A comparison of two methods for quantifying ant diversity and community in an East Himalayan montane forest mosaic

A comparison of two methods for quantifying ant diversity and community in an East Himalayan montane forest mosaic | Research Square window.SnipcartSettings = { analytics: { enabled: false } }; (function() { var accessVector = localStorage.getItem('access_vector') || ''; window.dataLayer = window.dataLayer || []; if (accessVector) { window.dataLayer.push({ user: { profile: { profileInfo: { snid: accessVector } } } }); } })(); (function(w,d,s,l,i){w[l]=w[l]||[];w[l].push({'gtm.start':new Date().getTime(),event:'gtm.js'});var f=d.getElementsByTagName(s)[0],j=d.createElement(s),dl=l!='dataLayer'?'&l='+l:'';j.async=true;j.src='https://www.googletagmanager.com/gtm.js?id='+i+dl;f.parentNode.insertBefore(j,f);})(window,document,'script','dataLayer','GTM-K279D39R'); Browse Preprints In Review Journals COVID-19 Preprints AJE Video Bytes Research Tools Research Promotion AJE Professional Editing AJE Rubriq About Preprint Platform In Review Editorial Policies Our Team Advisory Board Help Center Sign In Submit a Preprint Cite Share Download PDF Research Article A comparison of two methods for quantifying ant diversity and community in an East Himalayan montane forest mosaic Chuan-Jing Zhang, Yi-Ting Cheng, Xian-Shu Luo, Yao Chen, Yu-Chao He, and 4 more This is a preprint; it has not been peer reviewed by a journal. https://doi.org/ 10.21203/rs.3.rs-2261097/v1 This work is licensed under a CC BY 4.0 License Status: Published Journal Publication published 09 Aug, 2023 Read the published version in Journal of Insect Conservation → Version 1 posted 7 You are reading this latest preprint version Abstract Sampling efficiency, composition and detection biases associated with pitfall-trap and sample plot methods were compared at seven montane sites at Lasha Mountain, Yunnan, China. On average, sample plot samples contained 1.5 times more taxa than pitfall-trap samples; however, we found no significant difference between of alpha and beta diversity in pitfall-trap and sample plot site samples. Rarefaction-interpolations curves revealed significantly higher total diversity from sample plot methods; that sample plot methods would require three times more sampling to reach asymptote of true diversity; and that sample plot samples achieve higher sample coverage across sample sizes. Permutational multivariate analysis of variance showed community composition and dominant species differed between methods. Of all taxa collected, the two methods had 16 species in common, accounting for 52% of the total species; 29% were exclusive to sample plot samples and 16% were exclusive to pitfall traps. Implications for insect conservation : Our findings suggest that results from the two methods cannot be directly compared and are imperfect substitutes to one another. For long-term monitoring of biodiversity, we suggest integrating multiple complementary methods, including standardised active collection methods, such as the sample plot method, to achieve more complete representation of ant composition and diversity. Ant survey sampling method ground-dwelling epigaeic hypogaeic Formicidae Hengduan Mountains Figures Figure 1 Figure 2 Figure 3 Figure 4 Introduction Global biodiversity is declining at an unprecedented rate (IPBES 2019). Monitoring biodiversity is a key part of addressing the trend by providing facts of the occurring changes and mechanisms of change to bring about science-based policy and land management decisions that benefit biodiversity (Noss 1990 ; Hobbie et al. 2003 ; Kuussaari et al. 2009 ; Cardoso et al. 2011 ; Navarro et al. 2017 ; Guerra et al. 2021 ). The continuing challenge is measuring biodiversity in ways that are useful, accurate, and repeatable (Purvis & Hector 2000 ; Lindenmayer & Likens 2010 ). Ants, in particular, are an insect group of immense ecological significance, contributing a large proportion of the global arthropod biomass (Schultheiss et al. 2022 ) and having critical ecological functions such as nutrient cycling, soil aeration, and seed dispersal (Hölldobler & Wilson 1990 ; Parr et al. 2016 ). Ants are relatively easy to collect, sensitive to environmental change and representative of soil diversity (Andersen et al. 2004 ; Andersen & Majer 2004 ; Li et al. 2015b ). As such, ants have long been an important insect group for biological monitoring and indicators of ecological change (bioindicators) (Andersen 1997 ; McGeoch 1998 ; Andersen et al. 2002 ; Underwood & Fisher 2006 ; Zhang & Ou 2006 ; Gerlach, Samways & Pryke 2013 ; Tiede et al. 2017 ). Several sampling methods have been developed to collect ants, each with their own strengths and weaknesses (Bestelmeyer et al. 2000 ). Understanding the limitations to sampling techniques is important to selecting a method, or combinations of methods, that will address survey objectives within a given environment, sampling effort and available expertise (Bestelmeyer et al. 2000 ; Gotelli et al. 2011 ; de Souza et al. 2012 ). To allow stricter comparison of individual studies, numerous researchers have advocated for the adoption of recognised standard protocols (e.g., Li, Chen & Xu 2009 ; Antoniazzi et al. 2020 ). Such protocols have been proposed (e.g., Ants of the Leaf Litter (ALL) Protocol: Agosti & Alonso 2000 ) and successfully applied (Vineesh, Sabu & Karmaly 2007 ; Lopes & Vasconcelos 2008 ; Bray 2014 ; Yeo et al. 2017 ) but universal application has lagged. In China, the ‘sample-plot’ is the most used method, followed by pitfall traps. Pitfall traps are small, covered pits set in the ground that target active epigaeic fauna (Bestelmeyer et al. 2000 ). The widely-used method provides relatively simple and cost-effective sampling of epigaeic fauna that allows for continuous sampling for a time period, both day and night (Majer 1997 ). Pitfall traps suffer for their sensitivity to species size, trappability and activity (Topping & Sunderland 1992 ; Majer 1997 ), as well as biases to do with trap diameter, trap depth, spacing and habitat complexity (Luff 1975 ; Adis 1979 ; Ward, New & Yen 2001 ; Jiménez-Carmona, Carpintero & Reyes-López 2020 ). The sample plot method—a modified ‘direct sampling’ method (Bestelmeyer et al. 2000 )—is a technique where specimens are collected systematically from microhabitats using a range of hand-collecting techniques within a specified area and time (Xu 2002 ). The method has advantages in efficiency but is vulnerable to the competence of the researchers and differences in habitats, reducing the comparability between samples and studies (Bestelmeyer et al. 2000 ). Here, we aimed to evaluate these two widely used ant sample methods in China by assessing their results in terms of ant diversity and community composition in a forest mosaic in Yunling Nature Reserve, Yunnan Province. We expected that sample plot methods would collect more ant species than pitfall traps, where hand sorting could allow for sampling of more cryptic, submissive, and less abundant ants (Parr & Chown 2001 ; Longino, Coddington & Colwell 2002 ; de Souza et al. 2012 ; Mark & Guenard 2017). We also expected that sampling methods may selectively filter the ant community present so that combining methods could improve the survey and generate a closer estimation of true ant community (Longino & Colwell 1997 ). Our results will provide a measure of the sampling effort required and reliability of the combination of methods to characterise and monitor ground-dwelling ant diversity in a montane forest mosaic. Materials and Methods Study area The study was conducted at Lasha Mountain (拉沙山) (N26°20', E99°15') in Yunling Nature Reserve, Yunnan, China, within the Mountain Area of Southwest China biodiversity hotspot (Myers et al. 2000 ). At 75,894 km 2 , the Yunling Nature Reserve comprises a mosaic of land covers, including forest, regenerating forest, pastoral grazing land, cropland, and permanent settlements. The study area is a ca. 1400 ha catchment ranging from 2500 to 3700 m asl. Forest vegetation dominates most of the mid-elevations, transitioning from deciduous broadleaved forest ( Betula alnoides, Acanthopanax gracilistylus, Acer oliverianum ) at lower elevations, through mixed deciduous-conifer forest to conifer forest ( Abies georgei, A. fabri, Tsuga dumosa ) at the highest elevations. The forest undergrowth primarily comprises Rhododendron spp. and bamboo ( Fargesia strigosa, F. edulis, F. solida ). Few local families live in the lowest part of the catchment where there is a mixture of grazing, cropland and forest margins. The forest is used by local people for resources such a wood, traditional foods and medicine (Huang et al. 2017 ). The climate is characterised by alternating dry and wet seasons, with an annual mean precipitation of 910 mm (Wang et al. 2012 ; Li et al. 2019 ). Sample design We sampled ants using two common sampling methods (pitfall trap and sample plot) across the study area. Sample sites were established every 200 m elevation from 2500 m to 3700 m (totalling 7 sampling sites) using the stream as a centreline. Each sample site comprised a 60 m × 60 m plot, evenly divided across the stream. Five pitfall traps were stratified across each plot. The open-topped attractant bottles (11 cm in diameter, 15.5 cm in height) were embedded into the ground until flush with the natural soil surface and covered with 15-cm diameter plastic plates to keep rain out. Traps were baited with 50–70 ml liquid attractant composed of 1:1:4:16 solution of ethanol, sugar, vinegar, water (Li et al. 2017 ; Fang & Xu 2021 ). Trap surveys were conducted from August to September in 2018, 2019 and 2020. We emptied traps every two weeks during each sampling period. Sample plot sampling was undertaken in each 60 m × 60 m plot at five 5 m × 5 m sampling sub-plots. Subplots were paired with pitfall traps, buffered by 10 m. After an initial surface inspection, ants were collected by hand from litter, decaying wood and under stones for 1 hr (1 person hour). Using hand tools, soil was excavated to 20 cm to search for and collect ants from subterranean nests. A maximum thirty individuals were collected from a single nest. Foliage collections were made using a 2 m × 2 m white curtain was placed flat around the sample sub-plot and small trees and shrubs up to 5 m were vigorously shaken or beaten to capture ants that fell on the curtain. In total, each plot was investigated for 2 hr. Collected specimens from both methods were stored in 75% ethanol and transferred to a freezer at -10°C (Xu 2002 ) before being identified. Statistical analysis All analyses were performed using R v4.2.0 (R Core Team 2020 ). To reduce the effects of spatial autocorrelation of response variables, we pooled pitfall trap data by site and year. Sampling yielded 28 sampling units in total from 21 pitfall trap units and 7 sample plot units. For community analysis, we reduced data to presence-absence to reduce the effects of ant biology that lead to non-random distribution at some scales (Agosti et al. 2000 ). To visualise the number of species of ants detected with each method, a Venn diagram was drawn using the ‘eulerr’ package in R (Larsson 2021 ). We drew these for all species, as well as species classed as “dominant”, “common”, and “rare.” We defined dominant species as those with a total relative abundance exceeding 10%; common species contributed between 1% and 10% total abundance, and rare species contributed less than 1% to the total population sampled (Li et al. 2015a ). The effects of method on total measured alpha (α) and beta (β) diversity were tested using a Linear Mixed-Effects Model, using the function lmer in the ‘lme4’ package in R (Bates et al. 2015 ) with a parametric analysis of variance (ANOVA). Method blocked into year and elevation were introduced as random effects in the model. Normality and homogeneity of variance were checked by performing Quartile-Quartile (QQ) plots of the residuals and fitted models using the function plotresid in the package ‘RVAideMemoire’ (Hervé 2018 ). We identified and removed outliers in the model using the function romr.fnc in the package ‘LMERConvenienceFunctions’ (Tremblay & Ransijn 2015 ). Pairwise comparison of estimated marginal means (EMMs) were performed using the ‘emmeans’ package (Length 2022 ). We used rarefaction analysis to compare estimated asymptotic species diversity between the sample plot and pitfall trapping methods, a technique based on species frequency (i.e., presence-absence) that avoids biases caused by insufficient or differing sampling efforts (Gotelli & Colwell 2001 ). To do this, we used sample-sized-based rarefaction-extrapolation analysis based on sampling-unit-based incidence data (Chao et al. 2014 ). Hill species diversity (or the effective number of species) in the zero (q = 0), first (q = 1), and second (q = 2) orders (or species richness, the exponential Shannon entropy, and inverse Simpson index, respectively) were computed in the iNEXT package (Hsieh, Ma & Chao 2016 ). In addition, we compared sampling coverage (based on incidence) of each method across Hill species diversities to our sampling effort, permitting estimation of the proportion of the total community represented by the sampling effort and an assessment of sample completeness (sample coverage) across sampling units (Chao & Jost 2012 ; Chao et al. 