The Taxome Project* -- Butterflies

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Butterflies and taxonomy
There are about 17,500 described species of butterflies (including skippers) worldwide (see "the scale of the problem", Ackery et al. 1999), compared with maybe 1,413,000 species of all organisms described (Wilson 1992). Nearly half of all butterflies are estimated to be in the new world tropics (7,500 spp., see Heppner 1991, Lamas 1997, Robbins & Opler 1997, Lamas et al. submitted). Butterflies represent around 1% of all species known.

In the future, with most species of organisms described, it is to be hoped that taxonomy - i.e. information related to identification and descriptions of taxa, or nameable groups of individual organisms - will be available free of charge for government, academic, commercial and private use. This seems most in line with current taxonomic practice (where all that is required is library access, and no patenting of naturally-occurring organisms is allowed), and indeed with current practice for another fundamental kind of information, genome sequences. The most convenient system for presenting taxonomic information would be as an internationally-moderated website (Godfray 2002a,b).

A modest proposal
We propose using butterflies to provide a model "Taxome Project", a complete taxonomy, for a sizeable chunk of fauna. The work would contain  links to key genetic information of use in phylogeny reconstruction, and, indeed, almost any other information relevant to the taxon available on the word-wide web. An exemplar large-scale complete taxomony is needed as a demonstration for all of taxonomy, in a similar way that an example eukaryotic genome was provided by the genome of Drosophila melanogaster.

Why butterflies?
Butterflies are popular with the public, roughly twice as diverse as birds (9,000 spp., Wilson 1992) or ants (11,000 spp., Clarke 2002), and are important in conservation issues; for instance, in Britain, "Butterfly Conservation", the British conservation society, is second in membership only to the Royal Society for the Protection of Birds (RSPB).

Butterflies and moths are also among the earliest examples and most important organisms for the understanding of evolution, with the great discoveries in mimicry by Bates and Müller being followed rapidly by encyclopaedic analysis of sexual dimorphism and sexual selection among butterlies by Charles Darwin.  Today, a number of Lepidoptera, particularly butterflies, are model systems for evolution and ecology. Genetically also, they are important, and were the first organisms in which sex chromosomes were discovered . Currently, there is a Lepidoptera Genome project focusing particularly on the silkmoth, but including other Lepidoptera (Mita et al. 2001), and a commercial collaboration to sequence the Heliothis genome (Genoptera 2002). More recently, the call for proposals for funding Bacterial Artificial Chromosome (BAC) libraries (NSF 2001) has led to funding for a butterfly moth genomes, Heliconius, Heliothis and Manduca, which happen to have particular developmental genetic interests (info gleaned at Leiden butterfly conference; source reasonably reliable; final results to be announced).

Butterflies, perhaps because they are mobile exotherms, respond extremely rapidly to environmental disturbance or degradation, especially those involving temperature changes, and are well studied ecologically. In Europe, many butterflies have undergone well-documented extinctions or range contractions, almost all of which due to changes in agricultural practice and degradation of the environment (Thomas & Lewington 1991). Similarly, via an army of amateur recording enthusiasts, we know more about the impact of global warming on the poleward and elevational expansions and contractions of butterfly distributions than for any other organisms on the planet (Parmesan 1996, Parmesan et al. 1999, Thomas et al. 2001).

Because Europe holds most of the historical collections made especially during the days of empires and before their decline, in particular in museums in London, Paris, and various cities in Germany, a European effort is essential if historical information, particularly information about historic type specimens and other historical data, is to become generally accessible. This is not to belittle important partners, which will include the USA, Russia, China and Japan, and indeed any country or people with expertise or information on the topic of taxonomy. However, Europe, with its long-established museums has perhaps most to offer to the world community by organization of their museum resources, and a program generated from within Europe would receive world-wide support.

The Taxome Project: democratization and global dissemination of European scientific resources
Currently, the developing world criticises countries with major museums for making the information hard to extract for scientists in poorer countries with little access to libraries and obscure taxonomic literature, even though many of the data and type specimens may come from their own countries. The Taxome Project would to much to democratise this information (i.e. "data repatriation").

European countries have signed international agreements on biodiversity which mandate that they should manage their own natural resources, and also help provide other countries with information which they hold to manage theirs. Central to this effort is taxonomy, the identification and enumeration of subspecies, species and higher taxa, without which any biodiversity management is impossible. The Taxome Project aims to provide this information in an accessible, open format.

Rejuvenation of taxonomy as science**
There are likely to be many excellent scientific uses of such large-scale taxonomic databases. General patterns of evolution, speciation, and biogeography will emerge from such detailed data.

