Inventory and Bioindicator Sampling: Testing Pitfall and Winkler Methods with Ants in a South African Savanna

The sampling efficiency and consistency of pitfall traps and Winkler samples for inventory, bioindicator and ecological studies in savanna habitats was compared using ants. Pitfall traps are often used for ant collecting while Winkler litter sampling has until now had rather limited use. We test Winkler sampling for the first time in a South African savanna. Pitfall traps were more efficient and productive than Winkler sampling for epigaeic ants, with a greater total species richness and higher abundance of ants recorded. Winkler samples contributed few additional species. The relative abundance of different sized ants was different with the two collection methods. Winkler sampling was found to catch greater numbers of smaller ants than pitfall trapping, whereas pitfall trapping caught more larger ants. The standard collecting Winkler quadrat size of 1m² did not perform as well as 2× m² quadrats combined for one sample.     

A comparison of pitfall traps with bait traps for studying leaf litter ant communities.

A comparison of pitfall traps with bait traps for sampling leaf litter ants was studied in oak-dominated mixed forests during 1995-1997. A total of 31,732 ants were collected from pitfall traps and 54,694 ants were collected from bait traps. They belonged to four subfamilies, 17 genera, and 32 species. Bait traps caught 29 species, whereas pitfall traps caught 31 species. Bait traps attracted one species not found in pitfall traps, but missed three of the species collected with pitfall traps. Collections from the two sampling methods showed differences in species richness, relative abundance, diversity, and species accumulation curves. Pitfall traps caught significantly more ant species per plot than did bait traps. The ant species diversity obtained from pitfall traps was higher than that from bait traps. Bait traps took a much longer time to complete an estimate of species richness than did pitfall traps. Little information was added to pitfall trapping results by the bait trapping method. The results suggested that the pitfall trapping method is superior to the bait trapping method for leaf litter ant studies. Species accumulation curves showed that sampling of 2,192+/-532 ants from six plots by pitfall traps provided a good estimation of ant species richness under the conditions of this study.     

Tropical ant communities are in long-term equilibrium

Communities change with time. Studying long-term change in community structure permits deeper understanding of community dynamics, and allows us to forecast community responses to perturbations at local (e.g. fire, secondary succession) and global (e.g. desertification, global warming) spatial scales. Monitoring efforts exploring the temporal dynamics of indicator taxa are therefore a critical part of conservation agendas. Here, the temporal dynamics of the Otongachi leaf litter ant community, occurring in a cloud forest in coastal Ecuador, were explored. By sampling this community six times over eleven years, I assessed how the ant fauna caught by Winkler traps (more diverse and cryptic fauna) and caught by pitfall traps (larger, more mobile fauna) changed over time. The Otongachi leaf litter ant community was dynamic. Although species richness in the community remained constant, temporal turnover of species was high: on average, 51% of the ant species in Winkler traps, and 56% of those in pitfall traps, were replaced with other ant species from one year to the other. Shifts in the rank abundance of species in the community were also large across the eleven years and, on average, shifts in the rank abundance of species collected by Winkler traps doubled those occurring in pitfall traps from one census to the other. In spite of these trends, the Otongachi ant fauna showed no (Winkler) or weak (pitfall) evidence of directional change (towards a new community). Thus, this tropical ant community can be divided in two community compartments. The Winkler compartment composed by a more diverse and cryptic ant fauna appears to be resilient and stable in time. The pitfall compartment composed by larger and more mobile ants may be prone to respond to disturbance. This study suggests that 1) species appearing/disappearing from a site may be rather the rule, difficult to separate from responses to ecological stress. 2) Conclusions made in short-term studies, or studies comparing two (e.g. before and after) snapshots of a community, should thus be revisited. Finally, 3) the ant fauna caught by pitfall traps (a rather simple and cheap survey method) is the most likely community compartment to indicate ecological perturbation. This study adds to the growing evidence that using ants as ecological indicators should incorporate long-term temporal dynamics.     