2014 ; Hsieh, Ma & Chao 2016 ). Significant differences in estimated diversity and sampling coverage between methods were judged by non-overlapping confidence intervals (Chao, Chiu & Jost 2014 ). To evaluate differences in ant composition of samples obtained from the two methods, we performed non-metric multidimensional scaling (NMDS: calculated with Jaccard dissimilarity index) with the function metaMDS of the R package ‘vegan’ (Oksanen et al. 2019 ). We considered a species x sample matrix with presence-absence data (30 species x 28 samples). A dummy species was added to the species matrix to mitigate the impact of pitfall traps that had zero abundances (Clarke, Somerfield & Chapman 2006 ). We used the function ordiellipse of the vegan package to draw ellipses representing 95% CI around centroids. To test for differences between groups (methods) we used a permutational multivariate analysis of variance (PERMANOVA) using distance matrices, which was performed with the function adonis2 in vegan (Oksanen et al. 2019 ). Elevation was set as a random factor. P values were obtained using 9999 permutations of residuals. To determine which species contributed most to the observed multivariate differences between pitfall trapping and sample plot methods, we used similarity percentage analysis (SIMPER) (Clarke 1993 ). Results Ant collection Using two sampling methods, we found a total of 10,206 ant (Insecta: Formicidae) specimens belonging to four subfamilies (Myrmicinae, Formicinae, Dolichoderinae, and Ponerinae), 17 genera and 30 species (Table 1 ). Of these, 6312 and 3894 individuals were collected from pitfall trap and sample plot methods respectively. We recorded, 21 species from pitfall traps and 25 species from sample plots, of which only 16 species (53%) were shared (Fig. 1 ). The most abundant (‘dominant’) species across sampling methods were Pheidole nietneri Emery, 1901, Myrmica rugosa Mayr, 1865, Myrmica kozlovi Ruzsky, 1915, Myrmica bactriana Ruzsky, 1915, Formica fusca Linnaeus, 1758, together accounting for 70% of all the specimens collected (Table 1 , Fig. 1 ). We recorded 11 rare species from pitfall traps and 14 rare species from sample plots, seven of which were shared. Rare species contributed 5% to the total specimen abundance. Table 1 Number and account of species in Sample-plot method and Pitfall-trap method. For Classes, D = dominant species (> 10% of total); C = common species (1–10% of total); R = rare species (< 1% of total); (%) = % contribution to total abundance. Subfamily/species Class Pitfall Trap Sample Plot Sum (%) Dolichoderinae Tapinoma sinense Emery, 1925 C 0 106 106 1.04 Ponerinae Ponera bawana Xu, 2001 R 1 0 1 0.01 Myrmicinae Aphaenogaster caeciliae Viehmeyer, 1922 C 100 119 219 2.2 Aphaenogaster lepida Wheeler, 1930 R 0 3 3 0.03 Myrmica bactriana Ruzsky, 1915 D 490 697 1187 11.6 Myrmica jessensis Forel, 1901 R 0 1 1 0.01 Myrmica kozlovi Ruzsky, 1915 D 257 988 1245 12.2 Myrmica margaritae Emery, 1889 R 35 2 37 0.4 Myrmica pararitae Radchenko, 2008 R 1 98 99 1.0 Myrmica ritae Emery, 1889 R 38 26 64 0.6 Myrmica rugosa Mayr, 1865 D 1344 207 1551 15.2 Myrmecina striata Emery, 1889 R 3 0 3 0.03 Pheidole nietneri Emery, 1901 D 1955 2 1957 19.2 Pheidole pieli Santschi, 1925 R 0 1 1 0.01 Perissomyrmex bidentatus Zhou & Huang, 2006 R 0 8 8 0.08 Temnothorax sp.1 R 1 10 11 0.1 Temnothorax sp.2 R 1 0 1 0.01 Tetramorium kraepelini Forel, 1905 R 43 5 48 0.5 Stenamma bhutanense Baroni Urbani, 1977 R 1 49 50 0.5 Formicinae Camponotus anningensis Wu & Wang, 1989 R 0 52 52 0.5 Camponotus herculeanus Linnaeus, 1758 C 19 90 109 1.1 Lasius flavus Fabricius, 1782 R 0 2 2 0.02 Lasius coloratus Santschi, 1937 R 9 39 48 0.5 Lasius alienus Foerster, 1850 C 686 273 959 9.4 Nylanderia bourbonica Forel, 1886 R 0 2 2 0.02 Nylanderia flavipes Smith, 1874 C 53 96 149 1.5 Paraparatrechina aseta Forel, 1902 C 0 202 202 2.0 Prenolepis angularis Zhou, 2001 R 70 0 70 0.7 Formica fusca Linnaeus, 1758 D 360 816 1176 1.5 Formica sinensis Wheeler, 1913 C 844 0 844 8.3 Total 6312 3894 10206 Species diversity Comparisons of sampled alpha and beta diversity between to two methods did not differ Fig. 2 ); however, our Hill number series analysis (q = 0, 1, 2) showed significantly higher species diversity of ants from sample plot methods than pitfall trap methods (non-over-lapping in confidence intervals) for three estimators of diversity across sampling effort and coverage (Fig. 3 ). Total estimated species richness with iNEXT was 35 ± 7.7 s.e. for SP and 25 ± 4.7 s.e. for PF (Supplementary Information 1). Results were consistent when comparing diversity at a given sampling effort and coverage (Fig. 3 B–C). The sampling coverage method showed an approximate completeness of 76% in sample plot method and 95% in pitfall trap method using the true sampling effort (SP, n = 7; PF n = 21). Community composition The NMDS plots based on species presence-absence showed some clustering of the observed species composition according to method (Fig. 4 ). PERMANOVA results with elevation held constant showed PF and SP assemblages differed significantly (p < 0.05). The NMDS analysis had a low stress value (0.07) which indicates that the ordination summarised the observed distances between samples well (Fig. 4 ). In pitfall traps, we found high numbers of Pheidole nieneri (1955) and Myrmica rugosa (1344), whereas in sample plots we collected high numbers of Myrmica kozlovi (988) and Formica fusca (816) (Table 1 ). The SIMPER analysis showed that Myrmica kozlovi , M. bactirana and M. rugosa contributed the most to the difference in communities between sample methods, collectively contributing 25% to total between method community variation (Supplementary Information 2). Discussion Comparison of sampling efficiency for two widely used sample methods in detecting ant diversity in a high-elevation montane forest mosaic indicate the sample plot methods are more efficient and productive than pitfall trapping for collecting ant diversity. With data representing three sampling seasons each spanning two months, pitfall traps collected 20% fewer species than a single effort of sample plot sampling (sample plot = 25 species; pitfall trap = 21 species). Using rarefaction analysis, we found the sample plot method estimated a significantly higher site species richness and diversity than pitfall traps. To collect a similar richness using pitfall traps would have required a near doubling of the sampling effort. These findings are consistent with Gotelli et al. ( 2011 ) who identified standardised hand-sampling methods as the most efficient method for ant surveys. Our findings are also consistent with recent studies comparing pitfall trapping and quadrat methods for sampling ants carried out across rainforest habitats (Mbenoun et al. 2021 ) and along a gradients of increasing vegetation disturbance (Fotso Kuate et al. 2015 ) in equatorial Africa, as well as a comparison of pitfall trapping to standardised hand collecting in pine forests in Spain (Abril & Gómez 2013 ). The sample-plot method is an active species-collection method that employs a set of standardised hand-sampling techniques (i.e., quadrat sampling (Bestelmeyer et al. 2000 ), nest excavation (Romero & Jaffe 1989 ), and foliage beating (Harris, Collis & Magar 1972 )) that target available microhabitats such as decaying woody material, litter, surface soil and foliage within a defined space and time (Xu 2002 ). Different sampling methods often yield a distinctive set of species, and many authors advocate using complementary methods to gain the greatest coverage of species, including improved representation of species that are rare, occupy specialised microhabitats, or are patchily distributed (Gotelli et al. 2011 ; Antoniazzi et al. 2020 ). Indeed, we found the sample plot method was slightly more proficient than pitfall trapping at collecting rare ants. Of the 18 ‘rare’ species observed, seven were distinctive to the sample plot method in comparison to the four species unique to pitfall trapping. However, surprisingly, the representative range of ecological niches observed (Gotelli et al. 2011 ; Mark & Guenard 2017) appeared remarkably undifferentiated between the two sampling methods (Supplementary Information 3). Both methods captured species that nest and forage across a range of microhabitats soil, litter and canopy. Pitfall traps target epigaeic (surface and litter foraging) ants; they are selective and spatially constrained, a problem that becomes more pronounced with increasingly complex habitats (Luff 1975 ; Majer 1997 ; Gotelli & Colwell 2001 ; Gotelli et al. 2011 ). Still, numerous studies suggest pitfall traps are a preferred method for comparing ant assemblages among some habitats (Steiner et al. 2005 ; Oliveira et al. 2009 ; Hoffmann & Pettit 2022 ) or are an effective complement to other methods as part of an integrated survey (Bestelmeyer et al. 2000 ; de Souza et al. 2012 ). Several authors have highlighted the need to identify added information gained from additional sampling methods to reduce sampling effort, costs, and the risk of sampling redundancy (Tista & Fiedler 2011 ; de Souza et al. 2012 ). We found pitfall traps were generally cheap and easy to install; they were effective in estimating ant species alpha and beta richness, capturing unique species, and identifying a distinctive ant assemblage, as determined by the presence of non-shared species (PERMANOVA, p < 0.05). However, if we increased the sampling effort of the sampling plot method to equal the sampling completeness of the pitfall trapping (95%), we would expect an increase of shared species, as well as further rare, cryptic or specialist species (Gotelli et al. 2011 ; Antoniazzi et al. 2020 ). Only with additional survey could pitfall trapping be confirmed redundant to sample plots in this habitat. The pitfall trap method was, on a sampling unit basis, more time-consuming and labour intensive than the sample plot method, since traps had to be cleared and maintained regularly and samples were generally slower to process in the lab due to sample bycatch, litterfall and detritus in the sample. In contrast, sampling plot methods were completed in the field in two hours and collections returned to the lab ready for identification. Our experience highlighted the efficiency of methods is not independent of the site location and environment; travel time to the traps was an important constituent to efficiency. The sample plot method does have limitations. Sample plot surveys can only be conducted in the daytime (i.e., adequate lighting) and in fair weather (Li et al. 2015b ). There may also be long-term effects from destructive sampling, an important consideration for any monitoring in sensitive environments (Bowie & Frampton 2004 ; Zaller et al. 2015 ). Also, consideration needs to be given to its susceptibility to differences in collector efficiency or expertise (Longino, Coddington & Colwell 2002 ; Sørensen, Coddington & Scharff 2009 ; Gotelli et al. 2011 ; Antoniazzi et al. 2020 ). In this study, ant samples were collected by a student with limited expertise and training, indicating the method has wide applications for monitoring high-elevation forests. For remote sites, particularly in areas with prominent local or indigenous groups presence, such as Lasha Mountain, a test will be if long-term biological monitoring can be facilitated by interested locals. Lasha Mountain has communities representing multiple ethnic nationalities that depend on the local biodiversity and the sustainable management of grazing land, soil, and water for their livelihoods. Establishing robust methods that can involve indigenous or local people can benefit research as well as improve situations for local people. This study demonstrates the sample plot method, a standardised hand-sampling methodology, is more efficient and productive than pitfall trapping for ants in this high-elevation forest system. Where resources limitations restrict the scope of sampling methods, the sample plot method is the most appropriate choice, providing a higher sampling efficiency on a sample unit basis. However, our results indicate pitfall trapping provides additional information on ant assemblages not detected in sample plot methods, indicating that where possible, adding complementary methods, such as pitfall trapping, to the sampling protocol improves sensitivity and accuracy of long-term biological monitoring. Declarations Acknowledgments The authors would like to express their gratitude to the Institute of Eastern-Himalaya Biodiversity Research and the Administration of Yunling Provincial Nature Reserve. We gratefully acknowledge the invaluable help of Professor. Zheng-Hui Xu, Southwest Forestry College for the species identification. References Abril S, Gómez C (2013) Rapid assessment of ant assemblages in public pine forests of the central Iberian Peninsula. For Ecol Manag 293:79–84 Adis J (1979) Problems of interpreting arthropod sampling with pitfall traps. Zoologischer Anzeiger - A Journal of Comparative Zoology 202:177–184 Agosti D, Alonso LE (2000) The ALL protocol: A standard protocol for the collection of ground-dwelling ants. Ants: Standard Methods for Measuring and Monitoring Biodiversity (ed. D.E. Wilson), pp. 204–206 Agosti D, Majer JD, Alonso LE, Schultz T (2000) Ants: standard methods for measuring and monitoring biodiversity. In: Wilson DE (ed) Biological Diversity Handbook Series. Smithsonian Institution, Washington DC, p 280 Andersen AN (1997) Using ants as bioindicators: multiscale issues in ant community ecology.Conservation ecology, 1 Andersen AN, Fisher A, Hoffmann BD, Read JL, Richards R (2004) Use of terrestrial invertebrates for biodiversity monitoring in Australian rangelands, with particular reference to ants. Austral Ecol 29:87–92 Andersen AN, Hoffmann BD, Müller WJ, Griffiths AD (2002) Using ants as bioindicators in land management: simplifying assessment of ant community responses. J Appl Ecol 39:8–17 Andersen AN, Majer JD (2004) Ants show the way Down Under: invertebrates as bioindicators in land management. Front Ecol Environ 2:291–298 Antoniazzi R, Ahuatzin DA, Pelayo-Martínez J, Ortiz-Lozada L, Leponce M, Dáttilo W (2020) On the effectiveness of hand collection to complement baits when studying ant vertical stratification in tropical rainforests. Sociobiology 67:213–222 Bates D, Mächler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1–48 Bestelmeyer BT, Agosti D, Alonso LE, Brandao CRF, Brown WL, Delabie JHC, Silvestre R (2000) Field techniques for the study of ground-dwelling ants: an overview, description, and evaluation. Ants: Standard Methods for Measuring and Monitoring Biodiversity , pp. 122–144 Bowie MH, Frampton CM (2004) A practical technique for non-destructive monitoring of soil surface invertebrates for ecological restoration programmes. Ecol Manag Restor 5:34–42 Bray D (2014) Modification of the ALL protocol to characterise the overall ant assemblage in temperate eucalypt forest. Australian Entomol 41:15–28 Cardoso P, Erwin TL, Borges PAV, New TR (2011) The seven impediments in invertebrate conservation and how to overcome them. Biol Conserv 144:2647–2655 Chao A, Chiu C-H, Jost L (2014) Unifying species diversity, phylogenetic diversity, functional diversity, and related similarity and differentiation measures through Hill numbers. Annu Rev Ecol Evol Syst 45:297–324 Chao A, Gotelli NJ, Hsieh TC, Sander EL, Ma KH, Colwell RK, Ellison AM (2014) Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecol Monogr 84:45–67 Chao A, Jost L (2012) Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93:2533–2547 Clarke KR (1993) Non-parametric multivariate analyses of changes in community structure. Aust J Ecol 18:117–143 Clarke KR, Somerfield PJ, Chapman MG (2006) On resemblance measures for ecological studies, including taxonomic dissimilarities and a zero-adjusted Bray–Curtis coefficient for denuded assemblages. J Exp Mar Biol Ecol 330:55–80 Ding YQ (1980) Principles of Mathematical Ecology of Insect Populations. Science Press, Beijing de Souza JLP, Baccaro FB, Landeiro VL, Franklin E, Magnusson WE (2012) Trade-offs between complementarity and redundancy in the use of different sampling techniques for ground-dwelling ant assemblages. Appl Soil Ecol 56:63–73 Fang H, Xu ZH (2021) Formicidae insects research of China based on bibliometrics. J Southwest Forestry Univ 41:14 Fotso Kuate A, Hanna R, Tindo M, Nanga S, Nagel P (2015) Ant Diversity in Dominant Vegetation Types of Southern Cameroon. Biotropica 47:94–100 Gerlach J, Samways M, Pryke J (2013) Terrestrial invertebrates as bioindicators: an overview of available taxonomic groups. J Insect Conserv 17:831–850 Gotelli NJ, Colwell RK (2001) Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecol Lett 4:379–391 Gotelli NJ, Ellison AM, Dunn RR, Sanders NJ (2011) Counting ants (Hymenoptera: Formicidae): Biodiversity sampling and statistical analysis for myrmecologists. Myrmecological News 15:13–19 Guerra CA, Bardgett RD, Caon L, Crowther TW, Delgado-Baquerizo M, Montanarella L, Navarro LM, Orgiazzi A, Singh BK, Tedersoo L, Vargas-Rojas R, Briones MJI, Buscot F, Cameron EK, Cesarz S, Chatzinotas A, Cowan DA, Djukic I, Van Den Hoogen J, Lehmann A, Maestre FT, Marín C, Reitz T, Rillig MC, Smith LC, De Vries FT, Weigelt A, Wall DH, Eisenhauer N (2021) Tracking, targeting, and conserving soil biodiversity: A monitoring and indicator system can inform policy. Science 371:239–241 Harris JWE, Collis DG, Magar KM (1972) Evaluation of the tree-beating method for sampling defoliating forest insects. Can Entomol 104:723–729 Hervé M (2018) RVAideMemoire: testing and plotting procedures for biostatistics. R package version 0.9–69 , 3 Hobbie JE, Carpenter SR, Grimm NB, Gosz JR, Seastedt TR (2003) The US Long Term Ecological Research program. Bioscience 53:21–32 Hoffmann BD, Pettit M (2022) Towards precision ecology: Relationships of multiple sampling methods quantifying abundance for comparisons among studies.Plos One, 17 Hölldobler B, Wilson EO (1990) The ants. Harvard University Press Hsieh TC, Ma KH, Chao A (2016) iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods Ecol Evol 7:1451–1456 Huang ZP, Scott MB, Li YP, Ren GP, Xiang ZU, Cui LW, Xiao W (2017) Black-and-white snub-nosed monkey ( Rhinopithecus bieti ) feeding behavior in a degraded forest fragment: clues to a stressed population. Primates ecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (2019) In: Brondizio ES, Settele J, Díaz S, Ngo HT (eds) Global assessment report on biodiversity and. IPBES secretariat, Bonn, Germany, p 1148 Jiménez-Carmona F, Carpintero S, Reyes-López JL (2020) Ant sampling: the importance of pitfall trap depth as a bias factor. Entomol Exp Appl 168:703–709 Kuussaari M, Bommarco R, Heikkinen RK, Helm A, Krauss J, Lindborg R, Öckinger E, Pärtel M, Pino J, Rodà F, Stefanescu C, Teder T, Zobel M, Steffan-Dewenter I (2009) Extinction debt: a challenge for biodiversity conservation. Trends in Ecology and Evolution 24:564–571 Larsson J (2021) eulerr: Area-Proportional Euler and Venn Diagrams with Ellipses Length RV (2022) emmeans: Estimated Marginal Means, aka Least-Squares Means Li AN, Xu ZH, Xu GL, Zheng Y, Zhuang JX, Li B (2017) Ant diversity of Tongbiguan Nature Reserve and adjacent area in Yunnan. J Southwest Forestry Univ 37:135–141 Li N, Huang Z-P, Xiao W, Cui L-W (2019) Seed dispersal by black-and-white snub-nosed monkey ( Rhinopithecus bieti ) at Lashashan, Yunnan, China. Chinese Journal of Ecology , 8 , 2019 Li Q, Chen YQ, Xu ZH (2009) Research methods on ant community. Chin J Ecol 28:1862–1870 Li Q, Lu ZX, Zhang W, Ma YY, Feng P (2015a) Communities of ground-dwelling ants in different plantation forests in arid-hot valleys of Jinsha River, Yunnan Province. China Scientia Silvae Sinicae 51:137–145 Li Q, Lu ZX, Zhang W, Ma YY, Feng P (2015b) Ground-dwelling ants as bioindicators during 30-year vegetation restoration in a savanna area, Yunnan. Acta Ecol Sin 35:6199–6207 Lindenmayer DB, Likens GE (2010) The science and application of ecological monitoring. Biol Conserv 143:1317–1328 Longino JT, Coddington J, Colwell RK (2002) The ant fauna of a tropical rain forest: estimating species richness three different ways. Ecology 83:689–702 Longino JT, Colwell RK (1997) Biodiversity assessment using structured inventory: Capturing the ant fauna of tropical rainforest. Ecol Appl 7:1263–1277 Lopes CT, Vasconcelos HL (2008) Evaluation of three methods for sampling ground-dwelling ants in the Brazilian Cerrado. Neotrop Entomol 37:399–405 Luff ML (1975) Some features influencing the efficiency of pitfall traps. Oecologia 19:345–357 Majer J (1997) The use of pitfall traps for sampling ants - A critique. Mem Museum Vic 56:323–329 Mark K, Guenard B, Formicidae (2017) Myrmecological News , 25 , 1–16 Mbenoun PS, Tadu Z, Djieto Lordon C, Mony R, Kenne M, Tindo M (2021) Efficiency of sampling methods for capturing soil-dwelling ants in three landscapes in southern Cameroon. Soil Organisms 93:115–132 McGeoch MA (1998) The selection, testing and application of terrestrial insects as bioindicators. Biol Rev Camb Philos Soc 73:181–201 Myers N, Mittermeler RA, Mittermeler CG, Da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403:853–858 Navarro LM, Fernández N, Guerra C, Guralnick R, Kissling WD, Londoño MC, Muller-Karger F, Turak E, Balvanera P, Costello MJ, Delavaud A, Serafy E, Ferrier GY, Geijzendorffer S, Geller I, Jetz GN, Kim W, Kim ES, Martin H, McGeoch CS, Mwampamba MA, Nel TH, Nicholson JL, Pettorelli E, Schaepman N, Skidmore ME, Pinto ASousa, Vergara I, Vihervaara S, Xu P, Yahara H, Gill T, Pereira HM (2017) Monitoring biodiversity change through effective global coordination. Curr Opin Environ Sustain 29:158–169 Noss RF (1990) Indicators for Monitoring Biodiversity: A Hierarchical Approach. Conserv Biol 4:355–364 Oksanen J, Blanchet F, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin P, O’Hara R, Simpson G, Solymos P (2019) vegan: Community Ecology Package Oliveira M, Lucia TMC, Marinho C, Delabie J, Morato ER (2009) Ant diversity in an area of the Amazon Forest in acre, Brazil. Sociobiology 54:243–267 Parr C, Eggleton P, Davies A, Evans TA, Holdsworth S (2016) Suppression of savanna ants alters invertebrate composition and influences key ecosystem processes. Ecology 97:1611–1617 Parr CL, Chown SL (2001) Inventory and bioindicator sampling: testing pitfall and Winkler methods with ants in a South African savanna. J Insect Conserv 5:27–36 Purvis A, Hector A (2000) Getting the measure of biodiversity. Nature 405:212–219 R Core Team (2020) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria Romero H, Jaffe K (1989) A comparison of methods for sampling ants (Hymenoptera, Formicidae) in savannas. Biotropica 21:348–352 Schultheiss P, Nooten S, Wang R, Wong M, Brassard F, Guénard B (2022) The abundance, biomass, and distribution of ants on Earth. Proceedings of the National Academy of Sciences , 119 , e2201550119 Sørensen L, Coddington J, Scharff N (2009) Inventorying and Estimating Subcanopy Spider Diversity Using Semiquantitative Sampling Methods in an Afromontane Forest. Environ Entomol 31:319–330 Steiner FM, Schlick-Steiner BC, Moder K, Bruckner A, Christian E (2005) Congruence of data from different trapping periods of ant pitfall catches (Hymenoptera: Formicidae). v . 46 Tiede Y, Schlautmann J, Donoso DA, Wallis CI, Bendix J, Brandl R, Farwig N (2017) Ants as indicators of environmental change and ecosystem processes. Ecol Ind 83:527–537 Topping CJ, Sunderland KD (1992) Limitations to the use of pitfall traps in ecological studies exemplified by a study of spiders in a field of winter wheat. J Appl Ecol 29:485–491 Tremblay A, Ransijn J (2015) LMERConvenienceFunctions: Model selection and post-hoc analysis for (G) LMER models. R package version , 2 Tista M, Fiedler K (2011) How to evaluate and reduce sampling effort for ants. J Insect Conserv 15:547–559 Underwood EC, Fisher BL (2006) The role of ants in conservation monitoring: if, when, and how. Biol Conserv 132:166–182 Vineesh PJ, Sabu TK, Karmaly KA (2007) Community structure and functional group classification of litter ants in the montane evergreen and deciduous forests of Wayanad region of Western Ghats, Southern India. Orient Insects 41:427–442 Wang SJ, Huang ZP, He YC, He XD, Li DH, Sun J, Cui LW, Xiao W (2012) Mating behavior and birth seasonality of black-and-white snub-nosed monkeys ( Rhinopithecus bieti ) at Mt. Lasha. Zoological Res 33:241–248 Ward DF, New TR, Yen AL (2001) Effects of pitfall trap spacing on the abundance, richness and composition of invertebrate catches. J Insect Conserv 5:47–53 Xu Z (2002) A study on the biodiversity of Formicidae ants of Xishuangbanna Nature Reserve. Yunnan Science and Technology Press, Yunnan Yeo K, Delsinne T, Konate S, Alonso LL, Aïdara D, Peeters C (2017) Diversity and distribution of ant assemblages above and below ground in a West African forest–savannah mosaic (Lamto, Côte d’Ivoire). Insectes Sociaux 64:155–168 Zaller JG, Kerschbaumer G, Rizzoli R, Tiefenbacher A, Gruber E, Schedl H (2015) Monitoring arthropods in protected grasslands: Comparing pitfall trapping, quadrat sampling and video monitoring. Web Ecol 15:15–23 Zhang HY, Ou XH (2006) Using insect for indicator to monitor and assess forest ecosystem health. World Forestry Research 19:22–25 Additional Declarations No competing interests reported. Supplementary Files SupplementaryInformation.pdf Cite Share Download PDF Status: Published Journal Publication published 09 Aug, 2023 Read the published version in Journal of Insect Conservation → Version 1 posted Editorial decision: Major revision 27 Dec, 2022 Reviews received at journal 05 Dec, 2022 Reviewers agreed at journal 25 Nov, 2022 Reviewers invited by journal 23 Nov, 2022 Editor assigned by journal 11 Nov, 2022 Submission checks completed at journal 11 Nov, 2022 First submitted to journal 10 Nov, 2022 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-2261097","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":151426327,"identity":"977990a8-df81-44dc-83d6-514fe90ed524","order_by":0,"name":"Chuan-Jing Zhang","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Chuan-Jing","middleName":"","lastName":"Zhang","suffix":""},{"id":151426328,"identity":"350fdab9-ab96-4fdf-ac8e-37ceae9da76d","order_by":1,"name":"Yi-Ting Cheng","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yi-Ting","middleName":"","lastName":"Cheng","suffix":""},{"id":151426330,"identity":"d06178eb-bd54-4b25-94b3-38afbd32214f","order_by":2,"name":"Xian-Shu Luo","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Xian-Shu","middleName":"","lastName":"Luo","suffix":""},{"id":151426332,"identity":"c9f1ef24-c955-4653-a92a-29cceb44d245","order_by":3,"name":"Yao Chen","email":"","orcid":"","institution":"Administration