Grazing history influences biodiversity: a case study on ground-dwelling arachnids (Arachnida: Araneae, Opiliones) in the Natural Park of Alpi Marittime (NW Italy)

Alpine pastures are typical examples of “high nature value farmlands”, representing important habitats harbouring unique biological communities. Disturbance induced by overgrazing influences significantly ecosystem processes, in which invertebrates play a major role. To develop new models of sustainable management of pastures, more knowledge is needed of the animal communities, essential to the ecological functioning of pasture ecosystems. The apparent poor ecological state of several pastures in the Natural Regional Park of Alpi Marittime (NW-Italy) lead us to evaluate the influence of grazing history on biodiversity, using spider and harvestmen assemblages as key groups. Four different pastoral types characterized by four different grazing histories were identified using the Daget-Poissonet method. Spiders and harvestmen were collected by means of pitfall traps. Arachnid assemblages were characterized in terms of composition, abundance, species richness, richness of endemic species and taxonomic relatedness. Generalized linear mixed models (GLMM) were used to test differences among assemblages occurring in each pastoral type. Furthermore, we tested differences in terms of plant communities (species richness and percentage of zoogenic species). Specificity and fidelity of every spider and harvestmen species within pastoral types were explored by the IndVal (Indicator Value) procedure. Fifty-eight species of spiders and seven species of harvestmen were collected (2,304 individuals). Pastoral types related to intensive grazing were characterized by the dominance of diurnal wanderer spiders (namely Lycosidae) while, conversely, nocturnal wanderers (mainly Gnaphosidae) were more abundant in extensive pastoral types. Results show that both species richness and spider abundance were higher in abandoned areas of extensive grazing, while endemic assemblages were richer in extensive grazed areas, which also hosted the most diverse plant community. Furthermore, most of the indicator species of spiders of this type were endemic, characterized by more demanding ecological requirements.     

Global analysis reveals that cryptic diversity is linked with habitat but not mode of life

The ubiquity of genetically distinct, cryptic species is limiting any attempt to estimate local or global biodiversity as well as impeding efforts to conserve species or control pests and diseases. Environmental factors or biological traits promoting rapid diversification into morphologically similar species remain unclear. Here, using a meta‐analysis of 1230 studies using DNA sequences to search for cryptic diversity in metazoan taxa, we test two hypotheses regarding the frequency of cryptic taxa based on mode of life and habitat. First, after correcting for study effort and accounting for higher taxonomic affinities and biogeographical region of origins, our results do not support the hypothesis that cryptic taxa are more frequent among parasitic than free‐living taxa. Second, in contrast, the results support the hypothesis that cryptic taxa are more common in certain habitats than others: for a given study effort, more cryptic taxa are found in freshwater than in terrestrial or marine taxa. These findings suggest that the greater heterogeneity and fragmentation of freshwater habitats may promote higher rates of genetic differentiation among its inhabitants, a general pattern with serious implications for freshwater conservation biology.     

Non-volant mammalian diversity in fragments in extreme eastern Amazonia

Current deforestation practices are likely to result in fragmentation of much of Amazonia. Extreme eastern Amazonia (here referring to the region east of the Tocantins River) is the most populated area of the Brazilian Amazon and, unfortunately, represents the likely future scenario for remaining Amazonia. Although data are available on mammals in central Amazon fragments (the BDFFP project, ), surprisingly little is known about mammalian distribution and responses to fragmentation in eastern Amazonia. As an initial step towards understanding these responses, we compiled available data on mammalian assemblage composition in four fragments east of the Tocantins River, in northeastern Pará, Brazil, between 2002 and 2006. These fragments are privately owned and embedded within a matrix of secondary forest, pasture, slash-and-burn agriculture, and roads. Survey methods included diurnal line transect censusing, nocturnal censusing, live trapping of small mammals, opportunistic observations, and interviews with local informants. Despite environmental stresses, nearly all of the expected large mammalian fauna was recorded at least once, providing reason for optimism. We documented 58 species of mammals in nine orders, and most species we failed to encounter were small nocturnal taxa for which our sampling effort likely was inadequate. Although preliminary in nature, this study highlights the need for comprehensive faunal surveys and complementary ecological research on the fauna of extreme eastern Amazonia. It also indicates that the terrestrial fauna of the region shows indications of resilience in spite of a long history of exploitation.     

