Using standardized sampling designs from population ecology to assess biodiversity patterns of therophyte vegetation across scales

Aim The analysis of diversity across multiple scales is hampered by methodological difficulties resulting from the use of different sampling methods at different scales and by the application of different definitions of the communities to be sampled at different scales. It is our aim to analyse diversity in a nested hierarchy of scales by applying a formalized sampling concept used in population ecology when analysing population structure. This concept involved a precise definition of the sampled vegetation type by the presence of a target species, in our case Hornungia petraea. We compared separate indices of inventory diversity (i.e. number of species) and differentiation diversity (i.e. extent of change in species composition or dissimilarity) with indices derived from species accumulation curves and related diversity patterns to topographical plot characteristics such as area and distance. Location Ten plots were established systematically over a distance of 100 km each in the distribution centre of H. petraea in Italy (i.e. Marche and Umbria) and in a peripheral exclave in Germany (i.e. Thuringia and Saxony‐Anhalt). Methods We used a nested sampling design of 10 random subplots within plots and 10 systematically placed plots within regions. Internal α‐diversity (species richness) and internal β‐diversity (dissimilarity) were calculated on the basis of subplots, α‐, β‐ and γ‐diversity on the basis of plots in Italy and Germany. In addition, indices of inventory diversity and differentiation diversity were derived by fitting species accumulation curves to the Michaelis–Menten equation. Results There was no significant difference in the internal α‐diversity between German and Italian plots but the α‐ and γ‐diversity were higher in Italy than in Germany. In Germany, the internal β‐diversity and β‐diversity were lower than in Italy. The differentiation diversity increased with increasing scale from subplots over plots to regions. The same results were obtained by calculating species accumulation curves. Significant positive correlations were encountered between the internal α‐diversity and α‐diversity in both countries, while the internal β‐diversity and internal α‐diversity showed a correlation only for the Italian plots. Similarity decay was found for German plots with respect to inter‐plot distance and for Italian plots with respect to altitudinal difference and to a smaller degree to distance between plots. Main conclusions The design chosen and the consistent analysis of species accumulation curves by the Michaelis–Menten equation yielded consistent results over different scales. The specific therophyte vegetation type in this study reflected diversity patterns also observed in other studies, e.g. a greater differentiation diversity in central than in peripheral habitats and a trend of increasing species richness towards lower altitudes. No asymptotic saturation of species richness between different scales was observed. Indications were found that the absolute level of inventory diversity at a particular scale and the completeness of the sampling procedure are the main clues for explaining the relationship between inventory and differentiation diversity at this particular scale.     

DNA barcoding of economically important freshwater fish species from north‐central Nigeria uncovers cryptic diversity

This study examines the utility of morphology and DNA barcoding in species identification of freshwater fishes from north‐central Nigeria. We compared molecular data (mitochondrial cytochrome c oxidase subunit I (COI) sequences) of 136 de novo samples from 53 morphologically identified species alongside others in GenBank and BOLD databases. Using DNA sequence similarity‐based (≥97% cutoff) identification technique, 50 (94.30%) and 24 (45.30%) species were identified to species level using GenBank and BOLD databases, respectively. Furthermore, we identified cases of taxonomic problems in 26 (49.00%) morphologically identified species. There were also four (7.10%) cases of mismatch in DNA barcoding in which our query sequence in GenBank and BOLD showed a sequence match with different species names. Using DNA barcode reference data, we also identified four unknown fish samples collected from fishermen to species level. Our Neighbor‐joining (NJ) tree analysis recovers several intraspecific species clusters with strong bootstrap support (≥95%). Analysis uncovers two well‐supported lineages within Schilbe intermedius. The Bayesian phylogenetic analyses of Nigerian S. intermedius with others from GenBank recover four lineages. Evidence of genetic structuring is consistent with geographic regions of sub‐Saharan Africa. Thus, cryptic lineage diversity may illustrate species’ adaptive responses to local environmental conditions. Finally, our study underscores the importance of incorporating morphology and DNA barcoding in species identification. Although developing a complete DNA barcode reference library for Nigerian ichthyofauna will facilitate species identification and diversity studies, taxonomic revisions of DNA sequences submitted in databases alongside voucher specimens are necessary for a reliable taxonomic and diversity inventory.     