of Yunling Provincial Nature Reserve","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yao","middleName":"","lastName":"Chen","suffix":""},{"id":151426334,"identity":"473c68d9-3b26-4ae1-a9ca-138f79632f87","order_by":4,"name":"Yu-Chao He","email":"","orcid":"","institution":"Administration of Yunling Provincial Nature Reserve","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yu-Chao","middleName":"","lastName":"He","suffix":""},{"id":151426335,"identity":"e23bf2ec-d6ff-4aa9-8aea-81ce438034e4","order_by":5,"name":"Yan-Pang Li","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Yan-Pang","middleName":"","lastName":"Li","suffix":""},{"id":151426336,"identity":"c296d9b2-f798-4cc0-af70-ca22d27d8e1d","order_by":6,"name":"Zhi-Pang Huang","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Zhi-Pang","middleName":"","lastName":"Huang","suffix":""},{"id":151426337,"identity":"6a035ba9-8644-400e-a3d8-2f7db11c0ae9","order_by":7,"name":"Matthew B. Scott","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAABAklEQVRIiWNgGAWjYHAD5gMMDAYIbgIelYwNIIqHjS2BZC08BijiOLXotp8//oDhj52cvXzP5w8/CmzyzNvbHz74uccuj4H98NENWLSYnUlmbGBsSzbmYePdJtljkFYsc+aMsWHPs+RiBp60tBvYtBwAaWk4kNgD1MLMYHA4cYZEDpsEz4EDiQ0SPGZYtZx/DPTLH5AWnsefwVrknz//+QeflhtAWxjYwFoYpCG2MJgx47XlxmPDGYkgvxxLMwP5JXEGT46xtMyB5MQ2XH45n/jgwwdgiLE3H3784ccfm8QZ7McffnxzwC6xn/3wMWxawCABqygbLuWjYBSMglEwCggCAJ4+Zj0SaaQXAAAAAElFTkSuQmCC","orcid":"","institution":"New Zealand Forest Research Institute Ltd (Scion)","correspondingAuthor":true,"submittingAuthor":false,"prefix":"","firstName":"Matthew","middleName":"B.","lastName":"Scott","suffix":""},{"id":151426338,"identity":"7b1a675b-b1f8-42b0-aa65-f76c6e73c329","order_by":8,"name":"Wen Xiao","email":"","orcid":"","institution":"Institute of Eastern-Himalaya Biodiversity Research","correspondingAuthor":false,"submittingAuthor":false,"prefix":"","firstName":"Wen","middleName":"","lastName":"Xiao","suffix":""}],"badges":[],"createdAt":"2022-11-11 01:29:08","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-2261097/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-2261097/v1","draftVersion":[],"editorialEvents":[{"content":"https://doi.org/10.1007/s10841-023-00501-y","type":"published","date":"2023-08-09T21:57:07+00:00"}],"editorialNote":"","failedWorkflow":false,"files":[{"id":29043622,"identity":"e8fd91c8-e5ab-45f1-b742-c57cb82ec3cd","added_by":"auto","created_at":"2022-11-14 18:18:20","extension":"png","order_by":1,"title":"Figure 1","display":"","copyAsset":false,"role":"figure","size":64108,"visible":true,"origin":"","legend":"\u003cp\u003eNumbers of unique and common ant species for pitfall trap (PF) and sample plot (SP) samples are depicted in a Venn diagram for All, ‘Dominant’, ‘Common’, and ‘Rare’ species.\u003c/p\u003e","description":"","filename":"Figure1.png","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/350b3950d335eb75e6e4dd8f.png"},{"id":29044385,"identity":"eac61cd5-aee5-4eca-a83f-43a0867b8b15","added_by":"auto","created_at":"2022-11-14 18:26:20","extension":"png","order_by":2,"title":"Figure 2","display":"","copyAsset":false,"role":"figure","size":25597,"visible":true,"origin":"","legend":"\u003cp\u003eEffect of method to total alpha (species richness) and beta diversity. PF: pitfall trap; SP: sample plot. Values are means ± standard errors. Differences of estimated marginal means according to Tukey post-hoc tests were not found significant; \u003cem\u003en.s.\u003c/em\u003e: not significant.\u003c/p\u003e","description":"","filename":"Figure2.png","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/c39d770892f5e575ee93bf51.png"},{"id":29045133,"identity":"63319e9b-3f8b-431c-a31c-e3266ff1a70e","added_by":"auto","created_at":"2022-11-14 18:34:21","extension":"png","order_by":3,"title":"Figure 3","display":"","copyAsset":false,"role":"figure","size":158099,"visible":true,"origin":"","legend":"\u003cp\u003eSample-based rarefaction (solid lines) and extrapolation (dotted lines) of ants collected using pitfall trap (PF) and sample-plot (SP) methods, with 95% unconditional confidence intervals (shading). Diversity was estimated for species richness, exponential Shannon’s index, and inverse Simpson’s index. (A) A comparison of estimated asymptotic or true diversities for two methods (PF and SP); (B) a comparison of estimated point diversities for sampling units for methods PF and SP; (C) an assessment of sample completeness (sample coverage) across increasing sample units.\u003c/p\u003e","description":"","filename":"Figure3.png","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/2e786fd35d2e3986ef77f91c.png"},{"id":29043623,"identity":"58624a6c-3edb-4bd4-860c-ce1a5f5e94c5","added_by":"auto","created_at":"2022-11-14 18:18:20","extension":"png","order_by":4,"title":"Figure 4","display":"","copyAsset":false,"role":"figure","size":54859,"visible":true,"origin":"","legend":"\u003cp\u003eNMDS plot for ants at Lasha Mountain using two methods, pitfall trapping (PF) and sample plot (SP). The ellipses represent 95% confidence limits around the centroids of each method.\u003c/p\u003e","description":"","filename":"Figure4.png","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/4f081777271a7ed76fa87b9f.png"},{"id":44735102,"identity":"4558023a-be80-4045-b53a-08905e5a1070","added_by":"auto","created_at":"2023-10-16 22:23:39","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":588863,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/f09a16d6-4c58-4c28-9e69-5279366b64d6.pdf"},{"id":29043626,"identity":"d73b9de4-05f2-4fda-8c9a-6241dca97411","added_by":"auto","created_at":"2022-11-14 18:18:21","extension":"pdf","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":134122,"visible":true,"origin":"","legend":"","description":"","filename":"SupplementaryInformation.pdf","url":"https://assets-eu.researchsquare.com/files/rs-2261097/v1/e402c59b981849481a1d9b56.pdf"}],"financialInterests":"No competing interests reported.","formattedTitle":"A comparison of two methods for quantifying ant diversity and community in an East Himalayan montane forest mosaic","fulltext":[{"header":"Introduction","content":"\u003cp\u003eGlobal biodiversity is declining at an unprecedented rate (IPBES 2019). Monitoring biodiversity is a key part of addressing the trend by providing facts of the occurring changes and mechanisms of change to bring about science-based policy and land management decisions that benefit biodiversity (Noss \u003cspan citationid=\"CR56\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Hobbie et al. \u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2003\u003c/span\u003e; Kuussaari et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Cardoso et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Navarro et al. \u003cspan citationid=\"CR55\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Guerra et al. \u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). The continuing challenge is measuring biodiversity in ways that are useful, accurate, and repeatable (Purvis \u0026amp; Hector \u003cspan citationid=\"CR61\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Lindenmayer \u0026amp; Likens \u003cspan citationid=\"CR45\" class=\"CitationRef\"\u003e2010\u003c/span\u003e). Ants, in particular, are an insect group of immense ecological significance, contributing a large proportion of the global arthropod biomass (Schultheiss et al. \u003cspan citationid=\"CR64\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and having critical ecological functions such as nutrient cycling, soil aeration, and seed dispersal (H\u0026ouml;lldobler \u0026amp; Wilson \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e1990\u003c/span\u003e; Parr et al. \u003cspan citationid=\"CR59\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Ants are relatively easy to collect, sensitive to environmental change and representative of soil diversity (Andersen et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Andersen \u0026amp; Majer \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015b\u003c/span\u003e). As such, ants have long been an important insect group for biological monitoring and indicators of ecological change (bioindicators) (Andersen \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; McGeoch \u003cspan citationid=\"CR53\" class=\"CitationRef\"\u003e1998\u003c/span\u003e; Andersen et al. \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; Underwood \u0026amp; Fisher \u003cspan citationid=\"CR71\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Zhang \u0026amp; Ou \u003cspan citationid=\"CR78\" class=\"CitationRef\"\u003e2006\u003c/span\u003e; Gerlach, Samways \u0026amp; Pryke \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2013\u003c/span\u003e; Tiede et al. \u003cspan citationid=\"CR67\" class=\"CitationRef\"\u003e2017\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eSeveral sampling methods have been developed to collect ants, each with their own strengths and weaknesses (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). Understanding the limitations to sampling techniques is important to selecting a method, or combinations of methods, that will address survey objectives within a given environment, sampling effort and available expertise (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; de Souza et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). To allow stricter comparison of individual studies, numerous researchers have advocated for the adoption of recognised standard protocols (e.g., Li, Chen \u0026amp; Xu \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Antoniazzi et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Such protocols have been proposed (e.g., Ants of the Leaf Litter (ALL) Protocol: Agosti \u0026amp; Alonso \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2000\u003c/span\u003e) and successfully applied (Vineesh, Sabu \u0026amp; Karmaly \u003cspan citationid=\"CR72\" class=\"CitationRef\"\u003e2007\u003c/span\u003e; Lopes \u0026amp; Vasconcelos \u003cspan citationid=\"CR48\" class=\"CitationRef\"\u003e2008\u003c/span\u003e; Bray \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Yeo et al. \u003cspan citationid=\"CR76\" class=\"CitationRef\"\u003e2017\u003c/span\u003e) but universal application has lagged.\u003c/p\u003e \u003cp\u003eIn China, the \u0026lsquo;sample-plot\u0026rsquo; is the most used method, followed by pitfall traps. Pitfall traps are small, covered pits set in the ground that target active epigaeic fauna (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). The widely-used method provides relatively simple and cost-effective sampling of epigaeic fauna that allows for continuous sampling for a time period, both day and night (Majer \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Pitfall traps suffer for their sensitivity to species size, trappability and activity (Topping \u0026amp; Sunderland \u003cspan citationid=\"CR68\" class=\"CitationRef\"\u003e1992\u003c/span\u003e; Majer \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1997\u003c/span\u003e), as well as biases to do with trap diameter, trap depth, spacing and habitat complexity (Luff \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Adis \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e1979\u003c/span\u003e; Ward, New \u0026amp; Yen \u003cspan citationid=\"CR74\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Jim\u0026eacute;nez-Carmona, Carpintero \u0026amp; Reyes-L\u0026oacute;pez \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). The sample plot method\u0026mdash;a modified \u0026lsquo;direct sampling\u0026rsquo; method (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e)\u0026mdash;is a technique where specimens are collected systematically from microhabitats using a range of hand-collecting techniques within a specified area and time (Xu \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). The method has advantages in efficiency but is vulnerable to the competence of the researchers and differences in habitats, reducing the comparability between samples and studies (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eHere, we aimed to evaluate these two widely used ant sample methods in China by assessing their results in terms of ant diversity and community composition in a forest mosaic in Yunling Nature Reserve, Yunnan Province. We expected that sample plot methods would collect more ant species than pitfall traps, where hand sorting could allow for sampling of more cryptic, submissive, and less abundant ants (Parr \u0026amp; Chown \u003cspan citationid=\"CR60\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Longino, Coddington \u0026amp; Colwell \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; de Souza et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Mark \u0026amp; Guenard 2017). We also expected that sampling methods may selectively filter the ant community present so that combining methods could improve the survey and generate a closer estimation of true ant community (Longino \u0026amp; Colwell \u003cspan citationid=\"CR47\" class=\"CitationRef\"\u003e1997\u003c/span\u003e). Our results will provide a measure of the sampling effort required and reliability of the combination of methods to characterise and monitor ground-dwelling ant diversity in a montane forest mosaic.