How to evaluate and reduce sampling effort for ants

Representative sampling of ant communities is time-consuming and laborious. Due to their manifold habitat requirements and sociobiological attributes, a suite of sampling methods is necessary to detect nearly all ant species in a given habitat. We analysed how well different combinations and intensities of sampling methods are suited to assess ant species richness and community patterns. Sampling occurred at 24 sites in mountain and floodplain habitats in Austria using pitfall traps, Winkler litter extraction, hand sampling of foragers, and colony sampling. Of these, pitfall traps delivered the largest species numbers. We drew 19 subsets of sampling approaches and compared these to the complete data. Many subsets that allowed for substantial reduction of work effort nonetheless reflected well the beta diversity patterns of the whole dataset. In contrast, alpha diversity assessment was more sensitive to reduced sampling effort. Pitfall traps turned out to be indispensable to collect ant species for biodiversity studies, but the optimal combination of sampling methods varied between habitat types. In rugged montane habitats pitfalls should be complemented at least by colony sampling. In floodplain forests the number of pitfall trap replicates can be reduced, but inclusion of Winkler samples is advisable.     

Ant Diversity in Dominant Vegetation Types of Southern Cameroon

Ants have been shown as particularly affected by land disturbance through deforestation and conversion of forest to agriculture. The effect of land use change on ant diversity in the Congo Basin is not well known. We conducted intensive sampling along a gradient of increasing vegetation disturbance to test the effect of habitat disturbance on ant diversity and Functional Groups composition. Sampling was conducted in 30 plots (5 study sites × 3 habitat × 2 plots/habitat), replicated six times in 1 year. In each plot, ants were monitored with pitfall traps, quadrats and baits. We recorded 237 ant morphospecies grouped in 10 subfamilies and 43 genera. Myrmicaria opaciventris was the most abundant species followed by Anoplolepis tenella. Forest had greater ant diversity compared with fallows and mixed‐crop fields. Functional groups were dominated by Opportunists, followed by Omnivorous Arboreal Dominants and Generalized Mymicinae. Their composition was not affected by the disturbance, but occurrence of Specialist Predators decreased with increasing disturbance. Occurrence of Generalized Myrmicinae, Opportunists and Subordinate Camponotini increased with disturbance. These results indicate that forest conversion into mixed‐crop fields reduce ant diversity. It can also increase abundance of species with generalized diet that predominates where stress and disturbance limits other ants.     

What is the effect of soil use on ant communities?

Studies on ant communities in agroecosystems have contributed to the knowledge of the effect of agricultural activities on biological communities. The aim of this study is to explain the effect of soil use on ant communities. We tested the hypothesis that there was a decrease in ant species richness and a change in the species composition at habitats with more intense soil use. We collected ants using sardine baits, subterranean traps and direct sampling at four habitats with different soil use (secundary forest, Acacia forestry, initial stage of succession and mixed crops). The ant species richness did not decrease with intensity of soil use. In successional habitat the species numbers collected using sardine baits and subterranean traps were significantly different. Species composition of communities had a pronounced variation, with the epigaeic and hypogaeic ant faunas of the habitat with high intense soil use (mixed crops) had low similarity with ant communities of the three other habitats. The predator species were restricted to habitats with low intensity of soil use. Then, species composition could better reflect the functional changes on ant communities than species richness. Our data can help to choose the component of ant community that better reflect the response of biodiversity to agricultural impacts.     

Rapid inventory of the ant assemblage in a temperate hardwood forest: species composition and assessment of sampling methods

Ants are key indicators of ecological change, but few studies have investigated how ant assemblages respond to dramatic changes in vegetation structure in temperate forests. Pests and pathogens are causing widespread loss of dominant canopy tree species; ant species composition and abundance may be very sensitive to such losses. Before the experimental removal of red oak trees to simulate effects of sudden oak death and examine the long-term impact of oak loss at the Black Rock Forest (Cornwall, NY), we carried out a rapid assessment of the ant assemblage in a 10-ha experimental area. We also determined the efficacy in a northern temperate forest of five different collecting methods--pitfall traps, litter samples, tuna fish and cookie baits, and hand collection--routinely used to sample ants in tropical systems. A total of 33 species in 14 genera were collected and identified; the myrmecines, Aphaenogaster rudis and Myrmica punctiventris, and the formicine Formica neogagates were the most common and abundant species encountered. Ninety-four percent (31 of 33) of the species were collected by litter sampling and structured hand sampling together, and we conclude that, in combination, these two methods are sufficient to assess species richness and composition of ant assemblages in northern temperate forests. Using new, unbiased estimators, we project that 38-58 ant species are likely to occur at Black Rock Forest. Loss of oak from these forests may favor Camponotus species that nest in decomposing wood and open habitat specialists in the genus Lasius.     

