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AND THE SYSTEMATICS Vane-Wright /?. /. CONSERVATION OF BIOLOGICAL DIVERSITY 1 AUSTRACT This paper concerns the role of systematies in efforts to conserve biological diversity. BioHivtTsity is seen both as and an interdisciplinary science (involving ecology and population bi(»logy as well as systematies). as a socio-political activity (because of the strongly anihropocentric focus of the Convention on Biological Diversit}). S)stetnalics has a runnbcr of key roles play, especially with respect to maximizing our limited and fragmentary knowledge of l)iology to through the predictive power of natural classification, and in helping lo set priorities for conservation when, as is inevitably the case, resources are limiled. After examining ways in which systematisls must support the growing needs of society know more about the Earth's biota, concluded that, because of their unique insights into the subjtvl, to is it many systematists have an e(jually strong responsibility to lake an active lead in of the issues relating to the stud), use, and conservation of biological diversity. Dan Janz(Mi (1993) has asked "What does trop- itarianisni" (Ponitt. 1994) is the best strategy is & open Allen Edwards, 1003: leal society want from the taxononiist?" In relation to (lel>ate (e.g., to the urgent need to know more about biological Oates, 1995), but tbis is where most of the force — question raises further issues no- l)ehind national and international eonser\ation ef- diversity, this tably, is biodiversity only really impotlant in tbe forts is now^ located. The Global Environnu^nt Fa- tropies, and does taxonomy only have a supporting eility (Glowka et al., 1994), the interim financial role in the study of biodiversity? Here will discuss nundianism of the Convention (currently adminis- I three questions, taking the view that taxono- tered by the \ôrld Bank), is the largest single all made work on and need be souree of funding ever a\ailable for mists, syst(^matists in giMieral, to — proactive as w(dl as supportive in their work. Iiiodiversity a funding souree which systematists May As seiun surprisingly slow to exploit. Robert SnoLLD Take a Lkad the (1990) has observed, "Without taxonomy lo givt^ Systi::matists in how Stidy oe Biodiversity? shape to the bricks, and systematies to tell us put them together, the house of biological science lo The subtitle of Global Biodiversity Strategy (R<Md meaningless jumble. svstematies provides a If is key documents produced et 1992). oiw. of the al., the foundation of our understanding and ability lo UNCED before the 1902 conference, '^Guid(dines is about biological diversity, what role ^.,,,,,„,unieale and Earths for action to save, study, itse biotie systematists seek to play in thi^ study, us(n ^,^^^^^1^, and The Car- wealth sustainably equitably.'' earlier and consenatiou of the Earth's biotie wealth? (lUCN/UNEP/WWF, Earth 1991) has 'ng for the / "A The sustainable the subtitle strategy for living.'" WHAT BIODIVEKSITY? IS primary obj<ictives of the Convention on Biological much An Diversity are conservation of biological div<M- impression can readily be fornuMl from ''the the sustainable use of components of the modern bioscience literature that the study sity its . . . . . . and the fair and equitable sharing of the benefits of biological divtMsily is the preseiTC, no! of sys- The from use genetic n^sourees" (Glowka tematics, but of ecology. study of diversity n\^ts arising the of 1994). Thus many of these goals are un- neither with on(^ nor the other, but with both. Bio- et al., and ashamedly anihropocentric and j)rlmarily con- diversity exists at the inteiface of pattern pro- cerned with conservation for human l>en(^fit, rather cess, as for (^xample in the twin hi<Marehy envis- than preservation of wildlife for its own sake. aged by Eldredge and Salthe (1984; Table 1), in Whether "rampant, unapologetic phylogenetics and population genetics (the dislinc- or not this util- My sincere thanks are due to Peter leaven. Jay Savage, and Mick Rieliardson for the invilation to attend tbe 41st ' many new and Annual Systemalies Symposium. was a delight to in sueli stimulating eonipatiy. meet colleagues, It l)e am Dave Garden. have an opportunity lo see firsthanil the great achievements of the Missouri liolanieal 1 grateful to comments Mollis, Scott Miller, Nigel Stork, an<l Paul Williams for eonstruetive on the manuserij)t. SW7 The Museum, Oomwell Hoad. London 5BI). L.K. Biogeograpby and Conservation Taboratory, Nalural History ^ G Ann. Missouri Hot. 83: 47-57. 1996, arl:». 48 Annals of the Garden Missouri Botanical 'ral)l(' 1. Tlu' twin biological liierarcliy (l>ase<l on El- dissimilarity between species, and basic uncertain- & Irvilgf Sallhe. V)IU), number i ties about 1u)H \o delimit the of gejies even compound in well-linown organisms, any attempt to Genealogical lii(*rarchy Ecological hierarchy estimate protean occurs tlu^ divtM'sity that at the fpalleni] [process] genetic Undeteired, Wilson has level. (1992) esti- Codons Enzymes mated something in the ord(*r of IC' rmcleotide Cciu \isJ Cells among pairs as specifying the diversity s{)ecies, to- O Organisms rganisnis gether witli more than 10'" gene combinations po- [individuation] [[)hysi(>logical function] tentially available per typical species. Taxonomists Dcincs Po[)nlalions might seem like chemists with a knowledge of 10 Species Local ecosystems elements from th<^ Periodic Table, wliile g(m(4icists Monophylclic laxa Riolic regions * might compared All Entire biosphere l)e to physicists trying to under- life stand the Universe from the beliavior of just a sin- fundanu^ntal our ignorance gle* particle. Is of the VI.