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at http://www.nhm.ac.uk/research-curation/research/ historical-marine-collections/. At a time when conserving the biosphere is a global priority, NHCs (coupled with new techniques for exploiting preserved material) are a vital resource for establishing pre-anthropogenic baselines, addressing the shifting-baseline syndrome [6], and understanding temporal trends.
References
1 Johnson, K.G. et al. (2011) Climate change and biosphere response: unlocking the collections vault. Bioscience 61, 147153 2 Boakes, E.H. et al. (2010) Distorted views of biodiversity: spatial and temporal bias in species occurrence data. PLoS Biol. 8, e1000385

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3 Robbirt, K.M. et al. (2011) Validation of biological collections as a source of phenological data for use in climate change studies: a case study with the orchid Ophrys sphegodes. J. Ecol. 99, 235241 4 Inger, R. and Bearhop, S. (2008) Applications of stable isotope analyses to avian ecology. Ibis 150, 447461 5 Magurran, A.E. et al. (2010) Long-term datasets in biodiversity research and monitoring: assessing change in ecological communities through time. Trends Ecol. Evol. 25, 574582 6 Pauly, D. (1995) Anecdotes and the shifting baseline syndrome of sheries. Trends Ecol. Evol. 10, 430

0169-5347/$ see front matter 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.tree.2010.12.009 Trends in Ecology and Evolution, April 2011, Vol. 26, No. 4

Letters

The costs of describing the entire animal kingdom


Fernando Carbayo1 and Antonio C. Marques2
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Escola de Artes, Ciencias e Humanidades, Universidade de Sao Paulo, Av. Arlindo Bettio, 1000, 03828-000 Sao Paulo, Brazil Departamento Zoologia, Instituto de Biociencias, Universidade de Sao Paulo, Rua Matao, Trav. 14, 101, 05508-900, Sao Paulo, Brazil

Biodiversity is one of the major factors underpinning human wellbeing [1,2]. Although the basic approach to the conservation and appropriate use of biodiversity requires identication of its component taxa [http://www.cbd.int/gti/], this is, in general, not well advanced. With particular regard to the animal kingdom, only 1.4 million of the total 6.8 million species estimated are currently known [3]. By being made aware of these gures, government decision-makers might allocate resources more appropriately to alleviate the shortfall in taxonomic knowledge. We have attempted to estimate the resources required to carry out this task. To achieve this goal, we used Brazilian expenditure per taxonomist, which we argue represents the approximate average global expenditure per taxonomist. Our data were derived from a survey of 44 taxonomists (almost 9% of the Brazilian community of employed and doctoral taxonomists [4]). These data detailed investments in their: academic education; professional positions (total income during career); training for recently qualied taxonomists; and grants for biological sampling as well as for equipping and maintaining taxonomy laboratories. The survey also included the number of new species catalogued (totaling 1,093 species for 44 taxonomists), and the number of taxonomic publications and general publications generated. The analyses dealt with the most biodiverse animal groups, and were divided into three major sets of taxa (vertebrates, insects, and other invertebrates) to better evaluate variation. The extrapolations for the calculation of world costs were based on three main facts. First, 10% of animal species known worldwide occur in Brazil [5]. Second, the Brazilian community of taxonomists has been responsible for most taxonomic contributions to knowledge of the entire Latin American biota [6]. Also, Brazil is among the most active countries worldwide with respect
Corresponding author: Carbayo, F. (baz@usp.br).

to the description of species and number of taxonomists [http://www.mapress.com/zootaxa/support/Statistics.htm. and http://www.gti-kontaktstelle.de/english/taxonomy_E. html]. Third, the salaries of Brazilian researchers and professors fall close to the average salary for all professors worldwide [7]. Our data showed that a researcher described 24.8 species on average during his/her career (varying from 1 year to 46 years for researchers in a permanent position), with average expenditures per year of US$ 97,000. These values vary between taxonomic groups (Table 1). Therefore, we estimated that the total cost to describe unknown animal diversity would be US$ 263 billion, notably far higher than the US$ 5 billion proposed by Wilson [8] required to describe the entire biota. Historically, the total amount invested to describe one species of vertebrate is three-times the amount to describe an insect, and two-times the amount for other invertebrates [4]. This is because 31% of the total number of Brazilian animal taxonomists works on 5% of the known animal species, i.e. vertebrates [4] (on a worldwide basis, these numbers are 50% and 4%, respectively [9]). The future need for greater numbers of taxonomists working on invertebrate groups (particularly insects) is enormous. Considering the vast number of non-catalogued species and the present number of taxonomists, one invertebrate taxonomist would have to describe 292-times more species than a vertebrate taxonomist. Based on the present average expenditure required to describe one species, we estimate that the total Brazilian and world expenditure previously spent on cataloguing animal diversity totaled US$ 7 and US$ 68 billion, respectively (Table 1). Of this, an average of 15% was allocated to training taxonomists. Salaries to maintain full-time taxonomists in scientic institutions amounted to 50% of the total cost. However, two-thirds of their