Determining woodpecker diversity in the sub‐Himalayan forests of northern India using call playbacks

ABSTRACT Tropical forests have exceptional woodpecker diversity, but little is known about the abundance and diversity of woodpeckers in the Indian subcontinent, particularly for the Shorea robusta‐dominated moist deciduous forests of northern India. Our objective was to compare the number of woodpecker species and number of individuals detected using playback surveys and visual/aural transect surveys at five sites. Each site was surveyed 5–6 times along a 2000‐m transect, with woodpeckers detected using two methods: (1) visual and aural cues, and (2) playing back calls of 13 species at 400‐m intervals. Both methods involved similar effort per survey (100–110 min). During surveys, we detected 11 species of woodpeckers. More species and more than twice as many individuals were detected during playback surveys than during visual/aural surveys. In addition, species accumulation curves showed that we detected the species known to be present based on previous work faster with playback surveys than with visual/aural surveys at four of the five sites. During field trials, 97% of targeted individuals (N= 269) of 12 species responded to playback, and 83% of the responses occurred within 1 min of broadcast. The number of species of woodpeckers in our study area (11 species) was typical for a structurally diverse, tropical/subtropical moist broad‐leaved forest. Our results demonstrate that playback surveys are more efficient and accurate than visual/aural surveys, and that playback surveys can be useful for assessing and monitoring woodpecker diversity in tropical forests.     

利用mtCOI PCR-RFLP技术鉴定中国境内九个烟粉虱隐种

烟粉虱Bemisia tabaci (Gennadius)是一个物种复合体, 包括31个以上形态上无法区分的隐种, 其中少数隐种是世界性入侵害虫。目前, 在中国境内分布有2个入侵隐种和13个土著隐种。快速、 高效的鉴别方法对掌握烟粉虱田间发生规律及制定相关防控策略具有重要意义。然而到目前为止, 除了mtCOI基因测序比对外, 尚未有一种简便的方法可以有效地区分多个烟粉虱隐种。本研究采用mtCOI PCR-RFLP技术, 单独或组合使用TaqI, VspI, Van91I, NcoI和FokI这5种限制性内切酶, 酶切烟粉虱mtCOI的PCR扩增片段, 鉴别分布在中国境内的9个烟粉虱隐种。结果表明, 单独使用TaqI酶切, 可鉴别出MEAM1和China 1两个隐种, 使用TaqI+NcoI分步酶切可鉴定出MED和Asia 1两个隐种, 使用TaqI+Van91I分步酶切可鉴定出Asia II 3和Asia II 9两个隐种, 使用TaqI+VspI+FokI分步酶切可鉴定出Asia II 1, Asia II 6和Asia II 7 3个隐种。本研究为高效鉴别中国境内的多个烟粉虱隐种提供了方法。     

Surveying carnivores at large spatial scales: a comparison of four broad-applied methods

Reliable methods to estimate species richness are very important to managers and conservationists because they provide key data to make the right decisions in conservation programmes. In the case of carnivore mammals, traditional methods, such as direct count censuses, are not useful since these animals are usually scarce, elusive and nocturnal. Difficulties in carnivore sampling are compounded when monitoring programmes are developed at large spatial scales, where high economic costs and field efforts are necessary to achieve reliable richness or abundance estimates. These problems have highlighted the need to find more effective carnivore survey methods, especially in regions with high rates of landscape change, such as the Mediterranean basin. The present study, performed in a typical Mediterranean area, was the first in Europe to test simultaneously the relative efficiencies of four broad-applied sampling methods to detect carnivore species at large spatial scales. Sign surveys based on scat detection, scent stations, camera-trapping and live-trapping were investigated. We compared efficiencies using biological parameters and by considering both the logistic and economic costs of each method. Overall, scent stations and sign surveys were the most efficient methods both in economic and logistic terms. In addition, the use of scent stations may be necessary to detect species rarely detected by scats. Detailed and extensive training programmes for conducting sign surveys and scent stations may overcome perceived problems thus enhancing the widespread use of both methods. Our results are applicable not only to other Mediterranean areas, but also to other habitats and regions of the world. More research into the suitability of these and other methods in relation to different landscapes, seasons and species is required.     