- 11 ^ I.I biosphere really so extreme? .• .• tion 1ixM, ween [)[iylogenetic and tokog(*n(*tic relation- ^ ^ In 1973 a mealybug was discovered attacking Hennig, and ships: 1966), in historical versus eco- & cassava in Africa. By the end of 1989 this pest was biogeography (Myers l(»gical Ciller, 1988). o- /imnn a causing massive crop losses through(»ut the African J- Ur\l\i- vi• er Kieppei1 (1988) ilias discussedi i• n diepth a IIP . . . , iT troj)ics. Initial attempts to find a natural biocontrol number r.i c .......1 .. ot tin* sci• eiUuic and pluiosopnI i•cal1 i• ssues of r u agent faiiledi, untili • was realized that mealybuir ,1 ,, He ^ it th<^ the j)alterTi versus process d1ebi at.e. ob1 servedI .1 . J t^ tliat was . . .1^ „, ir part of an undescribed species complex, and I ^; f the analysis pat,t. ern andI process represent*s of dit- „ , . , . that only occuired naturally in the southern Neo- it and "ways fereiit incompatil)le of seeing," as 'Hlu & An (Cox Williams, r. , tropics 1981). appronriat(* c\\- lirst. emphIasi•ses d1i- sconl.-nunty, wh1 il1 e ^t1he ot.1her . ti i is wasp " cyilidi wâs then found, tested, and introduced 1based1 on ,t1h,e 1pri• nci• pl1 e ofr cont.i•nuit•.y, lîlnie .two are / r ^ hito Africa. T,h, e wasp ,has now spread over more „. 1 .. 1 rr conn)lcnu*ntary liccause pattern analysi•s offers itihe r ^han 12 million iknr.> andi gives effecti• ve control of o^,nl1,y, g,,u„;ulI e t. o common ancest.ry, and1 so gi• ves direc- i . • 11 ^he mealybu':^ throu";hout African ranee, with an 4: ' r its "»"' ^^^ tion t, o t,1lie anal1ysi•s of process, while process anal1- ^"'ô'^^ 1 , . annual cost benefit estimated $200,000,000 at way ysis is (he only of giving pattern a causal c\- & n (Herren Neuenscliwander, 1991). n r plI anat,•um. He rinally concl1 ud1edi, given we I1 iave ^ , Thi.s exampl,e, .invol, vi. ng a new pest and an ways we lilfiMcnt of seeing biology, that should ac- ( 11 unknown equally biocontrol us sonu^- aiient. tells cept, t,1fial, eacli lias i•.ts meri•t.s and1 ffaul1t.s, and1 ,t1hat. fe ^ 1 • . n tihing special about the nature of our ignorance. Bi- J re.a.^c..nU ;is. i;,n, c..o..m..p.lete Iby i-ftselIff, titl• ing compl1ementa. iy, r X 1 h, as to dieal wi•th a staggeri• rm number of spe- «w,e^ make <^*h>f1^y 1 shou..l1 dI ,,, use ofr Ibotiih approach1 es »to get. as — f^n & 1 1 cific^s there are thousands of mealybugs, and tens ^r .c.o..m.p.wh.b^»t».e. o ..;,>»..,.. /r>- a picture ol Ibi: of1 ogy as possi-bi l1 e (Kieppel1, fo ' -^ i7n i7n n ^^ thousands of encyrtid wasps. To solve the prob- !()«« ivoo, e..sopr^. 1/U— 1/lj. IJecause r .1 in.t•imat. e • . ot tins in- i enlonu.logists l.ad cut precisely which fin.l ^^'"'', I,, l.'m-lati.niship, essential systenuuists is tl.al w..rk it '"^^^ly'>"S ^^^^7 ^7^ ^^^^''^g ^'''•>' 1"^^*^' '^'^- together with ecologists and population biologists ^P^" to cies in natural habitat, and discover precisely its develop a fuller uiulerstanding of biological d(iiiver- '*"•"•' ^^"P" ^^'" ""' ""'"'^'''^ including uses—aturmost'ur- f ^'" '''"' ^^- sity,.1,11 its potential ... ^J'^'''' m^ The only reason this could be done 4quuincklv be- gently, to develop more efcfc ect^'ive st. ra.tegi• es fror -. is its , cause the existing classi.fication of f>ugs and wasps, consci-vation however incomplete, form is sufficient to a valid ^^"'' 0"«' mealybug had been /'7'^^""'''"; ^^^ SYSTEMATICS AND USE OK BIODIVERSITY ^'", TllF. accurately locat<Ml in the system, was possible to it number Cunetit estimates of tlu^ of extant spe- predict its geographical origin, and then what sort ;ies of organisms vary from 3 100 million; Ham- of insect to look out for as a natural biocontrol t(» mond (1992) gave a carefully considered estimate agent. Our detailed knowledge of the biology of all of 12.5 million, hut confidence limits on such fig- these myriad species inevitably lags behind our nn*s are incalculable. There greater agreement knowledge is of their classification, but the natural number about the of species tliat have been for- system allows us to extrapolate what knowledge we mally described (in the region of l.S-1.8 million) do have. synonymy but, ev(*ri so, the uncertainties of ensure Thus our ignorance of biology, while profound, is remain that the limits difficult to assess (Solow et manageable insofar as our classifications are pre- The complications al., 1995). of luMerozygosity, dictive. Far from being a passive pigeon-holing ac- variation in g(Mionu^ size and degree of similarity/ tivity as somi^ seem to believe, cdassification has all Volume Number Vane-Wright 49 83, 1 1996 Systematics and Conservation properties