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.     

Evaluation of three methods for sampling ground-dwelling Ants in the Brazilian Cerrado

Few studies have evaluated the efficiency of methods for sampling ants, especially in regions with highly variable vegetation physiognomies such as the Cerrado region of central Brazil. Here we compared three methods to collect ground-dwelling ants: pitfall traps, sardine baits, and the Winkler litter extractor. Our aim was to determine which method would be most appropriate to characterize the ant assemblages inhabiting different vegetation types. More species were collected with pitfall traps and with the Winkler extractor than with sardine baits. Pitfall traps collected more species in the cerrado (savanna) physiognomies, particularly in those with a poor litter cover, whereas the Winlker extractor was more efficient in the forest physiognomies, except the one subject to periodic inundations. There was a low similarity in species composition between forest and cerrado physiognomies, and this pattern was detected regardless of the method used to sampling ants. Therefore, even the use of a single, relatively selective method of collection can be enough for studies comparing highly distinct habitats and/or conditions. However, if the purpose of the sampling is to produce a more thoroughly inventory of the ant fauna, we suggest the use of a combination of methods, particularly pitfall traps and the Winkler extractor. Therefore, the Ants of the Leaf-Litter (ALL) Sampling Protocol appear to be an adequate protocol for sampling ants in the highly-threatened Brazilian cerrado biome.     

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.     

Exploring a new biodiversity frontier: subterranean ants in northern Australia

Although it is common for ant surveys to uncover previously uncollected species, a recent study of subterranean ants in Amazonian Ecuador has indicated that an entire ant fauna may remain largely undiscovered. Here we report on the first systematic investigation of subterranean ants in northern Australia, in order to assess the extent to which the high abundance and diversity of subterranean ants in Amazonia is apparent in tropical Australia. We use a novel sampling technique that combines elements of an attractant bait and a pitfall trap, and allows many traps to be deployed simultaneously. Our main study was conducted at three closely approximated sites in Darwin, where the local ant fauna has been intensively surveyed using conventional (above-ground) sampling techniques. The 720 traps deployed resulted in 421 species records, representing 29 species from 17 genera. Sixteen of these species have cryptobiotic morphology, with four recorded here for the first time. Remarkably, one of these four (a blind species of Solenopsis) was the second most frequently caught species in subterranean traps, with 70 records. Ant abundance, species richness and composition varied markedly between sites, despite site similarity in soils and vegetation. Total ant records were greater in the middle compared with start of the wet season, declined with depth, and were greater after 4 days than one. Sampling at six sites in the Mitchell Falls area of the northern Kimberley region, 1,200 km southwest of Darwin, also revealed several cryptobiotic species new to science, including a new genus record (Pseudolasius) for Western Australia. Our underground sampling has therefore revealed an abundant and diverse subterranean ant fauna in northern Australia, containing many cryptobiotic species not previously collected. We use our results to provide methodological guidelines for most effectively sampling this fauna. Combined with the Amazonian study, our findings indicate that a specialist subterranean ant fauna, including numerous species remaining to be discovered, might be a feature of tropical landscapes throughout the world.     

New Caledonia has a depauperate subterranean ant fauna, despite spectacular radiations above ground

Ants are often a target of tropical biodiversity assessment because of their ecological dominance and value as indicators of ecosystem health, but high microhabitat specificity, patchy distribution and cryptic habits of many species make effective sampling problematic. Although tropical ant faunas have long been known to show strong vertical stratification, only recently has it been recognised that this can include a high diversity of subterranean species that are poorly sampled using traditional methods. Global diversity patterns of subterranean ants and their responses to above-ground disturbance remain largely unknown. We describe ants collected in 360 subterranean traps distributed across 15 sites representing contrasting soil types (volcano-sedimentary and ultramafic) in New Caledonia, a recognised Global Biodiversity hotspot. New Caledonia has a diverse above-ground ant fauna that includes spectacular radiations of both Gondwanan and Indo-Malayan genera from all above-ground layers (litter, epigaeic and arboreal), and so it might be expected also to harbour a subterranean ant fauna of high biogeographic and conservation significance. We show that New Caledonia supports an extremely depauperate subterranean ant fauna, especially on ultramafic soils, with only two cryptobiotic species recorded in subterranean traps, and a trap success rate up to an order of magnitude lower than in comparable Australian studies. Our results show that there is an uncoupling of ant diversity above- and below- ground, such that a high diversity and conservation significance of ants above ground is not necessarily matched below.     