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eThe study was conducted at Lasha Mountain (拉沙山) (N26\u0026deg;20', E99\u0026deg;15') in Yunling Nature Reserve, Yunnan, China, within the Mountain Area of Southwest China biodiversity hotspot (Myers et al. \u003cspan citationid=\"CR54\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). At 75,894 km\u003csup\u003e2\u003c/sup\u003e, the Yunling Nature Reserve comprises a mosaic of land covers, including forest, regenerating forest, pastoral grazing land, cropland, and permanent settlements. The study area is a ca. 1400 ha catchment ranging from 2500 to 3700 m asl. Forest vegetation dominates most of the mid-elevations, transitioning from deciduous broadleaved forest (\u003cem\u003eBetula alnoides, Acanthopanax gracilistylus, Acer oliverianum\u003c/em\u003e) at lower elevations, through mixed deciduous-conifer forest to conifer forest (\u003cem\u003eAbies georgei, A. fabri, Tsuga dumosa\u003c/em\u003e) at the highest elevations. The forest undergrowth primarily comprises \u003cem\u003eRhododendron\u003c/em\u003e spp. and bamboo (\u003cem\u003eFargesia strigosa, F. edulis, F. solida\u003c/em\u003e). Few local families live in the lowest part of the catchment where there is a mixture of grazing, cropland and forest margins. The forest is used by local people for resources such a wood, traditional foods and medicine (Huang et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). The climate is characterised by alternating dry and wet seasons, with an annual mean precipitation of 910 mm (Wang et al. \u003cspan citationid=\"CR73\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Li et al. \u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec4\" class=\"Section2\"\u003e \u003ch2\u003eSample design\u003c/h2\u003e \u003cp\u003eWe sampled ants using two common sampling methods (pitfall trap and sample plot) across the study area. Sample sites were established every 200 m elevation from 2500 m to 3700 m (totalling 7 sampling sites) using the stream as a centreline. Each sample site comprised a 60 m \u0026times; 60 m plot, evenly divided across the stream. Five pitfall traps were stratified across each plot. The open-topped attractant bottles (11 cm in diameter, 15.5 cm in height) were embedded into the ground until flush with the natural soil surface and covered with 15-cm diameter plastic plates to keep rain out. Traps were baited with 50\u0026ndash;70 ml liquid attractant composed of 1:1:4:16 solution of ethanol, sugar, vinegar, water (Li et al. \u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Fang \u0026amp; Xu \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Trap surveys were conducted from August to September in 2018, 2019 and 2020. We emptied traps every two weeks during each sampling period.\u003c/p\u003e \u003cp\u003eSample plot sampling was undertaken in each 60 m \u0026times; 60 m plot at five 5 m \u0026times; 5 m sampling sub-plots. Subplots were paired with pitfall traps, buffered by 10 m. After an initial surface inspection, ants were collected by hand from litter, decaying wood and under stones for 1 hr (1 person hour). Using hand tools, soil was excavated to 20 cm to search for and collect ants from subterranean nests. A maximum thirty individuals were collected from a single nest. Foliage collections were made using a 2 m \u0026times; 2 m white curtain was placed flat around the sample sub-plot and small trees and shrubs up to 5 m were vigorously shaken or beaten to capture ants that fell on the curtain. In total, each plot was investigated for 2 hr.\u003c/p\u003e \u003cp\u003eCollected specimens from both methods were stored in 75% ethanol and transferred to a freezer at -10\u0026deg;C (Xu \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2002\u003c/span\u003e) before being identified.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec5\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eAll analyses were performed using R v4.2.0 (R Core Team \u003cspan citationid=\"CR62\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). To reduce the effects of spatial autocorrelation of response variables, we pooled pitfall trap data by site and year. Sampling yielded 28 sampling units in total from 21 pitfall trap units and 7 sample plot units. For community analysis, we reduced data to presence-absence to reduce the effects of ant biology that lead to non-random distribution at some scales (Agosti et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2000\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo visualise the number of species of ants detected with each method, a Venn diagram was drawn using the \u0026lsquo;eulerr\u0026rsquo; package in R (Larsson \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). We drew these for all species, as well as species classed as \u0026ldquo;dominant\u0026rdquo;, \u0026ldquo;common\u0026rdquo;, and \u0026ldquo;rare.\u0026rdquo; We defined dominant species as those with a total relative abundance exceeding 10%; common species contributed between 1% and 10% total abundance, and rare species contributed less than 1% to the total population sampled (Li et al. \u003cspan citationid=\"CR43\" class=\"CitationRef\"\u003e2015a\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe effects of method on total measured alpha (α) and beta (β) diversity were tested using a Linear Mixed-Effects Model, using the function \u003cem\u003elmer\u003c/em\u003e in the \u0026lsquo;lme4\u0026rsquo; package in R (Bates et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) with a parametric analysis of variance (ANOVA). Method blocked into year and elevation were introduced as random effects in the model. Normality and homogeneity of variance were checked by performing Quartile-Quartile (QQ) plots of the residuals and fitted models using the function \u003cem\u003eplotresid\u003c/em\u003e in the package \u0026lsquo;RVAideMemoire\u0026rsquo; (Herv\u0026eacute; \u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). We identified and removed outliers in the model using the function \u003cem\u003eromr.fnc\u003c/em\u003e in the package \u0026lsquo;LMERConvenienceFunctions\u0026rsquo; (Tremblay \u0026amp; Ransijn \u003cspan citationid=\"CR69\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Pairwise comparison of estimated marginal means (EMMs) were performed using the \u0026lsquo;emmeans\u0026rsquo; package (Length \u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eWe used rarefaction analysis to compare estimated asymptotic species diversity between the sample plot and pitfall trapping methods, a technique based on species frequency (i.e., presence-absence) that avoids biases caused by insufficient or differing sampling efforts (Gotelli \u0026amp; Colwell \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). To do this, we used sample-sized-based rarefaction-extrapolation analysis based on sampling-unit-based incidence data (Chao et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e). Hill species diversity (or the effective number of species) in the zero (q\u0026thinsp;=\u0026thinsp;0), first (q\u0026thinsp;=\u0026thinsp;1), and second (q\u0026thinsp;=\u0026thinsp;2) orders (or species richness, the exponential Shannon entropy, and inverse Simpson index, respectively) were computed in the \u003cem\u003eiNEXT\u003c/em\u003e package (Hsieh, Ma \u0026amp; Chao \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). In addition, we compared sampling coverage (based on incidence) of each method across Hill species diversities to our sampling effort, permitting estimation of the proportion of the total community represented by the sampling effort and an assessment of sample completeness (sample coverage) across sampling units (Chao \u0026amp; Jost \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2012\u003c/span\u003e; Chao et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2014\u003c/span\u003e; Hsieh, Ma \u0026amp; Chao \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2016\u003c/span\u003e). Significant differences in estimated diversity and sampling coverage between methods were judged by non-overlapping confidence intervals (Chao, Chiu \u0026amp; Jost \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e2014\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eTo evaluate differences in ant composition of samples obtained from the two methods, we performed non-metric multidimensional scaling (NMDS: calculated with Jaccard dissimilarity index) with the function \u003cem\u003emetaMDS\u003c/em\u003e of the R package \u0026lsquo;vegan\u0026rsquo; (Oksanen et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). We considered a species x sample matrix with presence-absence data (30 species x 28 samples). A dummy species was added to the species matrix to mitigate the impact of pitfall traps that had zero abundances (Clarke, Somerfield \u0026amp; Chapman \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2006\u003c/span\u003e). We used the function \u003cem\u003eordiellipse\u003c/em\u003e of the vegan package to draw ellipses representing 95% CI around centroids. To test for differences between groups (methods) we used a permutational multivariate analysis of variance (PERMANOVA) using distance matrices, which was performed with the function \u003cem\u003eadonis2\u003c/em\u003e in vegan (Oksanen et al. \u003cspan citationid=\"CR57\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Elevation was set as a random factor. P values were obtained using 9999 permutations of residuals. To determine which species contributed most to the observed multivariate differences between pitfall trapping and sample plot methods, we used similarity percentage analysis (SIMPER) (Clarke \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e1993\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec7\" class=\"Section2\"\u003e \u003ch2\u003eAnt collection\u003c/h2\u003e \u003cp\u003eUsing two sampling methods, we found a total of 10,206 ant (Insecta: Formicidae) specimens belonging to four subfamilies (Myrmicinae, Formicinae, Dolichoderinae, and Ponerinae), 17 genera and 30 species (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). Of these, 6312 and 3894 individuals were collected from pitfall trap and sample plot methods respectively. We recorded, 21 species from pitfall traps and 25 species from sample plots, of which only 16 species (53%) were shared (Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The most abundant (\u0026lsquo;dominant\u0026rsquo;) species across sampling methods were \u003cem\u003ePheidole nietneri\u003c/em\u003e Emery, 1901, \u003cem\u003eMyrmica rugosa\u003c/em\u003e Mayr, 1865, \u003cem\u003eMyrmica kozlovi\u003c/em\u003e Ruzsky, 1915, \u003cem\u003eMyrmica bactriana\u003c/em\u003e Ruzsky, 1915, \u003cem\u003eFormica fusca\u003c/em\u003e Linnaeus, 1758, together accounting for 70% of all the specimens collected (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e, Fig.\u0026nbsp;\u003cspan refid=\"Fig1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). We recorded 11 rare species from pitfall traps and 14 rare species from sample plots, seven of which were shared. Rare species contributed 5% to the total specimen abundance.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eNumber and account of species in Sample-plot method and Pitfall-trap method. For Classes, D\u0026thinsp;=\u0026thinsp;dominant species (\u0026gt;\u0026thinsp;10% of total); C\u0026thinsp;=\u0026thinsp;common species (1\u0026ndash;10% of total); R\u0026thinsp;=\u0026thinsp;rare species (\u0026lt;\u0026thinsp;1% of total); (%) = % contribution to total abundance.\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eSubfamily/species\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eClass\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003ePitfall Trap\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSample\u003c/p\u003e \u003cp\u003ePlot\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eSum\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003e(%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eDolichoderinae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eTapinoma sinense\u003c/em\u003e Emery, 1925\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e106\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e106\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.04\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003ePonerinae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePonera bawana\u003c/em\u003e Xu, 2001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eMyrmicinae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eAphaenogaster caeciliae\u003c/em\u003e Viehmeyer, 1922\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e100\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e119\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e219\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eAphaenogaster lepida\u003c/em\u003e Wheeler, 1930\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.03\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica bactriana\u003c/em\u003e Ruzsky, 1915\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e490\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e697\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1187\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica jessensis\u003c/em\u003e Forel, 1901\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica kozlovi\u003c/em\u003e Ruzsky, 1915\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e257\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e988\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1245\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e12.