of an intelligence system, often al- have (Farris, 1079, 1983), and ihe more effective tlie lowing us go far h(^yond the seemingly limited they he for making such predit-tions. to wîll informtition available. Although there are literally millions of undeserihed species, nearly animals all ERSm SYSTEMATICS AND CONSERVING RI()I)I\ and can he placed meaningful readily [)lants wîthiii Like hypotlu^se^s, good classifications There are two major strands to conseiTation hi- famili(*s. ''al- how low limited data to be used with remarkable effect. ology: wIum'c to conser\e living things, and to by allowing interpolations through data-gaps, and consenê iIkmu (Caughley, 1994). Tlu* latter repn^- (^xtra[)olations to be made to new situations for sents a major application of ecology and population which data are not available'' (Gregory, 1980). biology. Regarding the former, syst(unatists have One of the most frequ(Mitly proposed uses for bio- been closely involved with rec(^nt d(n(dopments diversity is biochemic-al {irospecting. Costa Rica's (e.g.. see Forcy et al., 1994), iniluding the fornm- new INBio has even fornKÎ contractual agreements with lation of a|)]>roaches to measin-ing diversity international pharmac(Hitical companies. Cam(^z that take account of phylogenelic relationships. and Gauld (1993), in <l(^scribing the Costa Rican According to Taylor (1978), the notion of diver- experience, sugg(^sted that the Hymenopt(Ma, with sity, as an interrelation of species richness and in- Henry tluMr multitude of species and pharniacologically dividual abuiulance, was recognized first l)y active venoms and other secretions, are a pot<Mi- Walter Biites. Bates's idea, to evaluate tlu* diversity new and compare excellent source of valuable chemicals. of a locality with that of another. tially it While this undoubtedly true, the same can be was later formalized as or-divcrsity (\^'hittaker, is W more said for almost any div(M-se group. 1965). hittak<M- (1972), while elaborating a Schulz (1993) examined the male phcro- complex schemt et al. iS mon(*-glan(l secretions of 10 Africcin milkweed bul- account of species turnover al varjîng spatial scales separating 214 substances in 14 chemical (MaguiTan, 1988, gave a review^ of the considerable t<;Hli(^s, classes. Individual speci(\s had from 12 to 59 com- variety of })ro(êdures that have now IxM'n proposed measurement pounds (excluding tetrahydrofurans, which were not for the of ''ecological divtM'sity"), also and systematically investigated), including a high pro- suggested that lime, in addition lo richness more portion restricted individual species, or just a spatial turnover, should be included in a coin- t(» few, usually closely relateil species. Among these plete expression of diversity. Time is the primary were some compounds rarely or never found in na- dimension of (^volution along which difh^rences be- ture before, such as the ketone 16-heptadecen-2- tween lineages accumulate. need t)ne h'oin Amauris hecale, and the monot(Mp<Mie {E)- Conservation biologists, eonc:enuMl with the 2,6-dimelhyl-5-octen-l,8-diol from Danaus chrysippus. to set priorities for the preservation oi genetic di- and we wished to prospect for closely related substances versity, have nn'ognized the ; prol)lem pro ll' in oihcv species, tlu* existing cladistic classifuation posed that this could be solved, to a Iirst approxi- & (Ackeiy Vane-Wright, 19&1) would give us imme- mation, by mobilizing information contained within — "The diate and obvious clues such as die chemically un- the taxonomic hierarchy: sizc^ ol the potential invt^stigated Amauris dannfelti and A. inferna for the genetic loss is related to the taxonomic hierarchy keton(% and Daiium gilippiis for the terpe^n<\ because, ideally al least, different positions in this This suggests that, armed with an appropriate hierarchy refhn't gn^ler or lesser degrees of genetic classification, chemical prospecting need not be difference and hcruê differences in such variables we we can moqihology, behaviour, physiology, chemist find somethin"; interesting, as it ''blind'': if look at closely related species with the height(^ned and ecology. Although the degree of diflereiux' (tlu^ expectation of finding more of the same or related gap) between genera and between spi'cies within among compounds. on the other hand, we wânt to select, genera varies l)oth within and classes, the If, say, the 10 most diffenmt species out of a sample cuiTent taxonomic hierarchy provides tlie ojdy con- thumb of 1000, maximize the chance of finding radically venient rule of for determining the relatives to different chemicals per unit we can make use size of a potential loss of genetic mat<^riar' (lUCN/ effort, W UNEP/ of the diversity measunMut^nts developed in system- \^T, 1980). alic conservation evaluation (see below). Either Vane-Wright et al. (1991) ])roposed a diversity wav. the natural classification offers the most intel- metric sensitive both to individual taxonomic (hi- number ligent basis for biochemical prospecting, or for any erarchical, not formal) rank and total of other use that depends on predicting the biological species found within an area. This