Abstract
When studying arthropod visitors of flowers, the sampling unit (individual flowers, groups of flowers, areas of plants, timed visits, etc.) depends upon the aim of the study and the sampling method employed. In this study, arthropods using flowers of the ice plant, Carpobrotus edulis, were recorded on the sand dunes at New Brighton in the south island of New Zealand. Of 3600 flowers, only 10% contained invertebrates and only 478 specimens were recorded in total. Of 32 arthropod species observed on this exotic plant, we consider at least 20 to be native to New Zealand and five species are probably New Zealand endemics. Based on an occupation rate of individual flowers of 10%, a binomial model indicated that a sample of 100 flowers would have <0.003% chance of containing no specimens, and 96% chance that 5–16 flowers would contain some animals. Species accumulation models (e.g. bootstrap, Chao and rarefaction) and models examining the likelihood of recording rare species indicated that after examining 2000 flowers, 80% of arthropod species would be recorded, and that only the rarest species in our study would fall below an 80% statistical power of detection. The results suggest that for this flower–invertebrate system, a scheme that involved 20 independent samples, each consisting of 100 flowers, would provide a good chance of (1) avoiding totally empty samples (2) collecting a high proportion of the total species present and (3) recording all but the very rarest species that occur in this system.
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References
Albrecht M, Schmid B, Hautier Y, Muller CB (2012) Diverse pollinator communities enhance plant reproductive success. Proc R Soc B 279:4845–4852
Anderson MJ, Santana-Garcon J (2014) Measures of precision for dissimilarity-based multivariate analysis of ecological communities. Ecol Lett 8:66–73
Au L (2000) Carpobrotus edulis in coastal california plant communities. Restor Reclam Rev 6:1–7
Bartomeus I, Vilà M, Santamaría L (2008) Contrasting effects of invasive plants in plant–pollinator networks. Oecologia 155:761–770
Bezemer TM, Harvey JA, Cronin JT (2014) The response of native insect communities to invasive plants. Ann Rev Entomol 59:119–141
Cao Y, Dudley-Williams D, Larsen DP (2002) Comparison of ecological communities: the problem of sample representativeness. Ecol Monogr 72:41–56
Cardoso P (2009) Standardization and optimization of arthropod inventories—the case of Iberian spiders. Biodivers Conserv 18:3949
Chacoff NP, Vazquez DP, Lomascolo SB, Stevani EL, Dorado J, Padro B (2012) Evaluating sampling completeness in a desert plant–pollinator network. J Anim Ecol 81:190–200
Chao A (1984) Non-parametric estimation of the number of classes in a population. Scand J Stat 11:265–270
Chao A, Jost L (2012) Coverage-based rarefaction and extrapolation: standardizing samples by completeness rather than size. Ecology 93:2533–2547
Chao A, Colwell RK, Lin C-W, Gotelli NJ (2009) Sufficient sampling for asymptotic minimum species richness estimators. Ecology 90:1125–1133
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
Chinnock RJ (1972) Natural hybrids between Disphyma and Carpobrotus (Aizoaceae) in New Zealand. N Z J Bot 10:615–625
Colwell RK, Coddington JA (1994) Estimating terrestrial biodiversity through extrapolation. Phil Trans R Soc B 345:101–118
Colwell RK, Chang XM, Chang J (2004) Interpolating, extrapolating, and comparing incidence-based species accumulation curves. Ecology 98:2717–2727
Cunningham RB, Lindenmayer DB (2005) Modelling count data of rare species: some statistical issues. Ecology 86:1135–1142
Delipetrou P (2006) Carpobrotus edulis. DAISIE. www.europe-aliens.org/pdf/Carpobrotus_edulis.pdf. Accessed 2 Feb 2017
Essenberg CJ (2012) Explaining variation in the effect of floral density on pollinator visitation. Am Nat 180:153–166
Falcao JCF, Dattilo W, Rico-Gray V (2016) Sampling effort differences can lead to biased conclusions on the architecture of ant–plant interaction networks. Ecol Complex 25(2016):44–52
Forster RR (1979) The spiders of New Zealand. Part V. Cycloctenidae, Gnaphosidae, Clubionidae. Otago Mus Bul 5:8–95
Garbuzov M, Ratnieks FLW (2014) Quantifying variation among garden plants in attractiveness to bees and other flower-visiting insects. Funct Ecol 28:364–374
Gerber E, Krebs C, Murrell C, Morietti M, Rocklin R, Schaffner U (2008) Exotic invasive knotweeds (Fallopia spp.) negatively affect native plant and invertebrate assemblages in European riparian habitats. Biol Conserv 141:646–654
Green RH, Young RC (1993) Sampling to detect rare species. Ecol Appl 3:351–356
Harder LD, Jordan CY, Gross WE, Routley MB (2004) Beyond floricentrism: the pollination function of inflorescences. Plant Species Biol 19:137–148
Henderson P, Seaby R (2008) A practical handbook for multivariate methods. Pisces Conservation Ltd., Lymington
Hodge S, Vink CJ (2016) Evidence of absence is not proof of absence: the case of the New Brighton katipō. N Z J Zool. doi:10.1080/03014223.2016.1227343
Hoehn P, Tscharntke T, Tylianakis JM, Steffan-Dewenter I (2008) Functional group diversity of bee pollinators increases crop yield. Proc R Soc B 275:2283–2291
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