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica margaritae\u003c/em\u003e Emery, 1889\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica pararitae\u003c/em\u003e Radchenko, 2008\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica ritae\u003c/em\u003e Emery, 1889\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.6\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmica rugosa\u003c/em\u003e Mayr, 1865\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1344\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e207\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1551\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e15.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eMyrmecina striata\u003c/em\u003e Emery, 1889\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e3\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.03\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePheidole nietneri\u003c/em\u003e Emery, 1901\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1955\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1957\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e19.2\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePheidole pieli\u003c/em\u003e Santschi, 1925\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePerissomyrmex bidentatus\u003c/em\u003e Zhou \u0026amp; Huang, 2006\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e8\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.08\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eTemnothorax\u003c/em\u003e sp.1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eTemnothorax\u003c/em\u003e sp.2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eTetramorium kraepelini\u003c/em\u003e Forel, 1905\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eStenamma bhutanense\u003c/em\u003e Baroni Urbani, 1977\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colspan=\"2\" nameend=\"c2\" namest=\"c1\"\u003e \u003cp\u003eFormicinae\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eCamponotus anningensis\u003c/em\u003e Wu \u0026amp; Wang, 1989\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eCamponotus herculeanus\u003c/em\u003e Linnaeus, 1758\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e109\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.1\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eLasius flavus\u003c/em\u003e Fabricius, 1782\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eLasius coloratus\u003c/em\u003e Santschi, 1937\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e9\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eLasius alienus\u003c/em\u003e Foerster, 1850\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e686\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e273\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e959\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.4\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eNylanderia bourbonica\u003c/em\u003e Forel, 1886\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e2\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.02\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eNylanderia flavipes\u003c/em\u003e Smith, 1874\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e96\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e149\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eParaparatrechina aseta\u003c/em\u003e Forel, 1902\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e202\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e202\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e2.0\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003ePrenolepis angularis\u003c/em\u003e Zhou, 2001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.7\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eFormica fusca\u003c/em\u003e Linnaeus, 1758\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e360\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e816\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1176\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.5\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u003cem\u003eFormica sinensis\u003c/em\u003e Wheeler, 1913\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003eC\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e844\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e0\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e844\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8.3\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003eTotal\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6312\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3894\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e10206\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec8\" class=\"Section2\"\u003e \u003ch2\u003eSpecies diversity\u003c/h2\u003e \u003cp\u003eComparisons of sampled alpha and beta diversity between to two methods did not differ Fig.\u0026nbsp;\u003cspan refid=\"Fig2\" class=\"InternalRef\"\u003e2\u003c/span\u003e); however, our Hill number series analysis (q\u0026thinsp;=\u0026thinsp;0, 1, 2) showed significantly higher species diversity of ants from sample plot methods than pitfall trap methods (non-over-lapping in confidence intervals) for three estimators of diversity across sampling effort and coverage (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003e). Total estimated species richness with iNEXT was 35\u0026thinsp;\u0026plusmn;\u0026thinsp;7.7 s.e. for SP and 25\u0026thinsp;\u0026plusmn;\u0026thinsp;4.7 s.e. for PF (Supplementary Information 1). Results were consistent when comparing diversity at a given sampling effort and coverage (Fig.\u0026nbsp;\u003cspan refid=\"Fig3\" class=\"InternalRef\"\u003e3\u003c/span\u003eB\u0026ndash;C). The sampling coverage method showed an approximate completeness of 76% in sample plot method and 95% in pitfall trap method using the true sampling effort (SP, n\u0026thinsp;=\u0026thinsp;7; PF n\u0026thinsp;=\u0026thinsp;21).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eCommunity composition\u003c/h2\u003e \u003cp\u003eThe NMDS plots based on species presence-absence showed some clustering of the observed species composition according to method (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). PERMANOVA results with elevation held constant showed PF and SP assemblages differed significantly (p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). The NMDS analysis had a low stress value (0.07) which indicates that the ordination summarised the observed distances between samples well (Fig.\u0026nbsp;\u003cspan refid=\"Fig4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). In pitfall traps, we found high numbers of \u003cem\u003ePheidole nieneri\u003c/em\u003e (1955) and \u003cem\u003eMyrmica rugosa\u003c/em\u003e (1344), whereas in sample plots we collected high numbers of \u003cem\u003eMyrmica kozlovi\u003c/em\u003e (988) and \u003cem\u003eFormica fusca\u003c/em\u003e (816) (Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e). The SIMPER analysis showed that \u003cem\u003eMyrmica kozlovi\u003c/em\u003e, \u003cem\u003eM. bactirana\u003c/em\u003e and \u003cem\u003eM. rugosa\u003c/em\u003e contributed the most to the difference in communities between sample methods, collectively contributing 25% to total between method community variation (Supplementary Information 2).\u003c/p\u003e \u003cp\u003e \u003c/p\u003e \u003c/div\u003e"},{"header":"Discussion","content":"\u003cp\u003eComparison of sampling efficiency for two widely used sample methods in detecting ant diversity in a high-elevation montane forest mosaic indicate the sample plot methods are more efficient and productive than pitfall trapping for collecting ant diversity. With data representing three sampling seasons each spanning two months, pitfall traps collected 20% fewer species than a single effort of sample plot sampling (sample plot\u0026thinsp;=\u0026thinsp;25 species; pitfall trap\u0026thinsp;=\u0026thinsp;21 species). Using rarefaction analysis, we found the sample plot method estimated a significantly higher site species richness and diversity than pitfall traps. To collect a similar richness using pitfall traps would have required a near doubling of the sampling effort. These findings are consistent with Gotelli et al. (\u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e) who identified standardised hand-sampling methods as the most efficient method for ant surveys. Our findings are also consistent with recent studies comparing pitfall trapping and quadrat methods for sampling ants carried out across rainforest habitats (Mbenoun et al. \u003cspan citationid=\"CR52\" class=\"CitationRef\"\u003e2021\u003c/span\u003e) and along a gradients of increasing vegetation disturbance (Fotso Kuate et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2015\u003c/span\u003e) in equatorial Africa, as well as a comparison of pitfall trapping to standardised hand collecting in pine forests in Spain (Abril \u0026amp; G\u0026oacute;mez \u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2013\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe sample-plot method is an active species-collection method that employs a set of standardised hand-sampling techniques (i.e., quadrat sampling (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e), nest excavation (Romero \u0026amp; Jaffe \u003cspan citationid=\"CR63\" class=\"CitationRef\"\u003e1989\u003c/span\u003e), and foliage beating (Harris, Collis \u0026amp; Magar \u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e1972\u003c/span\u003e)) that target available microhabitats such as decaying woody material, litter, surface soil and foliage within a defined space and time (Xu \u003cspan citationid=\"CR75\" class=\"CitationRef\"\u003e2002\u003c/span\u003e). Different sampling methods often yield a distinctive set of species, and many authors advocate using complementary methods to gain the greatest coverage of species, including improved representation of species that are rare, occupy specialised microhabitats, or are patchily distributed (Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Antoniazzi et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Indeed, we found the sample plot method was slightly more proficient than pitfall trapping at collecting rare ants. Of the 18 \u0026lsquo;rare\u0026rsquo; species observed, seven were distinctive to the sample plot method in comparison to the four species unique to pitfall trapping. However, surprisingly, the representative range of ecological niches observed (Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Mark \u0026amp; Guenard 2017) appeared remarkably undifferentiated between the two sampling methods (Supplementary Information 3). Both methods captured species that nest and forage across a range of microhabitats soil, litter and canopy.\u003c/p\u003e \u003cp\u003ePitfall traps target epigaeic (surface and litter foraging) ants; they are selective and spatially constrained, a problem that becomes more pronounced with increasingly complex habitats (Luff \u003cspan citationid=\"CR49\" class=\"CitationRef\"\u003e1975\u003c/span\u003e; Majer \u003cspan citationid=\"CR50\" class=\"CitationRef\"\u003e1997\u003c/span\u003e; Gotelli \u0026amp; Colwell \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2001\u003c/span\u003e; Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). Still, numerous studies suggest pitfall traps are a preferred method for comparing ant assemblages among some habitats (Steiner et al. \u003cspan citationid=\"CR66\" class=\"CitationRef\"\u003e2005\u003c/span\u003e; Oliveira et al. \u003cspan citationid=\"CR58\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Hoffmann \u0026amp; Pettit \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) or are an effective complement to other methods as part of an integrated survey (Bestelmeyer et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; de Souza et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). Several authors have highlighted the need to identify added information gained from additional sampling methods to reduce sampling effort, costs, and the risk of sampling redundancy (Tista \u0026amp; Fiedler \u003cspan citationid=\"CR70\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; de Souza et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2012\u003c/span\u003e). We found pitfall traps were generally cheap and easy to install; they were effective in estimating ant species alpha and beta richness, capturing unique species, and identifying a distinctive ant assemblage, as determined by the presence of non-shared species (PERMANOVA, p\u0026thinsp;\u0026lt;\u0026thinsp;0.05). However, if we increased the sampling effort of the sampling plot method to equal the sampling completeness of the pitfall trapping (95%), we would expect an increase of shared species, as well as further rare, cryptic or specialist species (Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Antoniazzi et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). Only with additional survey could pitfall trapping be confirmed redundant to sample plots in this habitat.