index, subse- propcMties of organisms. The better our classifu-a- quently called root-weight, was tlu^ fust formal tions are, the mon^ (^xplanatorj^ power they will measurement of taxic diversity. In a series of {)a- 50 Annals of the Garden Missouri Botanical liverwort liverwort moss moss horsetail horsetail fern fern cycad cycad ginkgo ginkgo conifer conifer angiosperm angiosperm Figure Ch<M)sing plants from based on chMlogram, which comhination would maximize 1. ifiree laiu! eight: llie biodiversity? Character richness (on the left) chooses liverwort, conifer or angiosperm, plus any one of the series moss- Ginkgo; character combination richness (right) selects livervsorl, fern, and conifer or angiosperm. (Assuming cliaraclcr all ehanges are associated only witli nodes shown and a comparabh' numlter of character changes occur at each node; & & based on Willian»s Humphries, IW-t; liierarchical relationships of plants from Humphries Parenti, 1986.) See text for ex[)Ianatioti. pers, notably ibosc of Williams (e.g., Williams, potential gi'iu^lic losis" (or âin). In practice, how- Hum- 1993; Williams et al., 1991, 1993, 1995; ever, is impossible to nuvisure character differ- it & pbries 199 Wêilzman Nixon ences on enough and Wîlliatns, (1992), directly a larg(^ scale, so the !), and Wluêler (1992), and Failb (e.g., 1992, 1994), distributioti chaiacttMs across taxa has to be (»f the eonc(*pt of taxic diversity has been refined and modeled. now agreed should be based is that ibis It — clarifuHl notably with n^sp<*ct to tbe fundamental on the genealogical bierarcby, as expressed more (jut^slion of what is being ineasmtHl, and wliy. an<] m<»re accural<dy by taxononuV ranks, clado- Humphn(»s al. (1995) concluded that the cur- grams, and phylogenetic trees. Debate continues, t^t rent goal is to assess oj)tion value. This concept however, n^garding wlii(li model of charact(*r means offers "a of assigning a value to risk aversion change should be applied empirical versus (e.g., tbe face of uncertainty'' (McNcely, 1988), and anagtMielic versus cladogenclic), and whetber or not iti can In* related to the task of "maximising the hu- dilfcrences should then be assesse<l in terms of man capacity adapt changing to to ecological con- character richness only, or character combination wc ditions'* (Reid, 1994). If this is accepted, then richness (Vt illiams et al., 1995). can abandon problem tbe insolubb* of trying to as- S*»mc idea of the two approaches given by Fig- is sign fixed vahu^s to individual species (Ehrenftdd, ure 1. Suppose we could only select for conscrva- and 1988) focus our attention instead at the level lion [\ucc of the eight land plant species sliown. If ot expn^ssible and luMllable characters (genes, species ri<'hness were the only criterion, then any traits, features, etc.; Faith, 1992, 1994; Wîlliams of the 56 combinations of 3 species from 8 would et al., 1995), which, collectively, can be considered to be e<jnally acc<'plal)le. But we interpret tree if lb<* r<^pres<'!il tln^ fimdamenlal cinrency units of option subt<Miding the eight species as a statement about value. their mutual phylogenetic relationships, then on the Thus a set of species can be evalualt^d in terms basis of just this information (making the assump- number of the total of different chara(*ters they rep- tion that a (ômparabb* numlxT of character changes The resent. impact of losing (or the addition of) any occur bctwtu^n each node shown), only 10 combi- partit ular species can be mtâsured in the nations will maximize character richness (livenvort way, thus fulfilling the need to assess 'Hhe size of plus conifer or angiosperm, plus any one from the Volume Number Vane-Wright 51 83, 1 1996 Systematics and Conservation TahU' 2. Compleme^ntarity: the eight plants listed in In reality both procedures have a plae(\ OfXcu it Figure have been allo<*ated to five hypothetieal areas; ây not be possible represent species, veg- I 1 to all = f= = = cyead,g liverwort,m moss, ginkgo, fern, y ptation types, or land forms from tlie outset, in ,h n conifer, a angiosperm horsetail, (Based on may which case a step-wise procedure be the most Underhill, 1994; see text for ex[>lanalion.) may appropriate. In other cases, be possible to it select from the beginning a set of areas to represent Spe- known vulnerable, biological attributes (or etc.) all n a cies: in f 1 y g I 1 in a region, in which ease a set-wise proc(Mlure will - + + + + Area I more usually ofler a efficient analysis. (In this par- + 2 -f- ticular example, is inttêsting to note that ap[>h- it + + + 3 -H eation of a taxic diversity index, such as character + + 4 -h and + + + richness, identifies both areas 2 3 as richer 5 than based (m the hierarchy given in Figure cirea 1, illustrating the point that species richness shouhl 1, regarded as the sole determinant of taxic series moss-G7n%r>). Alternatively, we could choose "<»* 1>^ niversity.j maximize character combination richness, to l»ut Both the step-wise and set-wise procedures re- then two possible combinations are selected: otilv degree the idea of c(miplementarity: the to flect angiosperm. (The liverwort, fern, plus conifer or which specified