Testing the performance of beta diversity measures based on incidence data: the robustness to undersampling

Aim Researchers measuring beta diversity have rarely concerned themselves with the problems of how complete the species lists of studied communities are, and of how the varying degrees of completeness can actually change estimates of beta diversity. No comprehensive assessment has been made regarding the behaviour of most beta diversity indices when applied to incomplete samples, a situation which is more common than usually recognized. Our objective was to assess the behaviour and robustness of a number of beta diversity measures for incidence data from undersampled communities. Location Mainland Portugal and the Azorean archipelago (North Atlantic). Methods Data from intensive sampling of spiders in mainland Portugal and arthropods in Azores were collected. We examined the properties of 15 beta diversity measures developed for incidence data. We simulated varying degrees of completeness, whereas computing beta diversity for selected pairs of samples. The robustness of these beta diversity accumulation curves was assessed for the purpose of finding the best measures for undersampled communities. Results The Harrison et al.β_‐2 and the Williams β_‐3 are particularly robust to undersampling. These measures are also insensitive to differences of alpha diversity (species richness) between communities, and therefore to nestedness. Colwell & Coddington β_cc and the related Jaccard β_j and Gaston et al.β_g performed best of the measures sensitive to alpha diversity. They performed poorly, however, when compared communities exhibited very low values of beta diversity. In such cases, the Routledge β_r performed the best. Main conclusions No index was found to perform without bias in all circumstances. Overall, β_‐2, β_‐3 and β_cc (or related measures β_j and β_g) are recommended as they seem to be the most robust to undersampling.     

Diversity dynamics of Zosterophyllopsida

The Zosterophyllopsida were major contributors to the diversification of early land plants. We present the first detailed analysis of the diversity dynamics of these plants from an updated database of all currently recognized zosterophyllopsid species. A set of quantitative methods classically used in palaeodiversity studies was applied to two data sets. The first one, ‘Zosterophyllopsida sensu stricto’, corresponds to the clade identified by Hao & Xue (The Early Devonian Posongchong Flora of Yunnan. (2013), Science Press). In the second, called ‘Zosterophyllopsida sensu lato’, barinophytalean‐type plants and taxa for which zosterophyllopsid affinities are suspected are added. The number of localities is used to explore sampling bias. Results show that sampling effect is minimal for the Early Devonian. For this time interval, both data sets record consistent patterns of changes suggesting that, whatever their affinities, all taxa included in the Zosterophyllopsida sensu lato show similar evolutionary trends. The diversity dynamics of zosterophyllopsids are characterized by a radiation during the Lochkovian, maximal values in the Pragian and a decline starting in the Emsian. The proportion of zosterophyllalean taxa with terminal sporangia is high until the Late Lochkovian when gosslingialean taxa without terminal sporangia evolved. During the Middle and Late Devonian, when diversity patterns are strongly affected by sampling, zosterophyllopsid diversity is low and characterized by a high proportion of barinophytacean and gosslingialean taxa, the latter becoming extinct in the Early Frasnian.     

Completeness of the fossil record and the validity of sampling proxies at outcrop level

Abstract: Most studies of the adequacy of the fossil record have been carried out at a global or continental scale, and they have used sampling proxies that generally do not incorporate all aspects of sampling (i.e. rock volume, accessibility, effort). Nonetheless, such studies have identified positive correlations between apparent diversity and various sampling proxies. The covariation of fossil and rock record signals has been interpreted as evidence for bias or for a common cause, such as sea level change, or as evidence that the signals are in some ways redundant with each other. Here, we compare a number of proxies representing the three main aspects of sampling, (1) sedimentary rock volume, (2) rock accessibility and (3) worker effort, with palaeodiversity in a geographically and stratigraphically constrained data set, the marine Lower Jurassic outcrop of the Dorset and East Devon Coast. We find that the proxies for rock volume and accessibility do not correlate well with the other sampling proxies, nor with apparent diversity, suggesting that the total amount of sedimentary rock preserved does not influence apparent diversity at a local scale, that is, the rock record at outcrop has been sampled efficiently. However, we do find some correlations between apparent diversity and proxies for worker effort. The fact that the proxies do not correlate significantly with each other suggests that none can be regarded as an all‐encompassing sampling proxy that covers all aspects of bias. Further, the presence of some correlations between sampling proxies and diversity most probably indicates the bonanza effect, as palaeontologists have preferentially sampled the richest rock units.     

