preservation. These include how strongly each commonly used dung fungal taxon relies on dung as a substrate and environmental factors affecting dung fungal reproduction and coprophilous fungi deposition. Certain laboratory preparation techniques adversely affect spore representation on pollen slides. The methods of analysis and quantification of spore records also impact our understanding. We describe good practice to increase precision of analytical methods. Due to limitations imposed by some of these factors, it is possible that an absence of dung fungi from a palaeoecological record does not imply an absence of herbivores. However, consideration of these factors and inclusion of as wide a range of coprophilous spore records as possible increases the reliability of such inferences.