areas, singly or in combination, root-weight the cladogram rooted be- ind(^x, if is an r^pr^^^'H the species or taxic diversity of entire tweiMi liverwort and moss, would restrict the choice — roup or set of groups. Comphnnentarity, first ap- to livei'wort, moss, and horsetail but this index is and Acket7 and Vane- by Kirkpatrick (1983) some no longer regarded as appropriate.) Despite l>l'»^^tl and formalized by Vane-U'right et when ^^'igl'* (1984), significant differences in these results, large Rebelo Margules numbers of taxa are itivolved, simple species rich- al. (1991; see also et al., 1988; & 1992; Pressey 1993; Faith, Siegfried, et al., ness for an area usually turns out to be a good & 1994; Vt'illiams Humphries, 1994) lias mu(4i In and approximation both character richness for & Hum- ('onmion with and 5-diversily but, crucially, in- combination richness (Williams (3- chkra(>ler numerical stead of just reducing taxon turnover to phries, 1994). We values or indices, information on the identity of now seem close to a satisfacloiy theon-tical luMween areas retained. May la>â is what has basis for least part of (1990) calltMl at emergence Although simple, the of dece'i.tively Calculus and perhaps the of biodiversity.^Ânotlier, been the idea of complementarity has significant for even more aspect of the proeediue, significant is biodiversity evaluation. This b<'cause has shitt- is provided by the concept of complementarity. it ed attention from assessing areas on an absolute Most Imfokiant? scale richness or scoring index) to a relational BionivKKsiTY (e.g., Wi[i:itK Is scale (taking account of spatial turnover). In this AND COMtM.KMKNTAKITY IMinJCATIOiNS FTS way, areas can be seen as part of a whole. For all may and In Table 2 the eiglil land plants in Figure 1 have example, while tropical forests coral reefs been allocated to five areas. Suppose you wen* told be tlu^ ri<'hest biological systems on Earth, the veiy you could only choose one area for consena- (hstinc! biota of other ecosystems, such as those of that which would Guided by ocean bottoms or the relatively species-poor higluT select? sju^cies tion, y<»u make richness alone, area would seem an obvious latitudes, also have a unique contribution to 1 What you were then asked add a sec- (Table Comph^mentarity provides a basic crite- choice. to 3). if ond area? The greatest number of additional spe- rion for efficient and goal-directed procedures of cies to the five already represented by area 1 is two area selection, (liverwort and moss), both of which can be added by area 2 or area 4. Area 1 plus area 2 or 4 can CONTEXT f'KlorUTIKS IN thus account for seven of the eight species; the we eighth (horsetail) could then be added by a third If take into account quantitative effects of area or however, you w(^re asked to select biodiversity (Cousins, 1991), particularly in rela- (3 5). If, & exampl(\ ecosystem senices (Fhrlich just sufficient areas from the outset to represent all tion, for to eight species, obvious by inspection of Table Daily, 1993), or the value of local biodiversity to is it 2 that areas 2 and 3 together include all of them, humanity (GadglL 1991, 1992), It is evident that more solution than adding areas of the Earth should be seen as important, giving a efficient final all areas step by step, starting with the richest first. Recognl/Ing unicjue value for a particular area does 52 Annals of the Garden Missouri Botanical Heywood Tal>lt' ',\. Kssenlially cxtralropiral plant familes (from c{ al., 1978). Most contain only one or a fow spot'ies Hemisphere Hemisphere S. N. Bipolar Troehodendraeeae La<'lt>ri(laceae l-ardi/,al)alaeea< Coniortegii Cercidijiliyl]areae Empetraeeae Maleslierbiaeeae Fucommiaeeae Juncaginaeeae Achariareae Leitn<Tiaeeae Tosidoniaeeae Gnil)l)iaceae aeoniaceae I Cephalotareae Diap<'iisiaeeae Penaeaceae Theligonaeeae Misodendraceae Hi[>puridaeeae Geissolomalaeeae Cynomoriaeeae Calyceractâe Cneoraeeae Limnanthaeeae Phrymaeeae A(]oxaeeae Hutomaeeae Scheuehzeriaeeai not tnran (hat adjacent arras are unimportant, or make eedain and lists to things possible. First fore- lliat they can he abused with impunity. Priorities most, according to Janzen, a "clearuMl set of uf)" shouhl not be seen as merely choosing a few of the names and managealde a needt^d sysletti is for "fil- richest, or even most complementary but ing, comparing, searching, recording and working sitt^s, more in diff<Tential allocation of resources (Vane- with the boun- s|)ecies that constitute the . . . . . . Wright, in press) to do tlu^ best wv can, in relation tiful biodiversity resource of tropical nations." to relative impotlance both in terms of patt<Mn and Based on Janzen's ideas, and those of others like process, across all areas of land an<l sea. Never- Stork (1991) and Nitdsen and West (1994), list I iheless, for the establishment of a network of spe- Ixdow some anvis which in systematists are IxMiig cial areas to act as reser\es to ensure that as much called upon respond, to of the irreplaceabh^ qualitatively different (genea- de Tlu'rc a need de- Is to 11 logical) elemt^nts diversity survive, and are thus <»f taxonomy make . mystifv and, in the