Kenkel NC, Juhasz-Nagy IP, Podani J (1989) On sampling procedures in population and community ecology. Vegetatio 83:195–207
Leather SR, Basset Y, Didham RK (2014) How to avoid the top ten pitfalls in insect conservation and diversity research and minimise your chances of manuscript rejection. Insect Conserv Divers 7:1–3
MacDonald KJ, Lennon ZJ, Bensemann LL, Clemens J, Kelly D (2015) Variable pollinator dependence of three Gastrodia species (Orchidaceae) in modified Canterbury landscapes. N Z J Ecol 39:208–213
Macfarlane RP (2005) New brighton sand dune invertebrates. A report prepared for Christchurch City Council. Christchurch
MacKenzie D (2005) What are the issues with presence-absence data for wildlife managers? J Wildl Manag 69:849–860
Magurran AE, Henderson PA (2003) Explaining the excess of rare species in natural species abundance distributions. Nature 422:714–716
Mawdsley JR (2003) The importance of species of Dasytinae (Coleoptera: Melyridae) as pollinators in Western North America. Coleopt Bull 57:154–160
McBride GB, Johnstone P (2011) Calculating the probability of absence using the credible interval value. N Z J Ecol 35:189–190
Millidge AF (1988) The spiders of New Zealand. Part VI. Linyphiidae. Otago Mus Bull 6:35–67
Moragues E, Traveset A (2005) Effect of Carpobrotus spp. on the pollination success of native plant species of the Balearic Islands. Biol Conserv 122:611–619
Neinhuis CM, Dietzsch AC, Stout CJ (2009) The impacts of an invasive alien plant and its removal on native bees. Apidologie 40:450–463
Novotny V, Basset Y (2000) Rare species in communities of tropical insect herbivores: pondering the mystery of singletons. Oikos 89:564–572
Prasad AV, Hodge S (2013a) The diversity of arthropods associated with the exotic creeping daisy Sphagneticola trilobata in Suva, Fiji Islands. Entomol Mon Mag 149:155–161
Prasad AV, Hodge S (2013b) Factors influencing the foraging activity of the allodapine bee Braunsapis puangensis on creeping daisy (Sphagneticola trilobata) in Fiji. J Hymenopt Res 35:59–69
Primack RB (1978) Variability in New Zealand montane and alpine pollinator assemblages. N Z J Ecol 1:66–73
Queheillalt DM, Cain JW III, Taylor DE, Morrison ML, Hoover SL, Tuatoo-Bartley N, Rugge L, Christopherson K, Hulst MD, Harris MR, Keough HL (2002) The exclusion of rare species from community-level analyses. Wildl Soc Bull 30:756–759
Regan TJ, McCarthy MA, Baxter PWJ, Panetta FD, Possingham HP (2006) Optimal eradication: when to stop looking for an invasive plant. Ecol Lett 9:759–766
Rivera-Hutinel A, Bustamante RO, Marin VH, Medel R (2012) Effects of sampling completeness on the structure of plant–pollinator networks. Ecology 93:1593–1603
Royale JA, Chandler RB, Yackulic C, Nichols JD (2012) Likelihood analysis of species occurrence probability from presence-only data for modelling species distributions. Methods Ecol Evol 3:545–554
Samways MJ, McGeoch MA, New TR (2010) Insect conservation. OUP, New York
Showler K (1989) The Himalayan balsam in Britain—an undervalued source of nectar. Bee World 70:130–131
Stary P, Tkalcu B (1998) Bumble-bees (Hym. Bombidae) associated with the expansive touch-me-not, Impatiens glandulifera in wetland biocorridors. Anz Schadlingskunde Pflanzenschutz Umweltschutz 71:85–87
Stouffer DB, Cirtwill AR, Bascompte J (2014) How exotic plants integrate into pollination networks. J Ecol 102:1442–1450
Suehs CM, Medail F, Affre L (2004) Invasion dynamics of two alien Carpobrotus (Aizoaceae) taxa on a Mediterranean island: I. genetic diversity and introgression. Heredity 92:31–40
Sunny A, Diwaka S, Sharma GP (2015) Native insects and invasive plants encounters. Arthropod-Plant Interact 9:323–331
Tikoca S, Hodge S, Tuiwawa M, Pene S, Clayton J, Brodie S (2016) An appraisal of sampling method and effort for investigating moth assemblages in a Fijian forest. Aust Entomol 55:455–462
Topp W, Kappes H, Rogers F (2008) Response of ground-dwelling beetle (Coleoptera) assemblages to giant hogweed (Reynoutria spp.) invasion. Biol Invasions 10:381–390
Vila M, Bartomeus I, Dietzsch AC, Petanidou T, Steffan-Dewenter I, Stout JC, Tscheulin T (2009) Invasive plant integration into native plant–pollinator networks across Europe. Proc R Soc B 276:3887–3893
Vink CJ, Sirvid PJ (1998) The Oxyopidae (lynx spiders) of New Zealand. N Z Entomol 21:1–9
Westphal C, Bommarco R, Carré G et al (2008) Measuring bee diversity in different European habitats and biogeographical regions. Ecol Monogr 78:653–671
Acknowledgements
The authors wish to thank Jason Roberts and Antony Shadbolt of Christchurch City Council, for permission to carry out the survey at New Brighton and providing a copy of the report by Rod MacFarlane, respectively. John Early of Auckland Museum provided identifications of the parasitoid Hymenoptera.
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Hodge, S., Curtis, N., Vink, C.J. et al. Native arthropods on exotic sand dune flowers: consideration of sample size and number for investigating rare species and sparse communities. Arthropod-Plant Interactions 11, 691–701 (2017). https://doi.org/10.1007/s11829-017-9521-9
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DOI: https://doi.org/10.1007/s11829-017-9521-9