Spatial and functional structure of an entire ant assemblage in a lowland Panamanian rainforest

Ants are a major ecological group in tropical rainforests. Few studies in the Neotropics have documented the distribution of ants from the ground to the canopy, and none have included the understorey. A previous analysis of an intensive arthropod study in Panama, involving 11 sampling methods, showed that the factors influencing ant β diversity (i.e., changes in assemblage composition) were, in decreasing order of importance, the vertical (height), temporal (season), and horizontal (geographic distance) dimensions. In the present study, we went one step further and aimed (1) to identify the best sampling methods to study the entire ant assemblage across the three strata, (2) to test if all strata show a similar horizontal β diversity and (3) to analyze the functional structure of the entire ant assemblage. We identified 405 ant species from 11 subfamilies and 68 genera. Slightly more species were sampled in the canopy than on the ground; they belonged to distinct sub-assemblages. The understorey fauna was mainly a mixture of species found in the other two strata. The horizontal β diversity between sites was similar for the three strata. About half of the ant species foraged in two (29%) or three (25%) strata. A single method, aerial flight interception traps placed alongside tree trunks, acting as arboreal pitfall traps, collected half of the species and reflected the vertical stratification. Using the functional traits approach, we observed that generalist species with mid-sized colonies were by far the most numerous (31%), followed by ground- or litter-dwelling species, either specialists (20%), or generalists (16%), and arboreal species, either generalists (19%) or territorially dominant (8%), and finally army ants (5%). Our results reinforce the idea that a proper understanding of the functioning of ant assemblages requires the inclusion of arboreal ants in survey programs.     

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.     

Hypogaeic pitfall traps: methodological advances and remarks to improve the sampling of a hidden ant fauna

Ant assemblages present a great vertical stratification, with microhabitats showing strong differences in relation to species composition. Among the microhabitats, the hypogaeic has been poorly studied. Hypogaeic or subterranean ants live in the deeper soil layers, which make the sampling logistics and operability a difficult work. The fact that the hypogaeic ant fauna is diversified and abundant, with low similarity to ant assemblages in other microhabitats, has promoted the development of several collecting techniques to sample this hidden ant fauna. Here, we verify the ability of hypogaeic pitfall traps to sample subterranean ants. In addition, we propose methodological advances and remarks about the use of this ant sampling technique.     

Sampling communities of ground-foraging ants: Pitfall catches compared with quadrat counts in an Australian tropical savanna

The relative abundances of ant species captured in pitfall traps was compared with those obtained by direct counts in quadrats at a savanna site in Kakadu National Park, Northern Territory. Two measures of abundance in traps were used, one based on total numbers of ants, the other on species frequency of occurrence. All species commonly recorded in quadrats were collected in traps, and their relative abundances were highly correlated on all occasions. Of the 20 most common species in quadrats, five occurred with a significantly different (in all cases lower) frequency in pitfall traps, but these species represented only 1.8–3.1% of total quadrat counts. Results from quadrats and pitfall traps were particularly similar (r > 0.8) when species-were classified into functional groups. Frequency data from traps may sometimes overestimate the abundance of widespread, solitary foraging species (e.g. ‘Chelaner’ and Tetramorium spp.) and underestimate species with large colony sizes (e.g. Iridomyrmex spp.). Data based on total numbers of ants in traps may be more prone to distortion arising from species differences in locomotor behaviour. Species counts in traps could be scaled to reduce these distortions. The finding that pitfall traps gave results comparable with those from quadrat counts provides support for the use of pitfall traps in studies of Australian ant communities in open habitats.     

Conservation value of road verges in the semi-arid Karoo, South Africa: ants (Hymenoptera: Formicidae) as bio-indicators

Ground-foraging ants (Hymenoptera: Formicidae) were used to assess the conservation value of road verges in a semi-arid region around Prince Albert, Western Cape Province, South Africa. Ant communities were sampled using pitfall traps on 50 sites along two roads. Four transects were sampled at each site, two in the road verge and two in adjacent rangeland. A total of 43,298 individual ants of 34 species were caught, with 31 and 32 species in the road verge and adjacent rangeland respectively, but on average road verges were the more species-rich. Road verges also contained relatively more rare species (i.e. those species that occurred in <10% of the traps. Ant species richness appeared to be influenced by food availability. Although ants appeared to be negatively impacted by higher grazing intensities in the rangeland, there was no difference in species richness between fenced and unfenced road verges. Species in the road verge may benefit directly from road kills, and indirectly from lower grazing pressure on plants, increased surface run-off and differences in soil surface temperatures. No alien ant species were observed or captured at any of the sites.     