process, taxo- r •1 avaiiab1 l1 e t. o iut. ure generations, analI ys.es.ba1sed on ' nonu.c products (/ sucih as i• di enti• h> cati- on systems and ^^ •, among compl1ementa, rity areas supportnig ^ y vultiera- * ^ , catalogues) nu)re accessible (Miller, 1994). Mucl ij -1 • -n .,* / 1 l)le attribut. es (gene, s[)<M'ies, ass(Mnb1 lI ages)\ will 1be r i i , . tins attiludt* etianê re ales to Catherine svstem- f ^ r *u **' of gnât. i• mportance the l1-imi*.ted1 resources if cur- V ^ . , . , i i ; atic data in the first place, and processing in a It renlly available for biodiversity conservation are to way moun that not only offers self-satisfaction (peer /n P1i)e uus(um11 Mt, o Amaxn• num elrleect. (Pressey et. ^ i19nt9x3'^; U ^* fo yi al., n . approval), ibut ali so the rapidly expandine: satisfi<\s i^ ressey, 1994). TI-ihe whI ol1e must» Ibe managed1 as r / i J *"*b ... <'f- i needs r of n(»n-specialists. teet.•ivel1 y and1 sensi.t..ivtMI y as possn>1 l1e; wi-t.1hin suchI a whole, network a of sptM'ial reserve or managenuMil Inipwied iaxon and recording, saniplinfj:, stor- sites for biodiversity then has special significance age. Better distributional data, including bionom- and can serve a useful function. ic information such as host associations, are etc., essential (\^1ieeler, 1995; McNeely, 1995). Existing What Should Do Foh Svstf:m atists Society? information often based on ad hoc sampling is procedures, resulting in a partial and disconnected ACTION hi: coverage. Systematists should become Involved Commenting on own "What his (lueslion, does with radical approaches and to rational cost-effec- tiopieal soci<^ty want from the taxonomist?'\ Dan tlve methods for data collection and mod- si)atial Janzen obsened 'The (1993) that wording of \\w. eling, as wtdl as application of techniques for im- chapler heading the message. do not ask 'What proving Is I eslimat<'s of distributional patterns based does the taxonomist have to offer tropical society?^ on existing data (Margules & Austin, 1991, 1994). Tropical socii^ty s needs recently have been, can be This need Includes dealing with \\w almost over- and should be a nuijor n'juv(^nating force sys- whelming numl)er in of (mainly species-level) taxa tematics." According to this view society and. In thai remain unrecognized and undifferentiated particular, tropical society, is l(K»king to systema- (Wilson, 1992). Once biological samples have been 1 Volume Number Vane-Wright 53 83, 1 1996 Systematics and Conservation — made, well-docu- taxonomic remain tlu'ir coiiliiiuing availability in traditional skills skills that will mented and properly curated collections funda- fundamental further development and appliea- to is problems mental to future work, including the extraction of tion of systematics to the of biodiversity. DNA information (such as data) not necessarily Training neiv systematists. Uruh^r this Invading considered the time of original acquisition (Van<*- at we must acknowledge the need not only to ])e in- & Wright Cranston, 1992). new v<»lved in training generations of systfMuatists, but also with institution building, such as the cre- Because knowledge Better systematic analysis. new museums ation of and refereruê eoUections. of the Earth biota will inevitably remain incom s Assistance with the development of national bio- have argued above plete, that is essential to I it & (Gamez place what knowledge we do have in as powerful a diversity institutes Gauld, 1993) is likely & to re^present a particular challe^g(^ predictive system as possible Nit^lsen \êsl, (cf. 1994). This applies not only to the primar}^ activity The Involvement biodiversity projects. icith of classification, where phylogenetic methodol(,gy & emergence of "biodiversity" as a topic (Harper should be pursued vigorously building a natural in manv Hawksworth, 1994) raises issues, not hvist of system, but also secondai7 disciplines such as to -^ ^ represent a tran- ^^j^j^.,^ ^,^^ f^.^^ ^j^^j ^^^^y ^^^jy biogeography where, for example, die possibility of "ban<l-wagon," likely to run out of steam or ^j^.^^^ m endemism much recognizing areas of stdl offers who become deeply commit- too i^^.^^^,^. ^,, t|,,,ê terms of predictive power, or information (Platmck, q^^ c-ontrai^, because the concept links ^^^, ^j^^. 1991). ... concerns over the preservation nature and use of its human J dinn'tly to the needs of society, repn^sents jT make it User-ori• ent, ed d1at,abIases. II n ord1er t. o I sys- ^ 11* new way r ^ fundamentally of thiiikinii; about bioloii;- t. emat,i•c, t. axonomi•c, dist. nl-iiut»i• onal1, and l)ionomi• c .. » u i. cal di.versit. y. Systemati.sts need to play thei. r part d]at.a, t. oget.1her wi-t.xh i• mcormation about. .t1he uses andJ .. . rrr in the support of biodiversity projects, includiii"; r • • val7 ues ot diiierent. organisms, as widitMi y avai*li abi li e ... . . 