Ant species diversity in the 'Grands Causses' (Aveyron, France): In search of sampling methods adapted to temperate climates

This study aimed at showing the applicability of using a combination of four sampling methods (i.e., Winkler extractors, pitfall traps, baiting and manual collection), something most often conducted in the tropics, to create an inventory of ant species diversity in temperate environments. We recorded a total of 33 ant species in the Grands Causses by comparing three vegetal formations: a steppic lawn ('causse' sensu stricto), which was the most species-rich (29 species), followed by an oak grove (22 species) and a pine forest (17 species). No sampling method alone is efficient enough to provide an adequate sampling, but their combination permits one to make a suitable inventory of the myrmecofauna and to obtain information on the ecology of these ant species.     

Rainfall Influences Ant Sampling in Dry Forests

The standardized 'Ants of the Leaf Litter' protocol aims to facilitate the use of ground-foraging and litter-dwelling ants in biodiversity assessment and monitoring programs. It was initially developed to characterize assemblages from tropical rain forests and is based on two main techniques: Winkler extractions and pitfall traps. Here, we tested to what extent this protocol was adapted to tropical dry forests and affected by the rainfall regime. Our 10 study sites were located along an aridity gradient (average annual rainfall: 350–1300mm) in the Gran Chaco. The number of species collected per sampling effort increased with aridity for pitfalls but followed an opposite trend for Winkler samples. This trend could be explained by the low daytime foraging activity in the leaf litter during drought periods. In arid and semiarid regions the good performance of pitfalls was probably related to their 24-h operation and to the attractiveness of the water they contained. Our results stress that the Winkler method used in the Ants of the Leaf Litter protocol may not be cost-effective during periods of drought and may lead to severe underestimations of litter ant diversity in tropical dry forests.     

Limited effects of dominant ants on assemblage species richness in three Amazon forests

1. Ants are highly interactive organisms and dominant species are considered to be able to control the species richness of other ants via competitive exclusion. However, depending on the scale studied, inter‐specific competition may or may not structure biological assemblages. To date, ant dominance–richness relationships have only been studied in small sample units, where a few dominant colonies could plausibly control most of the sample unit. 2. We conducted a comprehensive survey of terrestrial ant assemblages using bait, pitfall, and litter‐sorting methods in three sites in Brazilian Amazonia. Using a spatially structured rarefaction approach, based on sampling units with linear dimensions ranging from 25 to 250 m, the mesoscale patterns of ant dominance–richness relationships (sampling units covering hundreds of meters separated by kilometers) were investigated. 3. Interference–competition models (parabolic or negative linear relationships between species richness and the abundance of dominant ants) tended to be more frequent in smaller sample units or in assemblages sampled with interactive methods, such as baits. Using more inclusive sampling methods, the relationship was generally asymptotic rather than parabolic, with no reduction in species diversity because of the presence of dominants. Random co‐occurrence patterns of species within sites support the interpretation of a limited role for present‐day competition in structuring these assemblages. 4. Competition from dominant species may reduce species richness in small areas, especially when artificial baits are used, but appears to be less important than environmental constraints in determining ant species richness across scales of hectares and greater in these Amazon forests.     