Argentine ant invasion associated with loblolly pines in the southeastern United States: minimal impacts but seasonally sustained

Invasive ants are notorious for directly displacing native ant species. Although such impacts are associated with Argentine ant invasions (Linepithema humile) worldwide, impacts within natural habitat are less widely reported, particularly those affecting arboreal ant communities. Argentine ants were detected in North Carolina mixed pine-hardwood forest for the first time but were localized on and around loblolly pines (Pinus taeda), probably because of association with honeydew-producing Hemiptera. We explored the potential impacts of L. humile on arboreal and ground-foraging native ant species by comparing interspersed loblolly pines invaded and uninvaded by Argentine ants. Impacts on native ants were assessed monthly over 1 yr by counting ants in foraging trails on pine trunks and in surrounding plots using a concentric arrangement of pitfall traps at 1, 2, and 3 m from the base of each tree. Of floristics and habitat variables, higher soil moisture in invaded plots was the only difference between plot types, increasing confidence that any ant community differences were caused by Argentine ants. Overall patterns of impact were weak. Composition differed significantly between Argentine ant invaded and uninvaded trunks and pitfalls but was driven only by the presence of Argentine ants rather than any resulting compositional change in native ant species. Native ant abundance and richness were similarly unaffected by L. humile. However, the abundance of individual ant species was more variable. Although numbers of the arboreal Crematogaster ashmeadi (Myrmicinae) declined on and around invaded pines, epigeic Aphaenogaster rudis (Myrmicinae) remained the most abundant species in all plots. Argentine ant densities peaked in late summer and fall, therefore overlapping with most native ants. Unexpected was their continued presence during even the coldest months. We provide evidence that Argentine ants can invade and persist in native North Carolina forests, probably mediated by pine-associated resources. However, their localized distribution and minimal impact on the native ant fauna relative to previously described invasions requires further resolution.     

Is livestock grazing compatible with biodiversity conservation? Impacts on savanna ant communities in the Australian seasonal tropics

Grazing by domestic livestock is one of the most widespread forms of anthropogenic disturbance globally, and can have a major impact on biodiversity and therefore conservation values. Here we use ants to assess the extent to which livestock grazing is compatible with biodiversity conservation in a tropical savanna of northern Australia, where there is growing pressure to intensify pastoral production. We focus on the extent to which ant responses conform with four general patterns identified in a recent global review: (1) soil and vegetation type have a far bigger impact on ant community composition than does grazing; (2) grazing modifies ant species composition but often not species richness or total abundance; (3) a species’ response often varies among habitats; and (4) between 25–50% of the species that can be statistically analysed are responsive to grazing. We sampled ants using pitfall traps at 38 sites in two land systems, based on cross-fence comparisons of areas of different grazing intensities. A total of 130 ant species from 24 genera were recorded, with the fauna dominated by species of Iridomyrmex and Monomorium. Land system was the primary driver of variation in ant species richness and composition, and grazing intensity was related to neither species richness nor total abundance. Only 10% of common species appeared to be impacted by grazing. Overall, ant responses to grazing in our study region were generally consistent with the four global patterns, except that the local fauna seems to be particularly resilient. Such resilience indicates that current grazing management practices are compatible with the conservation of ant biodiversity.     

The ant fauna of Castelporziano Presidential Reserve (Rome,Italy) as a model for the analysis of ant community structure in relation to environmental variation in Mediterranean ecosystems

Ants are often used as ecological indicators in environmental monitoring programs, but rarely in Europe, probably because of a lack of baseline knowledge on the relationship between ant community structure and environmental variation. We present this baseline knowledge for Italian ecosystems that are typical of the Mediterranean climate, in order to assess the value of ants as ecological indicators in this biogeographic zone. Sampling areas were selected inside the Castelporziano Natural Reserve (Rome, Italy). The elevated variety of habitats and diverse ecological conditions, as well as the different levels of human impact in the Reserve, make this area a suitable location to develop and test biological indicators of environmental quality. The ant community structure was compared among several Mediterranean ecosystems using pitfall trapping, multidimensional scaling, the indicator species analysis and the Functional Groups Approach. Key factors for differences in ant community structure seem to be the presence/absence of water, the level of light and microhabitat diversity. Our results suggest that Castelporziano is an informative area in order to develop a study model for the Italian ant fauna and the Functional Group Approach is a promising procedure to associate differences in ant community structures with differences in habitat stress and disturbance.     