11 J such diverse activities as prepariiiir user-friendly as possi•ble, user-orient* ed el1 ect* roni' c dJat»alI)as(»s ; . . .1 identification systems, trainini^ parataxonomists, must be d1evel1 oped1 andI mad1e wid1el1 y avai-l1 ab1 lI e 1by being i^ nvolved wi.th surveys and i. nventory sclicmes, means 1994; appropriate Internet: Miller, (e.g., c{. setting up museums, reference collections, and oth- Cracraft, 1995). In order to do this, has to be it making and er information systems, conservation understood continuing problems occur that will environmental evaluations, and so on. Biodivt*rsitv over costs and intellectual property rights, and r a key social issue (Machlis, 1992), and vital is is /' it ,t1hese prol1 ^lems (inc1luding cost. recovery andI aca- ^ . . that systematists play their pail, not least by being demic need be recognition) solved. to sensitive and catering for user needs (including to Improved use of advanced technology. As local names, natural products information, [)art (^tc. of the inevitable change to electronic methods for things sonu^times considered outside our n^mit). analyzing and making systematic data m<»re storing, Bixause Involvement unth education. biodiver- widely available, ever)' opportunity shoulil be takt^n important and will remain so in the future, sity is make increasingly imaginative use of coniput<'r to systematists should expect to play a full and active One and video technology. of the most obvious ar- building public awareness (Cracraft. 199S), role in development mul- eas lies in the of fully illustrated, young including, in particular, the education of tiple-enti7, interactive keys (cuiTently based, for peo[)le Yen, 1994). (e.g.. CD-ROM example, on technology: ETI, 1992; e.g., & many Watson Dallwitz, 1993), but other oppor- ACTION Pt^) tunities exist, such as the producti<»n of special checklists or other products tailored The activities listed above are described in l(M7ns to fulfill unique needs, or the application of shape-analysis of responding to the iKêds of sot:it;ty for better, Other advances biotechnology, more comprehensive, and above more accessible to identification. in all leading to automated id(^ntification procedures information about the Earths biota and its signifi- based on blood or other tissue samj>h*s, or rapid cance. Appropriate reactions by syst(unatists to tlie and sequence needs undoubtedly form our increases in the quantity quality of of society part (»f re- data, must also be expected. The community of sys- sponsibilities (especially as society at large lias al- much tematists should embrace these positive and excit- ready {)aid for so of the collectitms, libraries, ing developments because, so long as they are and other paraphernalia essential for our opera- properly set up, such information systems will give tions). However, as I will argue in the last section, — systematists more time to develop lluMr basic and is also our respimsibility to be proactive to [)ut it 54 Annals the of Garden Missouri Botanical forward ru'w ideas and cicale heller atlilu<!es to fiber, more clean water, more clean air, while ah- own hio<Hversilv hv takin<r iniliatives based on our sorhing more pollutants, from a continually di- all unique minishing and insiglils. hiotie hase, quantilativt^y qualita- Take just one example. If we attempt to set eon- tively. senation priorities based on separate analyses of Faced with this alarming prospect, most nations th(^ distributions of spûies belonging to more than have now agreed, at least in principle, to I17 to take one laxonomic gnuip, is apparent that this typi- some form <if(*orr<M'tive action, to which almost uni- it cally leads to different, sometimes totally different, versal support least on paper) for the Convention (at conclusions about what a<'tions are needed on Biological Diversity bears witness. This requires (e.g.. Can Pren<l(Mgast et al., 1993). such conflicts be re- that each nation ratifying the Convention should so1v<mI? Systematisls have proposed two approaches pursue, through various provisions outlined, the to solving this problem: taxon summation (Vane- consei'vatiori of l)iodiversity, sustainable use of tlu' V^Vight et al., 1991), and the use of higher taxa its components, and the fair and equitable sharing (\^'ilhams el al., 1994). Although furtluT work on of b<»nefils arising from the utilization of geru^lic methods both (which an* not necessarily exclusive) resources. As is needed, the point here is that systematists tend have already suggest<'d. this approach to I propose to very different sorts of solutions than liiological diversity ra(hcally different from pre- is ecologists or population biologists. vious concerns of the conservation movement, such Many ideas will need thorough (^valuation bt'lore as the pntteclion of and endangered species. we settle on the most a[>propriale information and or the prest^nation of wilderness areas. The diffcr- metliods for priority anâs analysis. Only by mobi- ence the anlhropocenlric focus: human rtdatt^s to lizing systematic data, and creating the means to needs constitute both the threat and the solution, interpret in a logical and systematic way, will we That is not to say that consenation will be com- it be able to develop efTicient {ilans and monitoring promis<ul: on the i-ontrary, conservation has shifted, schemes A for cons(»rving biodiversity. primarv' goal in th(M)ry, from pressun^-group status to being part is simply to make the most of