Effects of environmental factors on the ant fauna of restinga community in Rio de Janeiro, Brazil

The effects of environmental factors on the richness, diversity and abundance of ants were studied in the Restinga da Marambaia, south coast of Rio de Janeiro State, Brazil. The samples were taken using pitfall traps in August/2004 (winter) and March/2005 (summer) in three different vegetation types: (1) herbaceous ridge palmoid (homogeneous habitat); (2) shrub dune thicket and (3) ridge forest (heterogeneous habitats). At each habitat a range of environmental attributes was recorded: soil temperature and humidity, percentage of soil covering by litter and litter depth. Ninety-two ant species belonging to 36 genera and eight subfamilies were recorded. Density of ant species and abundance varied significantly between habitats and seasons; ant diversity varied only between habitats. Homogeneous habitat had lower ant species density, abundance and diversity than heterogeneous habitats. The two first variables were positively correlated with litter depth and both were higher in summer than in winter samples. There were more species of Ponerinae and Ectatomminae in heterogeneous than in the homogeneous habitat, whereas the Formicinae species were more abundant in the later.     

Ants as Bioindicators of Natural Succession in Savanna and Riparian Vegetation Impacted by Dredging in the Jequitinhonha River Basin,Brazil

We investigated the response of ant species to landscape and geomorphologic parameters of a long‐term (7–11 years) restoration project in the Jequitinhonha River (Northern State of Minas Gerais, Brazil) margins, previously dredged by a diamond mining company. Geomorphological changes from the dredging were severe and the area is unlikely to be adequately restored, mainly due to the negative effects of flooding. Our hypothesis is that ant species assemblages bioindicate successional stages and soil characteristics. We studied the association of effects from the river's flooding zone, the native vegetation, and sedimentary grain size with that of ant species diversity, abundance, and composition. An ant sampling program was conducted in April 2005, using three methods: baits, pitfall traps, and direct collection. Grain size was measured by sieving. In total, 10,784 ants were sampled, belonging to 7 subfamilies, 24 genera and 45 morphospecies. Ant species richness was greater in the undisturbed savanna area than in the restored habitats, and equivalently greater in the ecotone and intermediate zone habitats than on the river bank, the poorest habitat. Atta sexdens rubropilosa indicated a condition related to small forest remnants having well‐structured soil. On the other hand, ants with a body length of under 0.5 cm (Dorymyrmex pyramicus and Pheidole fallax) predominated in sandy areas, where the majority of the granules were the finest. The lack of organic matter and soil structure for constructing suitable nests may prevent large ants from colonizing such areas, and thus inhibit the advance of natural succession.     

Sampling efficacy for the red imported fire ant Solenopsis invicta (Hymenoptera: Formicidae)

Cost-effective detection of invasive ant colonies before establishment in new ranges is imperative for the protection of national borders and reducing their global impact. We examined the sampling efficiency of food-baits and pitfall traps (baited and nonbaited) in detecting isolated red imported fire ant (Solenopsis invicta Buren) nests in multiple environments in Gainesville, FL. Fire ants demonstrated a significantly higher preference for a mixed protein food type (hotdog or ground meat combined with sweet peanut butter) than for the sugar or water baits offered. Foraging distance success was a function of colony size, detection trap used, and surveillance duration. Colony gyne number did not influence detection success. Workers from small nests (0- to 15-cm mound diameter) traveled no >3 m to a food source, whereas large colonies (>30-cm mound diameter) traveled up to 17 m. Baited pitfall traps performed best at detecting incipient ant colonies followed by nonbaited pitfall traps then food baits, whereas food baits performed well when trying to detect large colonies. These results were used to create an interactive model in Microsoft Excel, whereby surveillance managers can alter trap type, density, and duration parameters to estimate the probability of detecting specified or unknown S. invicta colony sizes. This model will support decision makers who need to balance the sampling cost and risk of failure to detect fire ant colonies.     