Annual cycle and diversity of species and infraspecific taxa of Ceratium (Dinophyceae) in the Ligurian Sea,northwest Mediterranean1

We examined the well‐documented and species‐rich dinoflagellate genus Ceratium Schrank in the northwest Mediterranean Sea as a possible model for marine phytoplankton diversity and as a biological indicator of global climate change. First, we investigated the influence of counting effort; we then documented temporal changes in Ceratium specific and infraspecific taxa over 2 years (2002 and 2003) in the Villefranche Bay based on a monthly net sampling. Finally, we tried to identify factors associated with shifts in biodiversity. The calculation of taxonomic diversity, regularity, and richness were highly dependent on counting effort. We determined that a minimal sample volume of ～70 L was needed to obtain a good estimation of species richness. The annual cycle was characterized by a seasonal trend of high winter species richness followed by low spring biodiversity. Infraspecific variability not only appeared to depend on water temperature but also seemed to be influenced by bottom‐up control and was strongly affected by top‐down control. Thus, the occurrence of high concentrations of salps (Thalia democratica) and copepods larger than 2 mm (Calanus helgolandicus) coincided with a drastic decrease of Ceratium abundance and diversity during spring 2003. Ceratium is sensitive to both abiotic and biotic factors and could prove to be a good candidate as a biological indicator of global change.     

Bias in freshwater biodiversity sampling: the case of Iberian water beetles

Extensive distributional data bases are key tools in ecological research, and good-quality data are required to provide reliable conservation strategies and an understanding of biodiversity patterns and processes. Although the evaluation of data bases requires the incorporation of estimates of sampling effort and bias, no studies have focused on these aspects for freshwater biodiversity data. We used here a comprehensive data base of water beetles from the Iberian Peninsula and the Balearic Islands, and examine whether these data provide an unbiased, reliable picture of their diversity and distribution in the study area. Based on theoretical estimates using the Clench function on the accumulated number of records as a surrogate of sampling effort, about a quarter of the Iberian and Balearic 50 × 50 km Universal Transverse Mercator grid cells can be considered well prospected, with more than 70% of the theoretical species richness actually recorded. These well-surveyed cells are not evenly distributed across biogeographical and physicoclimatic subregions, reflecting some geographical bias in the distribution of sampling effort. Our results suggest that recording was skewed by relatively simple variables affecting collector activity, such as the perceived 'attractiveness' of mountainous landscapes and protected areas with recently described species, and accessibility of sampling sites (distance from main research centres). We emphasize the importance of these evaluation exercises, which are useful to locate areas needed of further sampling as well as to identify potential biases in the distribution of current biodiversity patterns.     

How well documented is Australia's flora? Understanding spatial bias in vouchered plant specimens

Massive digitization of natural history collections (NHC) has opened the door for researchers to conduct inferential studies on the collection of biological diversity across space and time. The widespread use of NHCs in scientific research makes it essential to characterize potential sources of spatial bias. In this study, we assessed spatial patterns in records from the Australian Virtual Herbarium (AVH), based on >3 000 000 vouchered specimens of around 21 000 native plant species. The AVH is the main database for describing Australia's flora, and identifying its limitations is of paramount interest for the validity of conservation and environmental studies. We characterized how sampling effort is distributed across each Interim Bioregion of Australia (IBRA), then asked: (i) How complete are species inventories for each bioregion? We define completeness (C) as the ratio of observed to estimated species richness, using the Chao 1 estimator, (ii) How is sampling effort related to a commonly used Human Influence Index (HII)? and (iii) What is the probability that additional collections would result in the identification of previously unrecorded species in each bioregion? Sampling effort across bioregions is unequal, which partially reflects the collecting behaviour of naturalists in relation to species richness patterns. The density of records in bioregions ranges from 0.02–8.37 km^?2. At the bioregional scale, completeness is generally high with 79% of bioregions estimated to have records for at least 80% of their species. Completeness is partly explained by sampling effort (r = 0.43, p = 0.01), although some bioregions (e.g. Northern Kimberley and Burt Plain) have high completeness yet relatively low sampling effort. The inventory of Hampton, however, is substantially less complete than other bioregions (C = 0.66). Bioregions with high HII consistently have high completeness, while regions with low HII span the full range of completeness values. We calculated that an additional specimen collected from a bioregion has a 0.33% (Wet Tropics) to 11.7% (Arnhem Coast) probability of representing a new species for that region. Our assessment can assist with directing future systematic survey efforts by identifying bioregions where additional surveying may result in the greatest return, in terms of increasing knowledge of species richness and diversity.     