As an example, take species concepts. The so-called "phylogenetic species concept", or diagnostic species concept of Cracraft and others has been criticised in that it may lead to very large numbers of species names being required compared with current counts based on "polytypic" species. However, some of its practitioners argue that the phylogenetic concept will inflate the numbers of species of birds very little, and may only approximately double the numbers of "species" considered valid from 9,000 to about 20,000 bird species (Zink 1996). A guess based on Heliconiina would put the species inflation for the phylogenetic concept between about 4-fold (the basal Heliconiina) to about 7-fold (Heliconius, Eueides, and allies) (information from Brown 1979), but we won't know until we have an accurate database on species and subspecies names across a diverse phylogenetic group.

Of course, any study or commercial use of biology involves identification and taxonomy. Gene databases, for instance, rely on the information supplier's identifications, and  errors of identification of the taxonomic source of the sequence are likely to be more problematic than errors with the sequence read-out itself. The identification of organisms is so fundamental that it is not surprising it is one of the earliest sciences (taxonomists, including lepidopterists John Ray and James Petiver, along with Isaac Newton, were among the first generation of Fellows of the Royal Society of London founded in 1662 - see Salmon 2000).  Taxonomy was the first bioinformatics. However, historical precedent is no excuse for using an eighteenth-century means of information transfer in the twenty-first century. We need efficient means of taxomomic information retrieval now. The Taxome Project is long overdue.

*Provisional  name.  See Lederberg & McCray (2001).
**See also the discussion in the entomological newsletter of the Royal Entomological Society (Godfray 2002a; Ashburner 2002).  Since these words were written, Godfray's proposal for web-based taxonomy has come out in Nature (Godfray 2002b).  Taxonomy is becoming very topical.


Ackery,PR et al. (1999): [Papilionoidea]. In: Handbuch der Zoologie part 35. Lepidoptera. (Ed: Kristensen,NP) Springer, Berlin, 263-300.

Ashburner,M (2002): Comment on Godfray's article (of January 2002). Antenna 26, 73-74. Extract.

Brown,KS (1979): Ecologia Geográfica e Evolução nas Florestas Neotropicais. 2 vols. (Livre de Docencia) Universidade Estadual de Campinas, Campinas, Brazil.

Genoptera (2002): Bayer and Exelixis joint venture, Genoptera, is first to sequence _Heliothis_ moth genome. Genoptera Press Release 3 April 2002.

Godfray,HCJ (2002a): How might more systematics be funded? Antenna 26, 11-17. Reprint (by permission of the author).

Godfray,HCJ (2002b): Challenges for taxonomy. Nature 417(2 May), 17-19.

Heppner,JB (1991): Faunal regions and the diversity of Lepidoptera. Trop. Lepid. 2(Suppl.), 1-84.

Lamas,G (1997): Comparing the butterfly faunas of Pakitza and Tambopata, Madre de Dios, Peru, or why is Peru such a mega-diverse country? In: Tropical Biodiversity and Systematics. (Ed: Ulrich,H) (Proceedings of the International Symposium on Biodiversity and Systematics in Tropical Ecosystems, Bonn, 1994.) Zoologisches Forschunginstitut und Museum Alexander Koenig, Bonn, 165-168.

Lamas, Mielke, Robbins, & Callaghan.  Neotropical Checklist (submitted)

Lederberg,J; McCray,AT (2001): 'Ome sweet 'omics -- a genealogical treasury of words. The Scientist 15(7), 8.

Mita,K; Feyereisen,R; et al. (2001): International Lepidoptera Genome Project. Working draft for feedback and comment. 18 August 2001.

NSF (2001): Genomic Resources: Bacterial Artificial Chromosome Library Construction (BAC). Program Solicitation NSF 01-145 Directorate for Biological Sciences: Division of Integrative Biology and Neurosciences.

Parmesan,C (1996): Climate and species' range. Nature 382, 765-766.

Parmesan,C; Ryrholm,N; Stefanescu,C; Hill,JK; Thomas,CD; Descimon,H; Huntley,B; Kaila,L; Kullberg,J; Tammaru,T; Tennant,WJ; Thomas,JA; Warren,M (1999): Poleward shifts in geographical ranges of butterfly species associated with regional warming. Nature 399, 579-583.

Robbins & Opler 1997 Butterfly diversity and a preliminary comparison with bird and mammal diversity. IN: Biodiversity II, Understanding and Protecting Our Biological Resources. D. E. Wilson, M. L. Reaka-Kudla, & E. O. Wilson (eds.)., pp. 69-82, Joseph Henry Press, Washington, DC.

Salmon,MA (2000): The Aurelian Legacy. British Butterflies and their Collectors. Harley Books, Great Horkesley, Essex.

Thomas,CD; Bodsworth,EJ; Wilson,RJ; Simmons,AD; Davies,ZG; Musche,M; Conradt,L (2001): Ecological and evolutionary processes at expanding range margins. Nature 411, 577-581.

Thomas,JA; Lewington,R (1991): The Butterflies of Britain and Ireland. Dorling Kindersley, London. 224 pages.

Wilson,EO (1992): The Diversity of Life. Allen Lane, the Penguin Press, London. 424 pages.

Zink,RM (1996): Bird species diversity. Nature 381, 566.

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