High levels of cryptic species diversity uncovered in Amazonian frogs

One of the greatest challenges for biodiversity conservation is the poor understanding of species diversity. Molecular methods have dramatically improved our ability to uncover cryptic species, but the magnitude of cryptic diversity remains unknown, particularly in diverse tropical regions such as the Amazon Basin. Uncovering cryptic diversity in amphibians is particularly pressing because amphibians are going extinct globally at an alarming rate. Here, we use an integrative analysis of two independent Amazonian frog clades, Engystomops toadlets and Hypsiboas treefrogs, to test whether species richness is underestimated and, if so, by how much. We sampled intensively in six countries with a focus in Ecuador (Engystomops: 252 individuals from 36 localities; Hypsiboas: 208 individuals from 65 localities) and combined mitochondrial DNA, nuclear DNA, morphological, and bioacoustic data to detect cryptic species. We found that in both clades, species richness was severely underestimated, with more undescribed species than described species. In Engystomops, the two currently recognized species are actually five to seven species (a 150-250% increase in species richness); in Hypsiboas, two recognized species represent six to nine species (a 200-350% increase). Our results suggest that Amazonian frog biodiversity is much more severely underestimated than previously thought.     

Investigating detection success: lessons from trials using decoy rare plants

Imperfect detection leads to underestimates of species presence and decreases the reliability of survey data. Imperfect detection has not been examined in detail for boreal forest understory plants, despite widespread use of surveys for rare plants prior to development. We addressed this issue using detectability trials conducted in Alberta, Canada with decoy vascular plants. Volunteer observers searched in survey plots for species while unaware of their true presence or abundance. Our findings indicate that the detection of cryptic species is very low when abundance is low (0–35%) and plot size is large (<?50% in?≥?100 m²). Plant density (individuals per unit area) was the most important determinant of detection probability, where more abundant species were detected more often and with less survey effort. When abundance was held constant, diffusely arranged species were twice as likely to be detected compared to those in clumps. Detection of cryptic species can be low even when individuals are flowering, and even morphologically distinct species can go unnoticed in small plots. We suggest that future decoy trials investigate search strategies that could improve detection and that field surveys for vascular plants address imperfect detection through careful consideration of plot size, characteristics of the target species, and survey effort, both in terms of time expenditure within an area and the number of observers employed.     

Using infrared thermography to detect night‐roosting birds

Most birds sleep while roosting at night. Although a widespread behavior, few investigators have studied the nocturnal roosting behavior of birds. Studies conducted to date have either focused on species that roost communally or used radio‐telemetry to locate sleeping individuals of a few focal species. Portable thermal cameras capable of detecting infrared (IR) heat signals may provide a more efficient and less invasive means of detecting nocturnal‐roosting endotherms such as birds. Our objective was to assess the efficacy of using thermal cameras to detect roosting birds in a woodland bird community in southeastern Australia. To better understand the limitations of using thermography to detect roosting birds, paired bird surveys were conducted along 44 transects from May to September 2016 using both traditional survey techniques during the day and surveys with a thermal camera at night. We detected 195 birds representing 21 species at nocturnal roosts using IR thermography, with the detection rate of birds during nocturnal surveys approximately one‐third (29.1%) that during diurnal surveys. Detection rates during nocturnal surveys declined more steeply with distance from observers than for diurnal surveys. Detection rates were significantly higher during diurnal surveys for 14 species of woodland birds, but did not differ between diurnal and nocturnal surveys for eight other species. Roost height, roost visibility, bird mass, and cluster size (i.e., two or more birds in physical contact) did not differ between species categorized as having high or low detectability during nocturnal surveys. Variability among species in nocturnal‐detectability could not be attributed to roost‐site visibility, roost height, or bird size. Positive detection biases associated with diurnal behavior, such as movement and vocalizations, and limitations of current IR technology, e.g., low resolution, likely contributed to overall lower detection rates during nocturnal surveys. However, our results suggest that infrared thermography can be an effective and useful technique for detecting roosting birds and studying roosting behavior, as well as for population monitoring under certain conditions.     

Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness

Species richness is a fundamental measurement of community and regional diversity, and it underlies many ecological models and conservation strategies. In spite of its importance, ecologists have not always appreciated the effects of abundance and sampling effort on richness measures and comparisons. We survey a series of common pitfalls in quantifying and comparing taxon richness. These pitfalls can be largely avoided by using accumulation and rarefaction curves, which may be based on either individuals or samples. These taxon sampling curves contain the basic information for valid richness comparisons, including category–subcategory ratios (species-to-genus and species-to-individual ratios). Rarefaction methods – both sample-based and individual-based – allow for meaningful standardization and comparison of datasets. Standardizing data sets by area or sampling effort may produce very different results compared to standardizing by number of individuals collected, and it is not always clear which measure of diversity is more appropriate. Asymptotic richness estimators provide lower-bound estimates for taxon-rich groups such as tropical arthropods, in which observed richness rarely reaches an asymptote, despite intensive sampling. Recent examples of diversity studies of tropical trees, stream invertebrates, and herbaceous plants emphasize the importance of carefully quantifying species richness using taxon sampling curves.     

Successful carnivore identification with faecal DNA across a fragmented Amazonian landscape

The use of scat surveys to obtain DNA has been well documented in temperate areas, where DNA preservation may be more effective than in tropical forests. Samples obtained in the tropics are often exposed to high humidity, warm temperatures, frequent rain and intense sunlight, all of which can rapidly degrade DNA. Despite these potential problems, we demonstrate successful mtDNA amplification and sequencing for faeces of carnivores collected in tropical conditions and quantify how sample condition and environmental variables influence the success of PCR amplification and species identification. Additionally, the feasibility of genotyping nuclear microsatellites from jaguar (Panthera onca) faeces was investigated. From October 2007 to December 2008, 93 faecal samples were collected in the southern Brazilian Amazon. A total of eight carnivore species was successfully identified from 71% of all samples obtained. Information theoretic analysis revealed that the number of PCR attempts before a successful sequence was an important negative predictor across all three responses (success of species identification, success of species identification from the first sequence and PCR amplification success), whereas the relative importance of the other three predictors (sample condition, season and distance from forest edge) varied between the three responses. Nuclear microsatellite amplification from jaguar faeces had lower success rates (15-44%) compared with those of the mtDNA marker. Our results show that DNA obtained from faecal samples works efficiently for carnivore species identification in the Amazon forest and also shows potential for nuclear DNA analysis, thus providing a valuable tool for genetic, ecological and conservation studies.     

Cryptic species complexes,widespread species and conservation: lessons from Amazonian frogs of the Leptodactylus marmoratus group (Anura: Leptodactylidae)

Morphologically cryptic species act as a wild card when it comes to biodiversity assessments and conservation, with the capacity to dramatically alter our understanding of the biological landscape at the taxonomic, ecological, biogeographic, evolutionary, and conservation levels. We discuss the potential effects that cryptic species may have on biodiversity assessments and conservation, as well as some of the current issues involving the treatment of cryptic species both at taxonomic and conservation levels. In addition, using a combination of advertisement call and morphological data, we describe a new species of the Leptodactylus marmoratus group from the upper Amazon basin, and we assess how cryptic species can affect conservation assessments of species in the Leptodactylus marmoratus group by examining how recent findings affect our understanding of the distribution of what is assumed to be a widespread Amazonian species, Leptodactylus andreae.     

Resource utilization and guild structure of small vertebrates in the Amazon forest leaf litter

Ten species of small vertebrates that occurred together in leaf litter of the Amazon tropical rainforest were studied during the dry season in Rondônia, western Brazil. This vertebrate assemblage consisted of two diurnal lizards, two diurnal frogs, one nocturnal frog, and five frog species that are active by day and night. All species feed on small invertebrates, primarily insects and arachnids. There are significant differences in prey types and sizes eaten, and overlap values among species are relatively low. Niche breadths vary from unity (extremely low) to high (> 9.0). A pseudocommunity analysis using two different algorithms indicated that structure exists in the prey resource matrix of the vertebrate assemblage. The zero structure of the community matrix contributes to the observed structure, i.e. prey items not taken by certain species are important in maintaining structure. Several well-defined feeding guilds are apparent, and the feeding guilds do not appear to follow taxonomic lines.     

Nonparametric tests for homogeneity of species assemblages: a data depth approach

Testing homogeneity of species assemblages has important applications in ecology. Due to the unique structure of abundance data often collected in ecological studies, most classical statistical tests cannot be applied directly. In this article, we propose two novel nonparametric tests for comparing species assemblages based on the concept of data depth. They can be considered as a natural generalization of the Kolmogorov-Smirnov and the Cramér-von Mises tests (KS and CM) in this species assemblage comparison context. Our simulation studies show that the proposed test is more powerful than other existing methods under various settings. A real example is used to demonstrate how the proposed method is applied to compare species assemblages using plant community data from a highly diverse tropical forest at Barro Colorado Island, Panama.     