\u003c/p\u003e \u003cp\u003eThe pitfall trap method was, on a sampling unit basis, more time-consuming and labour intensive than the sample plot method, since traps had to be cleared and maintained regularly and samples were generally slower to process in the lab due to sample bycatch, litterfall and detritus in the sample. In contrast, sampling plot methods were completed in the field in two hours and collections returned to the lab ready for identification. Our experience highlighted the efficiency of methods is not independent of the site location and environment; travel time to the traps was an important constituent to efficiency.\u003c/p\u003e \u003cp\u003eThe sample plot method does have limitations. Sample plot surveys can only be conducted in the daytime (i.e., adequate lighting) and in fair weather (Li et al. \u003cspan citationid=\"CR44\" class=\"CitationRef\"\u003e2015b\u003c/span\u003e). There may also be long-term effects from destructive sampling, an important consideration for any monitoring in sensitive environments (Bowie \u0026amp; Frampton \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2004\u003c/span\u003e; Zaller et al. \u003cspan citationid=\"CR77\" class=\"CitationRef\"\u003e2015\u003c/span\u003e). Also, consideration needs to be given to its susceptibility to differences in collector efficiency or expertise (Longino, Coddington \u0026amp; Colwell \u003cspan citationid=\"CR46\" class=\"CitationRef\"\u003e2002\u003c/span\u003e; S\u0026oslash;rensen, Coddington \u0026amp; Scharff \u003cspan citationid=\"CR65\" class=\"CitationRef\"\u003e2009\u003c/span\u003e; Gotelli et al. \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Antoniazzi et al. \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In this study, ant samples were collected by a student with limited expertise and training, indicating the method has wide applications for monitoring high-elevation forests. For remote sites, particularly in areas with prominent local or indigenous groups presence, such as Lasha Mountain, a test will be if long-term biological monitoring can be facilitated by interested locals. Lasha Mountain has communities representing multiple ethnic nationalities that depend on the local biodiversity and the sustainable management of grazing land, soil, and water for their livelihoods. Establishing robust methods that can involve indigenous or local people can benefit research as well as improve situations for local people.\u003c/p\u003e \u003cp\u003eThis study demonstrates the sample plot method, a standardised hand-sampling methodology, is more efficient and productive than pitfall trapping for ants in this high-elevation forest system. Where resources limitations restrict the scope of sampling methods, the sample plot method is the most appropriate choice, providing a higher sampling efficiency on a sample unit basis. However, our results indicate pitfall trapping provides additional information on ant assemblages not detected in sample plot methods, indicating that where possible, adding complementary methods, such as pitfall trapping, to the sampling protocol improves sensitivity and accuracy of long-term biological monitoring.\u003c/p\u003e"},{"header":"Declarations","content":"\u003ch2\u003eAcknowledgments\u003c/h2\u003e \u003cp\u003eThe authors would like to express their gratitude to the Institute of Eastern-Himalaya Biodiversity Research and the Administration of Yunling Provincial Nature Reserve. We gratefully acknowledge the invaluable help of Professor. Zheng-Hui Xu, Southwest Forestry College for the species identification.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\u003cli\u003e\u003cspan\u003eAbril S, G\u0026oacute;mez C (2013) Rapid assessment of ant assemblages in public pine forests of the central Iberian Peninsula. For Ecol Manag 293:79\u0026ndash;84\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAdis J (1979) Problems of interpreting arthropod sampling with pitfall traps. Zoologischer Anzeiger - A Journal of Comparative Zoology 202:177\u0026ndash;184\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAgosti D, Alonso LE (2000) The ALL protocol: A standard protocol for the collection of ground-dwelling ants. \u003cem\u003eAnts: Standard Methods for Measuring and Monitoring Biodiversity\u003c/em\u003e (ed. D.E. Wilson), pp.\u0026nbsp;204\u0026ndash;206\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAgosti D, Majer JD, Alonso LE, Schultz T (2000) Ants: standard methods for measuring and monitoring biodiversity. In: Wilson DE (ed) Biological Diversity Handbook Series. Smithsonian Institution, Washington DC, p 280\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAndersen AN (1997) Using ants as bioindicators: multiscale issues in ant community ecology.Conservation ecology, \u003cb\u003e1\u003c/b\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAndersen AN, Fisher A, Hoffmann BD, Read JL, Richards R (2004) Use of terrestrial invertebrates for biodiversity monitoring in Australian rangelands, with particular reference to ants. Austral Ecol 29:87\u0026ndash;92\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAndersen AN, Hoffmann BD, M\u0026uuml;ller WJ, Griffiths AD (2002) Using ants as bioindicators in land management: simplifying assessment of ant community responses. J Appl Ecol 39:8\u0026ndash;17\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAndersen AN, Majer JD (2004) Ants show the way Down Under: invertebrates as bioindicators in land management. Front Ecol Environ 2:291\u0026ndash;298\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eAntoniazzi R, Ahuatzin DA, Pelayo-Mart\u0026iacute;nez J, Ortiz-Lozada L, Leponce M, D\u0026aacute;ttilo W (2020) On the effectiveness of hand collection to complement baits when studying ant vertical stratification in tropical rainforests. Sociobiology 67:213\u0026ndash;222\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBates D, M\u0026auml;chler M, Bolker B, Walker S (2015) Fitting linear mixed-effects models using lme4. J Stat Softw 67:1\u0026ndash;48\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBestelmeyer BT, Agosti D, Alonso LE, Brandao CRF, Brown WL, Delabie JHC, Silvestre R (2000) Field techniques for the study of ground-dwelling ants: an overview, description, and evaluation. \u003cem\u003eAnts: Standard Methods for Measuring and Monitoring Biodiversity\u003c/em\u003e, pp.\u0026nbsp;122\u0026ndash;144\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBowie MH, Frampton CM (2004) A practical technique for non-destructive monitoring of soil surface invertebrates for ecological restoration programmes. Ecol Manag Restor 5:34\u0026ndash;42\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eBray D (2014) Modification of the ALL protocol to characterise the overall ant assemblage in temperate eucalypt forest. Australian Entomol 41:15\u0026ndash;28\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eCardoso P, Erwin TL, Borges PAV, New TR (2011) The seven impediments in invertebrate conservation and how to overcome them. Biol Conserv 144:2647\u0026ndash;2655\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChao A, Chiu C-H, Jost L (2014) Unifying species diversity, phylogenetic diversity, functional diversity, and related similarity and differentiation measures through Hill numbers. Annu Rev Ecol Evol Syst 45:297\u0026ndash;324\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChao A, Gotelli NJ, Hsieh TC, Sander EL, Ma KH, Colwell RK, Ellison AM (2014) Rarefaction and extrapolation with Hill numbers: a framework for sampling and estimation in species diversity studies. Ecol Monogr 84:45\u0026ndash;67\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eChao A, Jost L (2012) Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93:2533\u0026ndash;2547\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eClarke KR (1993) Non-parametric multivariate analyses of changes in community structure. Aust J Ecol 18:117\u0026ndash;143\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eClarke KR, Somerfield PJ, Chapman MG (2006) On resemblance measures for ecological studies, including taxonomic dissimilarities and a zero-adjusted Bray\u0026ndash;Curtis coefficient for denuded assemblages. J Exp Mar Biol Ecol 330:55\u0026ndash;80\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eDing YQ (1980) Principles of Mathematical Ecology of Insect Populations. Science Press, Beijing\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ede Souza JLP, Baccaro FB, Landeiro VL, Franklin E, Magnusson WE (2012) Trade-offs between complementarity and redundancy in the use of different sampling techniques for ground-dwelling ant assemblages. Appl Soil Ecol 56:63\u0026ndash;73\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFang H, Xu ZH (2021) Formicidae insects research of China based on bibliometrics. J Southwest Forestry Univ 41:14\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eFotso Kuate A, Hanna R, Tindo M, Nanga S, Nagel P (2015) Ant Diversity in Dominant Vegetation Types of Southern Cameroon. Biotropica 47:94\u0026ndash;100\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGerlach J, Samways M, Pryke J (2013) Terrestrial invertebrates as bioindicators: an overview of available taxonomic groups. J Insect Conserv 17:831\u0026ndash;850\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGotelli NJ, Colwell RK (2001) Quantifying biodiversity: procedures and pitfalls in the measurement and comparison of species richness. Ecol Lett 4:379\u0026ndash;391\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGotelli NJ, Ellison AM, Dunn RR, Sanders NJ (2011) Counting ants (Hymenoptera: Formicidae): Biodiversity sampling and statistical analysis for myrmecologists. Myrmecological News 15:13\u0026ndash;19\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eGuerra CA, Bardgett RD, Caon L, Crowther TW, Delgado-Baquerizo M, Montanarella L, Navarro LM, Orgiazzi A, Singh BK, Tedersoo L, Vargas-Rojas R, Briones MJI, Buscot F, Cameron EK, Cesarz S, Chatzinotas A, Cowan DA, Djukic I, Van Den Hoogen J, Lehmann A, Maestre FT, Mar\u0026iacute;n C, Reitz T, Rillig MC, Smith LC, De Vries FT, Weigelt A, Wall DH, Eisenhauer N (2021) Tracking, targeting, and conserving soil biodiversity: A monitoring and indicator system can inform policy. Science 371:239\u0026ndash;241\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHarris JWE, Collis DG, Magar KM (1972) Evaluation of the tree-beating method for sampling defoliating forest insects. Can Entomol 104:723\u0026ndash;729\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHerv\u0026eacute; M (2018) RVAideMemoire: testing and plotting procedures for biostatistics. \u003cem\u003eR package version 0.9\u0026ndash;69\u003c/em\u003e, \u003cb\u003e3\u003c/b\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHobbie JE, Carpenter SR, Grimm NB, Gosz JR, Seastedt TR (2003) The US Long Term Ecological Research program. Bioscience 53:21\u0026ndash;32\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHoffmann BD, Pettit M (2022) Towards precision ecology: Relationships of multiple sampling methods quantifying abundance for comparisons among studies.Plos One, \u003cb\u003e17\u003c/b\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eH\u0026ouml;lldobler B, Wilson EO (1990) The ants. Harvard University Press\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHsieh TC, Ma KH, Chao A (2016) iNEXT: an R package for rarefaction and extrapolation of species diversity (Hill numbers). Methods Ecol Evol 7:1451\u0026ndash;1456\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eHuang ZP, Scott MB, Li YP, Ren GP, Xiang ZU, Cui LW, Xiao W (2017) Black-and-white snub-nosed monkey (\u003cem\u003eRhinopithecus bieti\u003c/em\u003e) feeding behavior in a degraded forest fragment: clues to a stressed population. \u003cem\u003ePrimates\u003c/em\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eecosystem services of the Intergovernmental Science-Policy Platform on Biodiversity and Ecosystem Services (2019) In: Brondizio ES, Settele J, D\u0026iacute;az S, Ngo HT (eds) Global assessment report on biodiversity and. IPBES secretariat, Bonn, Germany, p 1148\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eJim\u0026eacute;nez-Carmona F, Carpintero S, Reyes-L\u0026oacute;pez JL (2020) Ant sampling: the importance of pitfall trap depth as a bias factor. Entomol Exp Appl 168:703\u0026ndash;709\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eKuussaari M, Bommarco R, Heikkinen RK, Helm A, Krauss J, Lindborg R, \u0026Ouml;ckinger E, P\u0026auml;rtel M, Pino J, Rod\u0026agrave; F, Stefanescu C, Teder T, Zobel M, Steffan-Dewenter I (2009) Extinction debt: a challenge for biodiversity conservation. Trends in Ecology and Evolution 24:564\u0026ndash;571\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLarsson J (2021) eulerr: Area-Proportional Euler and Venn Diagrams with Ellipses\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLength RV (2022) emmeans: Estimated Marginal Means, aka Least-Squares Means\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi AN, Xu ZH, Xu GL, Zheng Y, Zhuang JX, Li B (2017) Ant diversity of Tongbiguan Nature Reserve and adjacent area in Yunnan. J Southwest Forestry Univ 37:135\u0026ndash;141\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi N, Huang Z-P, Xiao W, Cui L-W (2019) Seed dispersal by black-and-white snub-nosed monkey (\u003cem\u003eRhinopithecus bieti\u003c/em\u003e) at Lashashan, Yunnan, China. \u003cem\u003eChinese Journal of Ecology\u003c/em\u003e, \u003cb\u003e8\u003c/b\u003e, 2019\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi Q, Chen YQ, Xu ZH (2009) Research methods on ant community. Chin J Ecol 28:1862\u0026ndash;1870\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi Q, Lu ZX, Zhang W, Ma YY, Feng P (2015a) Communities of ground-dwelling ants in different plantation forests in arid-hot valleys of Jinsha River, Yunnan Province. China Scientia Silvae Sinicae 51:137\u0026ndash;145\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLi Q, Lu ZX, Zhang W, Ma YY, Feng P (2015b) Ground-dwelling ants as bioindicators during 30-year vegetation restoration in a savanna area, Yunnan. Acta Ecol Sin 35:6199\u0026ndash;6207\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLindenmayer DB, Likens GE (2010) The science and application of ecological monitoring. Biol Conserv 143:1317\u0026ndash;1328\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLongino JT, Coddington J, Colwell RK (2002) The ant fauna of a tropical rain forest: estimating species richness three different ways. Ecology 83:689\u0026ndash;702\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLongino JT, Colwell RK (1997) Biodiversity assessment using structured inventory: Capturing the ant fauna of tropical rainforest. Ecol Appl 7:1263\u0026ndash;1277\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLopes CT, Vasconcelos HL (2008) Evaluation of three methods for sampling ground-dwelling ants in the Brazilian Cerrado. Neotrop Entomol 37:399\u0026ndash;405\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eLuff ML (1975) Some features influencing the efficiency of pitfall traps. Oecologia 19:345\u0026ndash;357\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMajer J (1997) The use of pitfall traps for sampling ants - A critique. Mem Museum Vic 56:323\u0026ndash;329\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMark K, Guenard B, Formicidae (2017) \u003cem\u003eMyrmecological News\u003c/em\u003e, \u003cb\u003e25\u003c/b\u003e, 1\u0026ndash;16\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMbenoun PS, Tadu Z, Djieto Lordon C, Mony R, Kenne M, Tindo M (2021) Efficiency of sampling methods for capturing soil-dwelling ants in three landscapes in southern Cameroon. Soil Organisms 93:115\u0026ndash;132\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMcGeoch MA (1998) The selection, testing and application of terrestrial insects as bioindicators. Biol Rev Camb Philos Soc 73:181\u0026ndash;201\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eMyers N, Mittermeler RA, Mittermeler CG, Da Fonseca GAB, Kent J (2000) Biodiversity hotspots for conservation priorities. Nature 403:853\u0026ndash;858\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNavarro LM, Fern\u0026aacute;ndez N, Guerra C, Guralnick R, Kissling WD, Londo\u0026ntilde;o MC, Muller-Karger F, Turak E, Balvanera P, Costello MJ, Delavaud A, Serafy E, Ferrier GY, Geijzendorffer S, Geller I, Jetz GN, Kim W, Kim ES, Martin H, McGeoch CS, Mwampamba MA, Nel TH, Nicholson JL, Pettorelli E, Schaepman N, Skidmore ME, Pinto ASousa, Vergara I, Vihervaara S, Xu P, Yahara H, Gill T, Pereira HM (2017) Monitoring biodiversity change through effective global coordination. Curr Opin Environ Sustain 29:158\u0026ndash;169\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eNoss RF (1990) Indicators for Monitoring Biodiversity: A Hierarchical Approach. Conserv Biol 4:355\u0026ndash;364\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOksanen J, Blanchet F, Friendly M, Kindt R, Legendre P, McGlinn D, Minchin P, O\u0026rsquo;Hara R, Simpson G, Solymos P (2019) vegan: Community Ecology Package\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eOliveira M, Lucia TMC, Marinho C, Delabie J, Morato ER (2009) Ant diversity in an area of the Amazon Forest in acre, Brazil. Sociobiology 54:243\u0026ndash;267\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eParr C, Eggleton P, Davies A, Evans TA, Holdsworth S (2016) Suppression of savanna ants alters invertebrate composition and influences key ecosystem processes. Ecology 97:1611\u0026ndash;1617\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eParr CL, Chown SL (2001) Inventory and bioindicator sampling: testing pitfall and Winkler methods with ants in a South African savanna. J Insect Conserv 5:27\u0026ndash;36\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003ePurvis A, Hector A (2000) Getting the measure of biodiversity. Nature 405:212\u0026ndash;219\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eR Core Team (2020) R: A Language and Environment for Statistical Computing. R Foundation for Statistical Computing, Vienna, Austria\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eRomero H, Jaffe K (1989) A comparison of methods for sampling ants (Hymenoptera, Formicidae) in savannas. Biotropica 21:348\u0026ndash;352\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSchultheiss P, Nooten S, Wang R, Wong M, Brassard F, Gu\u0026eacute;nard B (2022) The abundance, biomass, and distribution of ants on Earth. \u003cem\u003eProceedings of the National Academy of Sciences\u003c/em\u003e, \u003cb\u003e119\u003c/b\u003e, e2201550119\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eS\u0026oslash;rensen L, Coddington J, Scharff N (2009) Inventorying and Estimating Subcanopy Spider Diversity Using Semiquantitative Sampling Methods in an Afromontane Forest. Environ Entomol 31:319\u0026ndash;330\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eSteiner FM, Schlick-Steiner BC, Moder K, Bruckner A, Christian E (2005) Congruence of data from different trapping periods of ant pitfall catches (Hymenoptera: Formicidae). \u003cb\u003ev\u003c/b\u003e. \u003cb\u003e46\u003c/b\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTiede Y, Schlautmann J, Donoso DA, Wallis CI, Bendix J, Brandl R, Farwig N (2017) Ants as indicators of environmental change and ecosystem processes. Ecol Ind 83:527\u0026ndash;537\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTopping CJ, Sunderland KD (1992) Limitations to the use of pitfall traps in ecological studies exemplified by a study of spiders in a field of winter wheat. J Appl Ecol 29:485\u0026ndash;491\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTremblay A, Ransijn J (2015) LMERConvenienceFunctions: Model selection and post-hoc analysis for (G) LMER models. \u003cem\u003eR package version\u003c/em\u003e, \u003cb\u003e2\u003c/b\u003e\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eTista M, Fiedler K (2011) How to evaluate and reduce sampling effort for ants. J Insect Conserv 15:547\u0026ndash;559\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eUnderwood EC, Fisher BL (2006) The role of ants in conservation monitoring: if, when, and how. Biol Conserv 132:166\u0026ndash;182\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eVineesh PJ, Sabu TK, Karmaly KA (2007) Community structure and functional group classification of litter ants in the montane evergreen and deciduous forests of Wayanad region of Western Ghats, Southern India. Orient Insects 41:427\u0026ndash;442\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWang SJ, Huang ZP, He YC, He XD, Li DH, Sun J, Cui LW, Xiao W (2012) Mating behavior and birth seasonality of black-and-white snub-nosed monkeys (\u003cem\u003eRhinopithecus bieti\u003c/em\u003e) at Mt. Lasha. Zoological Res 33:241\u0026ndash;248\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eWard DF, New TR, Yen AL (2001) Effects of pitfall trap spacing on the abundance, richness and composition of invertebrate catches. J Insect Conserv 5:47\u0026ndash;53\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eXu Z (2002) A study on the biodiversity of Formicidae ants of Xishuangbanna Nature Reserve. Yunnan Science and Technology Press, Yunnan\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eYeo K, Delsinne T, Konate S, Alonso LL, A\u0026iuml;dara D, Peeters C (2017) Diversity and distribution of ant assemblages above and below ground in a West African forest\u0026ndash;savannah mosaic (Lamto, C\u0026ocirc;te d\u0026rsquo;Ivoire). Insectes Sociaux 64:155\u0026ndash;168\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZaller JG, Kerschbaumer G, Rizzoli R, Tiefenbacher A, Gruber E, Schedl H (2015) Monitoring arthropods in protected grasslands: Comparing pitfall trapping, quadrat sampling and video monitoring. Web Ecol 15:15\u0026ndash;23\u003c/span\u003e\u003c/li\u003e \u003cli\u003e\u003cspan\u003eZhang HY, Ou XH (2006) Using insect for indicator to monitor and assess forest ecosystem health. World Forestry Research 19:22\u0026ndash;25\u003c/span\u003e\u003c/li\u003e\u003c/ol\u003e"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":false,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"journal-of-insect-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jico","sideBox":"Learn more about [Journal of Insect Conservation](http://link.springer.com/journal/10841)","snPcode":"10841","submissionUrl":"https://submission.nature.com/new-submission/10841/3","title":"Journal of Insect Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Ant survey, sampling method, ground-dwelling, epigaeic, hypogaeic, Formicidae, Hengduan Mountains","lastPublishedDoi":"10.21203/rs.3.rs-2261097/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-2261097/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eSampling efficiency, composition and detection biases associated with pitfall-trap and sample plot methods were compared at seven montane sites at Lasha Mountain, Yunnan, China. On average, sample plot samples contained 1.5 times more taxa than pitfall-trap samples; however, we found no significant difference between of alpha and beta diversity in pitfall-trap and sample plot site samples. Rarefaction-interpolations curves revealed significantly higher total diversity from sample plot methods; that sample plot methods would require three times more sampling to reach asymptote of true diversity; and that sample plot samples achieve higher sample coverage across sample sizes. Permutational multivariate analysis of variance showed community composition and dominant species differed between methods. Of all taxa collected, the two methods had 16 species in common, accounting for 52% of the total species; 29% were exclusive to sample plot samples and 16% were exclusive to pitfall traps.\u003c/p\u003e \u003cp\u003e \u003cb\u003eImplications for insect conservation\u003c/b\u003e: Our findings suggest that results from the two methods cannot be directly compared and are imperfect substitutes to one another. For long-term monitoring of biodiversity, we suggest integrating multiple complementary methods, including standardised active collection methods, such as the sample plot method, to achieve more complete representation of ant composition and diversity.\u003c/p\u003e","manuscriptTitle":"A comparison of two methods for quantifying ant diversity and community in an East Himalayan montane forest mosaic","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2022-11-14 18:18:15","doi":"10.21203/rs.3.rs-2261097/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"decision","content":"Major revision","date":"2022-12-28T01:55:04+00:00","index":"","fulltext":""},{"type":"editorInvitedReview","content":"","date":"2022-12-05T16:39:45+00:00","index":"hide","fulltext":""},{"type":"reviewerAgreed","content":"ec5d4a85-cda2-460c-9349-15b8a4d474fc","date":"2022-11-25T18:47:45+00:00","index":"hide","fulltext":""},{"type":"reviewersInvited","content":"","date":"2022-11-24T01:49:24+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2022-11-11T18:45:43+00:00","index":"","fulltext":""},{"type":"checksComplete","content":"","date":"2022-11-11T18:45:42+00:00","index":"","fulltext":""},{"type":"submitted","content":"Journal of Insect Conservation","date":"2022-11-11T01:16:13+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"journal-of-insect-conservation","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"jico","sideBox":"Learn more about [Journal of Insect Conservation](http://link.springer.com/journal/10841)","snPcode":"10841","submissionUrl":"https://submission.nature.com/new-submission/10841/3","title":"Journal of Insect Conservation","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"220d8946-3e11-4e5b-b7fd-86f4c630e007","owner":[],"postedDate":"November 14th, 2022","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"published-in-journal","subjectAreas":[],"tags":[],"updatedAt":"2023-10-16T22:08:31+00:00","versionOfRecord":{"articleIdentity":"rs-2261097","link":"https://doi.org/10.1007/s10841-023-00501-y","journal":{"identity":"journal-of-insect-conservation","isVorOnly":false,"title":"Journal of Insect Conservation"},"publishedOn":"2023-08-09 21:57:07","publishedOnDateReadable":"August 9th, 2023"},"versionCreatedAt":"2022-11-14 18:18:15","video":"","vorDoi":"10.1007/s10841-023-00501-y","vorDoiUrl":"https://doi.org/10.1007/s10841-023-00501-y","workflowStages":[]},"version":"v1","identity":"rs-2261097","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-2261097","identity":"rs-2261097","version":["v1"]},"buildId":"-HB7Z8yhvgn0wM9Nzuekk","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}