千岛湖陆桥岛屿地表蚂蚁群落物种多样性空间格局及其影响因素

为了探讨千岛湖岛屿景观参数对地表蚂蚁群落物种α和β多样性空间格局的影响, 作者分别于2017和2018年的5-8月, 采用陷阱法、凋落物分拣法和手捡法调查了千岛湖33个岛屿上的地表蚂蚁群落, 并依据食性将其划分为捕食性蚂蚁和杂食性蚂蚁。利用回归模型分析了全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁群落α和β多样性与岛屿景观参数的关系。结果表明, 岛屿面积对全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁的物种丰富度均有显著的正向影响, 而隔离度则无显著作用。蚂蚁群落的β多样性由空间周转组分主导。岛屿面积差对全部蚂蚁、捕食性蚂蚁和杂食性蚂蚁群落β多样性的嵌套组分有正向影响, 隔离度差只对杂食性蚂蚁的总体β多样性有正向影响。因此, 岛屿面积是影响千岛湖地表蚂蚁群落物种丰富度的主要因素, 并且岛屿面积通过嵌套组分来影响蚂蚁群落的β多样性, 表现出选择性灭绝过程。此外, 不同食性蚂蚁可能因为扩散能力的差异对岛屿景观参数产生不同的响应。     

Rainforest ants of the Tiwi Islands: a remarkable centre of endemism in Australia’s monsoonal tropics

The small and isolated rainforest patches that are embedded in the predominantly savanna landscape of Australia’s monsoonal tropics support a highly distinctive and biogeographically significant ant fauna. This fauna features shade-tolerant taxa of Indo-Malayan origin, in contrast to the arid-adapted, endemic Australian taxa that dominate the surrounding savanna. The Tiwi Islands north of Darwin in the Northern Territory (NT) receive the highest mean annual rainfall (up to 2,000 mm) in monsoonal Australia, and have a particularly extensive rainforest estate that has been poorly surveyed for invertebrates. Here we describe results from intensive ant surveys at 17 sites representing the full range of Tiwi rainforest types, using subterranean traps, Winkler sacs, pitfall traps and arboreal traps, supplemented by opportunistic hand collections. Our surveys yielded a total of 87 species from 37 genera, with the richest genera being Pheidole (9 species collected), Polyrhachis (8), Camponotus (5), Rhytidoponera (5) and Strumigenys (5). The overall structure and diversity of the Tiwi rainforest fauna is comparable to that of rainforest ant faunas on the Australian mainland. However, the species have exceptional biogeographic significance. At least 21 species across 12 genera have apparently never previously been collected, three species from subcoastal northeastern Australia are recorded for the first time in the NT, and the genera Mesoponera and Onychomyrmex are documented for the first time in the NT. There was a very low incidence of exotic species, which further highlights the conservation values of this remarkable fauna.     

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.     

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.     

Immediate impacts of invasion by Wasmannia auropunctata (Hymenoptera: Formicidae) on native litter ant fauna in a New Caledonian rainforest

Abstract For the last 30 years, Wasmannia auropunctata (the little fire ant) has spread throughout the Pacific and represents a severe threat to fragile island habitats. This invader has often been described as a disturbance specialist. Here we present data on its spread in a dense native rainforest in New Caledonia. We monitored by pitfall trapping the litter ant fauna along an invasive gradient from the edge to the inner forest in July 1999 and March 2000. When W. auropunctata was present, the abundance and richness of native ants drops dramatically. In invaded plots, W. auropunctata represented more than 92% of all trapped ant fauna. Among the 23 native species described, only four cryptic species survived. Wasmannia auropunctata appears to be a highly competitive ant that dominates the litter by eliminating native ants. Mechanisms involved in this invasive success may include predation as well as competitive interactions (exploitation and interference). The invasive success of W. auropunctata is similar to that of other tramp ants and reinforces the idea of common evolutionary traits leading to higher competitiveness in a new environment.     

陷阱法和Winkler法调查土壤节肢动物多样性比较:以千岛湖岛屿封闭生境研究为例

陷阱法和Winkler法是调查土壤动物的两种常规方法,然而这两种方法的调查效率各有优劣。本研究于2010年秋季,在千岛湖中心湖区选取了15个面积不同的岛屿,同时采用Winkler法和陷阱法采集岛屿上的土壤节肢动物,在岛屿这一封闭生境中比较两种方法收集土壤节肢动物的效率。结果表明,两种方法捕获土壤动物类群丰富度差异不显著,但多样性指数差异极显著。Winkler法对常规土壤节肢动物类群的采集效率优于陷阱法,尤其对运动较缓慢、活动范围较小的土壤节肢动物类群的采集具有优势;陷阱法则更优于采集运动能力较强、活动范围较大的类群。基于样方的稀疏曲线结果说明,Winkler法能用少量样方快速获取研究区域的土壤节肢动物群落基本组成,推荐在面积较小的岛屿上使用;而在面积较大的岛屿上使用陷阱法能够获取更多的类群。ChaoJaccard相似性系数比较则显示两种方法所取的土壤节肢动物相似性在大型岛屿上差异较大,说明大样本数据的采集需要两种方法同时使用能够提高数据的完整性和可靠性。本研究的结果为土壤节肢动物研究的方法选择提供了数据支持,具有一定的实际应用性。     

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.