Assessing the completeness of bryophytes inventories: an oceanic island as a case study (Terceira,Azorean archipelago)

How useful, complete or unbiased are comprehensive databases in order to provide reliable estimations of diversity? Using compiled data from bryophytes in Terceira Island (Azores), we specifically aim (1) to describe the register of species over time, (2) to assess the inventory completeness, i.e., the ratio between the observed and the maximum expected species, and (3) to locate the most promising areas for further surveys. First, each new recorded species was plotted against its collecting year, using the number of database-records as a surrogate of survey effort, to get the accumulation curves. These curves were then extrapolated to obtain the theoretical number of existing species according to Clench and exponential models. Spatial and habitat characteristics of the recorded taxa were also explored. Our results show an increasing trend in the rate of recorded species (c. five species per year), as well as a maximum of around a third of the theoretically “real” number of expected species that could yet remain unknown. Nevertheless, predictions of species richness were highly variable depending on the fitting curve used. Survey effort was similar between liverworts and mosses, as were inventory completeness values, but the rate of new recorded species was higher for mosses. Although bryologists visited preferably native habitats, we show that new species citations may also be found in modified habitats (e.g., exotic forests and semi-natural grasslands). We conclude that the analysis of extensive databases is a useful tool in revealing the recording and taxonomic gaps, further showing that bryophyte inventories could still be incomplete in Terceira Island. A strategy on how to improve species’ collections in remote areas is suggested, hoping to contribute to all-inclusive biodiversity studies in the Azores and elsewhere.     

Palaeodiversity and formation counts: redundancy or bias?

A key question in palaeontology is whether the fossil record taken at face value is adequate to represent true patterns of diversity through time. Some methods of assessing data quality have depended on the commonly observed covariation of palaeodiversity and fossiliferous formation counts through time, based on the assumption that the count of formations containing fossils, to a greater or lesser extent, drives diversity; but what if diversity drives formations? Close study of two fossil records, early tetrapods (Devonian–Jurassic) and dinosaurs, shows how the relationship between new taxa and new fossiliferous formations varies through research time. Initially, each new find represents a new fossiliferous formation and discovery follows the ‘bonanza’ model (fossils drive formations). In unexplored parts of the world, new taxa are identified frequently in new regions/formations. Only after time, in well‐explored continents such as Europe and North America, does collecting style switch to a mix of exploration for new formations and re‐sampling of known fossiliferous formations. Data are most striking for dinosaurs, where the Triassic–Jurassic record largely comprises finds from Europe and North America, where new formation discoveries reached their half‐life in 1914. This contrasts with the Cretaceous, which is dominated by rapidly rising discoveries from regions outside Europe and North America and the formation half‐life for these ‘new’ lands is 1986, showing that 50% of new Cretaceous dinosaur‐bearing formations were identified only in the past 30 years. The relationship between dinosaur‐bearing formations and palaeodiversity then combines three signals in variable amounts, reflecting the original diversity (relative abundances of particular taxa in different formations), redundancy (new fossiliferous formations accruing because of new fossil finds) and sampling (intensity of exploration for new fossiliferous formations, and of search within already‐sampled formations). For fossil vertebrates at least, formation counts of various kinds are poor predictors of sampling, missing, for example, the bonanza samples of Lagerstätten such as the Yixian Formation in China: thousands of specimens, dozens of species, but counted as one formation. These observations suggest that formation count cannot be regarded as an unbiased metric of sampling.     