Effect of collect time on communities of epigaeic ants (Hymenoptera: Formicidae) in areas of Eucalyptus cloeziana and cerrado

This study aimed at evaluating the effect of collect time (day and night) on ant fauna attracted to baits in areas of Eucalyptus cloeziana (Myrtaceae) and cerrado. The ants were collected in Fazenda Boa Vista, Mannesmann Fi - El Florestal Ltda, Paineiras, Minas Gerais State. Eigthteen sample units were collected: 12 in E. cloezina and six in cerrado. Each sample unit consisted of three plots (25 x 35 m each). The plot consisted of 34 baits distributed in a grid pattern at 5 m intervals. The sampling was carried out in the diurnal and nocturnal period. The results obtained revealed that both type of vegetation (cerrado x Eucalyptus) and the collect time (day x night) had a significative influence on the epigaeic ant fauna. The ordination (DCA) indicated that collect time effect was more important to fauna structuration than the vegetation effect. Brachymyrmex sp.1, Brachymyrmex sp.2, Camponotus crassus Mayr, Camponotus rufipes (Fabricius), Cephalotes pusillus (Klug) and Ectatomma brunneum Smith were indicator species of nocturnal period, and Camponotus renggeri-Emery, Camponotus atriceps (Smith), Camponotus melanoticus Emery and Paratrechina sp.1 were indicators of nocturnal period.     

Ant diversity (Hymenoptera: Formicidae) from capões in Brazilian Pantanal: relationship between species richness and structural complexity

Species richness of epigeic ants was surveyed in forest islands named cap?es of Brazilian Pantanal and related with their structural complexity. The ants were collected using pitfall traps in 28 cap?es from Rio Negro Farm, in Aquidauana municipality, Mato-Grosso do Sul state, Brazil. The structural complexity of cap?es was evaluated by measuring vegetation density and litter quantity near the pit-fall traps. Seventy-one species, distributed in 26 genera and seven sub-families were found. Ectatomma edentatum Roger (Formicidae: Ectatomminae) and one species of Pheidole were the most frequent species. Species richness was positively correlated only with herbaceous vegetation density of cap?es, supporting the idea that the increase in environmental heterogeneity diminishes species competition, allowing species co-occurrence.     

Sampling local ant diversities and the importance of trait analyses

Ants are an incredibly diverse and ubiquitous group of invertebrates in most terrestrial ecosystems. Although extensively sampled, the majority of ant inventories do not evaluate the effect of different sampling techniques in capturing nontraditional metrics of diversity. We aimed to quantify TD (taxonomic) and FD (functional) diversities for a local ant assemblage by integrating metrics and evaluating complementarity of pitfall traps and Winkler extractors for the leaf litter versus epigeic ant faunas and to determine the effect of sampling techniques on functional composition (community-weighted means of 11 morphological traits) and functional diversity (multi-trait morphospace measured with three different metrics). We sampled the local community in an Atlantic Forest fragment using week-long pitfall traps and 1m² leaf litter samples submitted to Winkler extractors and quantified the contribution on the technique to uniquely capture the ant morphospace by applying a new index (PWindex). Although ant TD overlapped, FD was significantly affected by the sampling technique. By controlling for TD effects, the community collected by each technique was differentially structured. Higher TD did not translate into wider morphospace for Winklers. Pitfalls recovered more functionally overdispersed assemblages. Pitfalls and Winklers overlapped in the sampling of the overall community, but each sampling method contributed with a unique spectrum to the ant morphospace. Our results suggest the importance of incorporating FD metrics in local ant inventories and the importance of sampling techniques when measuring the magnitude of FD and community structure. Our PWindex further illuminates sampling effects for ant assemblages. Abstract in Portuguese & Spanish is available with online material.     

Estimating density of ant nests using distance sampling

The quantification of ant nest densities is a useful but challenging task given the group’s high abundance and diversity of nesting sites. We present a new application of a distance-sampling method which follows standard distance analytical procedures, but introduces a sampling innovation that is particularly useful for ants; instead of having an observer look for ants we let ants find a bait station and measure the distances covered between nest and station. We test this method by estimating the density of epigaeic ant nests in an Amazon tropical forest site near Manaus, Brazil. We distributed 220 baits of canned sardine mixed with cassava flour among 10, 210-m long transects in old-growth upland forest. Forty-five minutes after baiting, we followed the ants’ trails and measured the linear distance between the bait and each nest’s entrance. We then used the freely available program DISTANCE to estimate the number of nests per unit area while accounting for the effect of distance on the probability that a colony will find a bait. There were found 38 species nesting in 287 different colonies, with an estimated 2.66 nests/m². This estimate fell within the 95 % confidence bounds of nest density predicted for a similar number of species based on a literature survey of ant species richness and nest density. Our sampling solution, however, takes less than 30 % of the time used by conventional sampling approaches for a similar area, with the advantage that it produces not only a point estimate but also a quantification of uncertainty about density.     