Lazy ecologist's guide to water beetle diversity: Which sampling methods are the best?

Biodiversity surveys of aquatic macroinvertebrates in standing water rely on various methods, but a thorough comparison of the techniques is lacking. This hampers analyses across surveys and impedes development of efficient sampling schemes. We compare the selectivity and efficiency of four methods commonly used to collect aquatic insects – activity traps (ATs), box trap (BT), handnetting (HN) and light trap (LT) – using a large dataset on water beetles in a site with ～100 species. We propose to use time investment as a natural basis to compare efficiency, since it applies to any method. The results inherently differ from results based on samples or individuals because methods are neither equally demanding nor equally rewarding. Most differences between methods arise from their size selectivity: ATs select for larger species, while HN and BT seem least selective. Attraction to light is taxon-specific and LT yields more depauperate samples than ATs, BT and HN, limiting the use of LT in community studies. To boost the development of cost-effective protocols, we also identify the best designs for rapid bioassesment by simulating short surveys from the data. Combinations of ATs and BT give most species; the results are robust to partitioning of effort between both methods. However, these rapid surveys miss on average more than 40% of all species in our study. Our results therefore emphasize that long-term studies using multiple methods are vital for measuring diversity in species-rich freshwater habitats.     

Factors influencing the nesting success of Lapwings Vanellus vanellus and behaviour of Red Fox Vulpes vulpes in Lapwing nesting sites

Capsule Lapwing nest predation was negatively correlated to nest density, while Lapwing alarm duration in response to foxes was positively correlated with the number of Lapwing broods present.

Aims To identify factors affecting Lapwing nest predation and Red Fox search effort.

Methods Lapwing nest success was monitored at four sites in 1996, seven sites in 1997 and six sites in 1998. In 1997 we mapped the position of all Lapwing nests in order to determine distances between nests, and the proximity of linear features and potential avian predator perches to each nest. From April to June 1998 we carried out 199 hours of nocturnal observations at six Lapwing nesting sites using night vision equipment.

Results The risk of nest predation was significantly higher for more isolated nests. Nocturnal observations showed that of all the nocturnal predators, foxes were the most active at Lapwing nesting sites. However, fox search effort in Lapwing colonies was relatively low, averaging 57 s/ha per visit. Foxes spent significantly longer foraging near breeding Lapwings (measured as duration of alarm calls) when more broods were present. Fox search effort (s/ha per hour of observation) tended to be greater in areas of high waterbird density.

Conclusion The lack of positive density-dependent nest predation, the relatively low search effort of foxes near Lapwing nesting sites and the high nest success sometimes achieved in areas with foxes all suggest that Lapwing nest predation by foxes is ‘incidental’. Lapwing chicks are probably more vulnerable to predation by foxes than clutches.     

Rapid assessment in conservation research: a critique of avifaunal assessment techniques illustrated by Ecuadorian and Madagascan case study data

The urgency of conservation concerns in the tropics, linked with the limitations imposed on research efforts by the tropical environment has resulted in the development of methods for rapid assessment of biological communities. One such method, the MacKinnon list technique, has been increasingly applied in avifaunal surveys worldwide. Using paired tropical bird data sets from Ecuadorian cloud forest and Madagascan littoral forest, we compare the performance of the MacKinnon list with that of the more standard method of point counts in indicating when a site has been adequately surveyed, estimating the magnitude of species richness, quantifying relative species abundance, and providing an α‐index of diversity. In species‐rich Ecuadorian cloud forest, neither method produced data indicating adequate survey effort, despite extensive sampling, whereas in the relatively species‐poor Madagascan littoral forests, data collected by both methods indicated that the area had been sufficiently surveyed with comparable sampling effort. Species richness estimates generated from MacKinnon list data provided a more accurate estimate of the magnitude of the species richness for the Ecuadorian avifauna, whereas estimates for the Madagascan avifauna stabilised with relatively few samples using either method. Data collected by each method reflected different patterns of relative abundance among the five most abundant species, with MacKinnon list data showing a bias towards solitary and territorial species and against monospecific flocking species relative to the point count data. As a consequence of this bias, MacKinnon list data also fail to reflect accurately the structure of communities as quantified by an index of community evenness. Point counts, on the other hand, failed to capture the full species complement of the species‐rich Ecuadorian study area. As techniques for the rapid assessment of unsurveyed areas, both methods are subject to biases that limit their value, if used alone, in collecting data of scientific and management value. We propose a hybrid rapid assessment methodology that capitalises on the strengths of both techniques while compensating for their weaknesses.     