Skeletal completeness of the non‐avian theropod dinosaur fossil record

Non‐avian theropods were a highly successful clade of bipedal, predominantly carnivorous, dinosaurs. Their diversity and macroevolutionary patterns have been the subject of many studies. Changes in fossil specimen completeness through time and space can bias our understanding of macroevolution. Here, we quantify the completeness of 455 non‐avian theropod species using the skeletal completeness metric (SCM), which calculates the proportion of a complete skeleton preserved for a specimen. Temporal patterns of theropod skeletal completeness show peaks in the Carnian, Oxfordian–Kimmeridgian and Barremian–Aptian, and lows in the Berriasian and Hauterivian. Lagerstätten primarily drive the peaks in completeness and observed taxonomic diversity in the Oxfordian–Kimmeridgian and the Barremian–Aptian. Theropods have a significantly lower distribution of completeness scores than contemporary sauropodomorph dinosaurs but change in completeness through time for the two groups shows a significant correlation when conservation Lagerstätten are excluded, possibly indicating that both records are primarily driven by geology and sampling availability. Our results reveal relatively weak temporal sampling biases acting on the theropod record but relatively strong spatial and environmental biases. Asia has a significantly more complete record than any other continent, the mid northern latitudes have the highest abundance of finds, and most complete theropod skeletons come from lacustrine and aeolian environments. We suggest that these patterns result from historical research focus, modern climate dynamics, and depositional transportation energy plus association with conservation Lagerstätten, respectively. Furthermore, we find possible ecological biases acting on different theropod subgroups, but body size does not influence theropod completeness on a global scale.     

DarkDivNet – A global research collaboration to explore the dark diversity of plant communities

DarkDivNet is a global research collaboration which explores dark diversity — the set of species that are absent from a site despite being suitable under the site conditions and present in the region. Participants of the network survey vascular plant diversity both at local (10  m × 10 m) and regional scales (radius 10 km) using a standardized approach. They also measure simple plant traits and collect soil samples. Observed and dark diversity together form the site‐specific species pool, and the ratio of observed diversity and dark diversity describes community completeness. We shall explore how observed and dark diversity, site‐specific species pool and community completeness vary across natural and anthropogenic gradients. We link plant diversity measures to the information obtained from environmental DNA: soil biota and plant taxa that occurred at the site before. We will refine existing dark diversity methods and use large vegetation databases to infer species habitat suitability. We expand the dark diversity concept from a purely taxonomy‐based approach to include the functional and phylogenetic aspects of diversity. DarkDivNet currently includes 161 planned sampling areas globally, but new participants are welcome. The main vegetation sampling period is scheduled until September 2020, with the first research papers being produced after that.     

Identifying recorder-induced geographic bias in an Iberian butterfly database

A database with comprehensive butterfly faunistic information from the Iberian Peninsula and the Balearic Islands was used to estimate inventory completeness as well as the environmental, spatial, and land-use effects on sampling intensities, on a 50×50 km UTM grid. The degree of sampling effort was assessed by means of accumulation curves based on the Clench function. Using the General Linear Model regression procedure, the effects of 22 variables on the estimated sampling efforts were assessed. This combination of methods is proposed as a preliminary step in biodiversity studies, in order to evaluate not only the degree of geographic coverage of existing faunistic data, but also the amount and nature of the bias on the faunistic work done throughout the last two centuries. The degree of spatial effects on the data was greater than the effects of environmental or land-use variables, although the latter two proved to be locally relevant. The results confirm previous findings that collecting is often skewed by relatively simple factors that affect collector activity, such as accessibility and attractiveness of sampling sites. With regard to Iberian and Balearic butterflies, adequate inventories on the scale investigated may probably suffice for further studies of the diversity of this insect group. Additionally, the results enabled us to develop general guide lines for the design of further faunistic work in the area.     

On formation‐based sampling proxies and why they should not be used to correct the fossil record

The fossil record is a unique resource on the history of life, but it is well known to be incomplete. In a series of high‐profile papers, a residual modelling technique has been applied to correct the raw palaeodiversity signal for this bias and incompleteness, and the claim is made that the processed time series are more accurate than the raw data. We apply empirical and simulation approaches to test for correlation and directionality of any relationships between rock and fossil data. The empirical data comprise samples of the global fossil record through the Phanerozoic, and we use simulations to assess whether randomly sampled subsets of modelled data can be improved by application of the residual modelling technique. Our results show that using formation counts as a sampling proxy to correct the fossil record via residual modelling is ill founded. The supposedly independent model of sampling is information‐redundant with respect to the raw palaeodiversity data it seeks to correct, and so the outputs are generally likely to be further from the truth than the raw data. We recommend that students of palaeodiversity cease to use residual modelling estimates based on formation counts, and suggest that results from a substantial number of papers published in the past ten years require re‐evaluation.     