Contrasting Effects of Fire on Arboreal and Ground‐Dwelling Ant Communities of a Neotropical Savanna

Ants are a dominant group in tropical savannas and here we examined the responses of the arboreal and ground‐dwelling ant fauna to a fire in a Neotropical savanna (cerrado) reserve in Central Brazil. Ants were collected using pitfall traps and baits placed in trees and on the ground beneath each tree. Of the 36 trees marked along two transects, half (from each transect) were burned and half not. The same trees were sampled 1 wk before and again 3 and 12 mo after the fire. Rarefaction curves and ordination analyses using data from all trees from each side of each transect indicated that overall ant species richness and composition did not change after fire. Fire, however, reduced the mean number of ant species per tree, and increased the mean number of species on the ground. Fire increased the average abundance of specialist predators, Camponotini, and opportunistic species, and decreased that of arboreal specialists. Changes in the ground‐dwelling fauna were only detected 12 mo after the fire, while those in the arboreal fauna occurred earlier and were no longer apparent 12 mo after the fire. We suggest that these contrasting results represent mainly an indirect response of the ant communities to fire‐induced changes in vegetation. Given the temporary and small scale nature of the effects detected and the overall resilience of the ant fauna, our results indicate that a single fire in the cerrado vegetation does not greatly impact the structure of ant communities in the short term.     

Hunting for Wasps In-between: the Use of the Winkler Extractor to Sample Leaf Litter Hymenoptera

Soil ecosystems are considered biodiversity hotspots. Most of terrestrial organisms depend almost exclusively on soils at least during some part of their life cycle. Although there is an intensive effort to understand the dynamics involving communities of soil animals, most groups are unappreciated in faunal inventories, such as some parasitic Hymenoptera with soil or litter hosts (SLH Hymenoptera), which are considered one of the largest and most diverse groups of soil- or litter-dwelling animals. Contrasting with flight interception traps, techniques that sample soil and litter specimens normally present low costs and easy transportation, being easily replicated within a site. Among them, the Winkler extractor is a useful technique to sample cryptic species associated with leaf litter. The aim of the present work was to describe the SLH Hymenoptera fauna collected in a pool of samples, using a technique to sample leaf litter arthropods. In this work, we report a total of 600 specimens belonging to Hymenoptera (excluding ants), belonging to 15 families distributed within 24 subfamilies, in samples of leaf litter obtained using the Winkler extractor. We observed that this technique is promising for sampling SLH Hymenoptera and should be considered as a complement for inventories of the Hymenoptera fauna.     

Influences on the structure of suburban ant (Hymenoptera: Formicidae) communities and the abundance of Tapinoma sessile

Urbanization can alter the organization of ant communities and affect populations of urban pest ants. In this study, we sampled ant communities in urban and suburban yards to understand the habitat factors that shape these communities and influence the abundance of a common pest species, Tapinoma sessile (Say). We used pitfall traps to sample ant communities and a combination of pitfall traps and baiting to collect T. sessile at 24 sites in Knoxville, TN. In total, we collected 46 ant species. Ant species richness ranged from seven to 24 species per yard. Ant species richness tended to be lowest near houses, whereas T. sessile abundance was highest near houses. The best predictors of ant species richness in yards were canopy cover and presence of leaf litter: ant species richness peaked at mid-levels of canopy cover and was negatively correlated with the presence of leaf litter. Tapinoma sessile abundance increased with presence of logs, boards, or landscaping timbers and leaf litter in yards. Our results indicate that ant communities and the abundance of particular pest species in these urban and suburban landscapes are shaped by many of the same factors that structure ant communities in less anthropogenically disturbed environments.