eDNA metabarcoding: a promising method for anuran surveys in highly diverse tropical forests

Understanding the geographical distribution and community composition of species is crucial to monitor species persistence and define effective conservation strategies. Environmental DNA (eDNA) has emerged as a powerful noninvasive tool for species detection. However, most eDNA survey methods have been developed and applied in temperate zones. We tested the feasibility of using eDNA to survey anurans in tropical streams in the Brazilian Atlantic forest and compared the results with short‐term visual and audio surveys. We detected all nine species known to inhabit our focal streams with one single visit for eDNA sampling. We found a higher proportion of sequence reads and larger number of positive PCR replicates for more common species and for those with life cycles closely associated with the streams, factors that may contribute to increased release of DNA in the water. However, less common species were also detected in eDNA samples, demonstrating the detection power of this method. Filtering larger volumes of water resulted in a higher probability of detection. Our data also show it is important to sample multiple sites along streams, particularly for detection of target species with lower population densities. For the three focal species in our study, the eDNA metabarcoding method had a greater capacity of detection per sampling event than our rapid field surveys, and thus, has the potential to circumvent some of the challenges associated with traditional approaches. Our results underscore the utility of eDNA metabarcoding as an efficient method to survey anuran species in tropical streams of the highly biodiverse Brazilian Atlantic forest.     

An evaluation of the gradsect biological survey method

Biological surveys are necessary to gather species distribution data for the identification of priority conservation areas. The rationale of the gradsect method is that sampling (transects) oriented along the steepest environmental gradient should detect the maximum number of species in an area. The efficiency of the gradsect survey method was evaluated by comparing it to random, systematic and habitat-specific survey methods, during faunal field surveys (target groups: birds and dung beetles). Three gradsects were positioned within the study area to follow the major physiographical characteristics, incorporate all environmental strata (land facets) and yet be as logistically convenient as possible. The efficiency of survey methods was expressed as the number of species recorded per sampling unit effort and illustrated using bootstrap estimations to plot species accumulation curves. The gradsect method proved to be as efficient as the habitat-specific survey method and consiste ntly more efficient than the systematic and random surveys for both taxa sampled. The present study therefore illustrates that the gradsect survey method provides a cost-effective and swift representative sample of regional fauna. Moreover, the results indicate that land-form sequences, specifically land facets, are useful surrogates when sampling environmental diversity where distinct environmental gradients such as altitude and rainfall are absent.     

The ‘standardized search’: An improved way to conduct bird surveys

Abstract Bird surveys are among the most widely used biodiversity inventories and serve as the basis for an increasing proportion of pure and applied ecological research. It is rarely possible to conduct exhaustive censuses of all individuals present at a particular site, so stopping rules are routinely used to determine when sampling should finish. Most bird survey methods use (implicit) effort‐based stopping rules, either fixed times, fixed sampling areas (quadrats) or both, to standardize samples of different sites. If between‐site variation is high, however, a fixed sampling effort will generate samples of variable completeness with samples from smaller, less complex sites being more representative and complete than samples from larger, more complex sites. More importantly, quadrat‐based methods shift the scope of the overall study from bird occurrence in sites to bird occurrence in quadrats within sites, diminishing the impact of the research given that results cannot be extrapolated to relevant biological and management scales. Here I advocate an alternative means of conducting bird surveys, whereby the entire site is sampled and a results‐based stopping rule is used to ensure sample completeness is uniform across all sites. For example, a researcher may decide to continue sampling each site until two or fewer previously unencountered species are recorded in a 40‐min period. Samples of different sites will vary in both area and duration but will all be equivalently accurate estimates of species richness. This approach allows the avifauna of entire sites (whether territories, woodland remnants or catchments) to be sampled and compared directly, generating results and implications at the appropriate scale. In addition to yielding reliable measures of species richness, data collected this way can be used to calculate estimates of sample completeness and species incidence, two valuable metrics for ecological studies. This paper includes detailed worked examples of how to conduct a ‘standardized search’ and calculate sample completeness and species incidence estimates. I encourage further research on bird survey methods, and suggest that most current methods are insufficient, inconsistent and unreliable.