Extrapolating inventory results into biodiversity estimates and the importance of stopping rules

Seven methods for predicting species diversity from inventory data were tested based on two model data sets. These data sets, derived from state automobile license plates observed in Mexico City and Lawrence, Kansas, had the advantage of providing known 'communities' to be sampled, allowing evaluation of different inference methods. Of the seven methods, those of Chao (1984 ), Clench ( Soberón & LLorente, 1993 ), and model M_th of CAPTURE ( Otis et al ., 1978 ) were the most robust. Error inherent in calculations based on raw data was reduced substantially using a series of bootstrap manipulations. We recommend that optimal design of inventories should include stopping rules based on precision of results rather than on effort expended, an approach that offers considerable advantages, in terms of both accuracy of results and efficiency of sampling efforts.     

Gauging megadiversity with optimized and standardized sampling protocols: A case for tropical forest spiders

Characterizing and monitoring biodiversity and assessing its drivers require accurate and comparable data on species assemblages, which, in turn, should rely on efficient and standardized field collection. Unfortunately, protocols that follow such criteria remain scarce and it is unclear whether they can be applied to megadiverse communities, whose study can be particularly challenging. Here, we develop and evaluate the first optimized and standardized sampling protocol for megadiverse communities, using tropical forest spiders as a model taxon. We designed the protocol COBRA‐TF (Conservation Oriented Biodiversity Rapid Assessment for Tropical Forests) using a large dataset of semiquantitative field data from different continents. This protocol combines samples of different collecting methods to obtain as many species as possible with minimum effort (optimized) and widest applicability and comparability (standardized). We ran sampling simulations to assess the efficiency of COBRA‐TF (optimized, non‐site‐specific) and its reliability for estimating taxonomic, phylogenetic, and functional diversity, and community structure by comparing it with (1) commonly used expert‐based ad hoc protocols (nonoptimized, site‐specific) and (2) optimal protocols (optimized, site‐specific). We then tested the performance and feasibility of COBRA‐TF in the field. COBRA‐TF yielded similar results as ad hoc protocols for species (observed and estimated) and family richness, phylogenetic and functional diversity, and species abundance distribution. Optimal protocols detected more species than COBRA‐TF. Data from the field test showed high sampling completeness and yielded low numbers of singletons and doubletons. Optimized and standardized protocols can be as effective in sampling and studying megadiverse communities as traditional sampling, while allowing data comparison. Although our target taxa are spiders, COBRA‐TF can be modified to apply to any highly diverse taxon and habitat as long as multiple collecting techniques exist and the unit effort per sample is comparable. Protocols such as COBRA‐TF facilitate studying megadiverse communities and therefore may become essential tools for monitoring community changes in space and time, assessing the effects of disturbances and selecting conservation areas.     

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.     

Assessment of sampling approaches for a multi‐taxa invertebrate survey in a South African savanna‐mosaic ecosystem

Invertebrate diversity is seldom included in conservation assessments, primarily because information is lacking. Broad surveys may be too costly, difficult or ineffective. Here we assess a ‘shopping basket’ approach, targeting 17 taxa using a range of methods. We sampled 43 one‐hectare sites stratified within 560 km² of heterogenous African savanna. We achieved up to 80% sampling completeness for epigaeic fauna, but generally much lower completeness (around 50%) for plant‐dwelling and flying taxa. For the former we identified duplication of methods, and for the latter, addition of methods and increased temporal variation rather than effort would improve completeness. Within a taxon, sampling 75% of species present required, on average, about 784 individuals. When considering the local richness, 75% completeness required about 27 individuals per species, but these figures require validation in other areas. About 58 sites were required to achieve 75% sampling completeness, translating to about one site per 10 km². The percentage of species sampled only in a particular month ranged between 4% and 46%, with greater temporal effects recorded for flying taxa than for epigaeic ones. The trend was similar for species unique to a particular year, with the most extreme case being 67% of the butterfly species sampled one year not previously recorded. We demonstrated and evaluated the feasibility of a simultaneous multi‐taxon survey approach to produce data useful for conservation planning and monitoring. We strongly recommend a quantified approach for surveys and inventories, with details such as specific methods decided based on the biome sampled, and taxonomic expertise available for identification.