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Table 1. Cost estimates for cataloguing animal diversity
World animal Brazilian animal Cost per species species species catalogued [3] catalogued [5] catalogued in Brazil No. species No. species US$ 1,000/species 61,995 4,000 122 Vertebrates 1,000,000 62,000 39 Insects 11,000 61 Other invertebrates 362,158 1,424,153 77,000 Total

Trends in Ecology and Evolution April 2011, Vol. 26, No. 4

Costs for cataloguing unknown Brazilian animal species US$ Billion 2.3 16.1 8.7 27.1

Total cost for cataloguing known world animal species US$ Billion 7.6 39.0 22.1 68.7

Estimated unknown world animal species [3] No. species 18,505 4,000,000 1,407,570 5,426,075

World total cost for cataloguing unknown world animal species US$ Billion 22.6 156.0 84.5 263.1

publications were not related to taxonomy, and these professionals had other duties such as teaching and supervising students, participating in community outreach projects, and bureaucratic commitments. The remaining 35% of the costs was for project expenses. Although this budget is huge, the main immediate obstacle to cataloguing animal diversity is undoubtedly the small and inadequate number of procient taxonomists (the taxonomic impediment). At the current rate (average of 16,000 species per year, see [3]), the present generation of trained taxonomists would take 360 years to fully catalogue world animal diversity. Increasing the number of working taxonomists would take a signicant amount of time because it takes a long time to train and develop taxonomists. A complete inventory of the animal diversity of the world might remain an elusive goal [10]. Even this considerable achievement would provide only the leftovers of biological diversity after the effects of evolution and human intervention on natural habitats have been considered. However, some crucial future actions should be considered for cataloguing biodiversity other than those related purely to funding for conservation and sustainable use of living organisms. The most essential action now would be a concerted effort to raise the image of taxonomy from being seen merely as an old and simple task of biologists that is unfashionable and horribly constricted to low-impact-factor journals to being viewed instead as a fundamental, indispensable, and vibrant branch of the life sciences.
Letters

Acknowledgements
We thank the respondents to the survey. We also thank Richard Wightwick and Rodrigo H. Willemart for offering a critique of a previous version of the manuscript, Jim Hesson for a review of the English, and an anonymous reviewer for valuable suggestions. FC was funded by FAPESP and ACM by FAPESP and CNPq.

References
1 United Nations Statistical Division (2005) Millennium Development Goal Indicators Database, Geneva 2 Chapin, F.S., III et al. (2009) Ecosystem stewardship: sustainability strategies for a rapidly changing planet. Trends Ecol. Evol. 25, 241249 3 Chapman, A.D. (2009) Numbers of Living Species in Australia and the World. Report for the Australian Biological Resources Study, Australian Biological Resources Study, Canberra 4 Marques, A.C. and Lamas, C.J.E. (2006) Taxonomia zoologica no Brasil: estado da arte, expectativas e sugestoes de acoes futuras. Pap. Av. Zool. 46, 139174 [In Portuguese] 5 Lewinsohn, T.M. and Prado P.I. (2004) Biodiversidade Brasileira. Sntese do Estado Atual do Conhecimento, Contexto [In Portuguese]. 6 Michan, L. and Llorente-Bousquets, J. (2010) Bibliometra de la sistematica biologica sobre America Latina durante el siglo XX en tres bases de datos mundiales. Int. J. Trop. Biol. 58, 531545 [In Spanish] 7 Russo, G. (2010) For love and money. Nature 465, 11041107 8 Wilson, E.O. (2000) A global map of biodiversity. Science 253, 756762 9 May, R.M. (2010) Tropical arthropod species, more or less? Science 329, 4142 10 Singh, J.S. (2002) The biodiversity crisis: a multifaceted review. Curr. Sci. India 82, 638647

0169-5347/$ see front matter 2011 Elsevier Ltd. All rights reserved. doi:10.1016/j.tree.2011.01.004 Trends in Ecology and Evolution, April 2011, Vol. 26, No. 4

Why research on traits of invasive plants tells us very little


Ken Thompson1 and Mark A. Davis2
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Department of Animal and Plant Sciences, University of Shefeld, Shefeld, UK, S10 2TN Macalester College, 1600 Grand Avenue, St. Paul, MN 55105, USA

Human impacts on the planet have never been greater; agriculture is now the major global land use, with correspondingly large increases in disturbance and changes to the global nitrogen and water cycles [1]. Effects on oras have been predictable, with robust, fast-growing, shortlived, fecund and effectively dispersed species characterisCorresponding author: Thompson, K. (ken.thompson@sheffield.ac.uk).

tic of disturbed, nutrient-rich habitats doing well, whereas slow-growing, long-lived and poorly dispersed species characteristic of nutrient-poor habitats are generally in retreat [2,3]. Paralleling the research documenting the traits of plants exploiting these disturbed and nutrient-rich environments has been an effort to determine the characteristic traits of invasive alien plants. This research has revealed
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