limited resources of the fabric of human society. Conservation, and that. Inevitably, will never be enough to do every- specifically the needs and us(^s of biodiversity, are thing that might be consid<Med d(*sirable. Syste*m- now in the realm of what might be termed social Agenda atics 2()()0 can lead here by promoting a engineering. nelw<»rk of systematisls lo create the widt^ range of So biological diversity of a sudden, big is, all and spe(Mes-lev(d highcr-cat(*gory databases which, business. In relation to biological sci(Mice in g<Mi- togclhcr with appropriate analytical procedures, eral, and to the community of professional conser- will be needed f<M- a comj>rebensive approach to vationists, ecologists, systematists, and so on, this conservation evaluation. good However, Jiews. the arrival of biodiversity is new as a poHtical issue does not just signal sonnies need we of funds. to appreciate fully (unless orn \'^c KFspoNsiniLiTit:^ as sysi'iimatists allow ourselves to be prey to the worst sort of cyn- human Tlie burgtMniIng population, driven by icism), that biodiversity couched within a first is consumerism and pov<Mty, having a massively truly social framework (we are thus operating be- is deleterious effect on biological diversity, through yond the strict confines of science, in the areas of industrial pollution, resource ap[)ropriation, over- policy and soclo-economics), and second, we that We iropjn'ng, and ecosystem transformation. sec lo- have new and exf)anded responsibilities that w(\ cal and global extinctions on thtM>ne hand, an<l the the community of syst<Mnatists, rrmst face up to. sprea<l of a bmit*^! range of synanthropic species This includ(*s the fact that much of the best data on tlie oth<M-, leading to extinction of taxa and even about biodivt^rsity lies buried in i>ur c(»lleclions and entire ecosystems, extirpation of myriad popula- libraries. tions, and widespread loss of complementarity. Di- In my icw, veiy much part of our respon- is \ it versity is being diminished and homogenized. sibilltit^s lo lake control and help shape policy As human demand own We for resources in both the de- through our initiatives. h(»ld the best in- veloped! and developing world continues to grow, so sight into the strengths and weaknesses of taxono- the rate of human-induced biodiversity loss accel- my and systematics. We, veiy largelv, are paid out The unknown, erates. end-jjoinl is as are the con- of public funds, and thus hold the responsibility we sequences. All can say is that the biosphere, t>n not only to react supportivcly to tlu* needs of society which our life is totally dependt^nt, will be called at large, but also lo mobilize, make us(* of, com- upon lo pn»vide more and mon* food, timber, and nuinicate, and even lobby for the uni(]ue insights Volume Number Vane-Wright 55 83, 1 1996 and Systematics Conservation we eluding Deatli and the Compass, translated by A. tliat only are in a position to formulate or bring I). âtes from an original Spanish version of Pen- 1*>.'>2]. Our go beyond simply to bear. responsibilities giv- guin Books, Lxjndon. when we ing support asked: nmst also ensure that Caugliley, G. 1994. Directions in conservation biology. J. our krtowledge and understanding are brouglit to 215-244. Aniin. Ecol. 63: — the fore, to be judged useful—or not as otiiers Cousins, S. H. 1994. Taxonomy and functional biolic 397-419 measurement, Ark work? decide or, will the Pj). in P. & Humphries Vane-Wright L. Forey, C. R. (editors), J. I. Death and Compass the the of a short stoiy is title Systematics and Conservation Evaluation. Oxford Univ. A by Jorge Luis Borges (1970). criminal investi- Press, Oxford. & gator has been set the task of trying to discover the Cox, D. Williams. 1981. An account of cassava J. J. murderer Jewish At scene mealybug (Hemijitera: Pseudocoecudae) witfi a descrip- of a acad(Miiic. the the of new 247—258. tion of a species. Bull. Entouiol. lies. 71: crime, and in response to a suggestion that the Cracraft, 1995. The urgency of Iniilding global capacity J. scholar was accidentally nmrdered by somebody re- for biodiversity science. Biodiversity and Conservation man ally int(Miding to rob the living next door, the 463-475. 4: Why investigator replies, "Possible but not interesting. Ehrenfcld, D. 1988. j)ut value on biodiversity? Pp. & 212-216 in E. 0, Wilson M, Peter (editors), Bio- F. YouTl reply that reality hasn't the least obligation diversity. National Acaflemy Press, Washington, D.C. And to be interesting. answer you that reality C I'll & Ehrlich, R. C. Daily. 1993. Popuhition extinction P. may may avoid that obbgation but that hypotheses Amhio 64— and saving biodiversity. 22: r>8. & not. In the hypothesis that you propose, chance in- Eldreilge, N. N. Salthe. 1984. Hierarchy and evo- S, & Here have dead lution. Oxford Surv. Evol. Biol. (R. Dawkins M. Kid- terv'enes copiously. w<^ a rabbi; 1 184-208. editors) Icy, 1: would prefer a purely rabbinical (Explanation, not CD-ROM ETi/ P;92. Birds of Europe (Macintosh Version the imaginaiT mischances of an imaginary robber." Amsterdam. University of 1,0). I'.TI, A multitude of taxa are under threat of death. In Faitb. 1992. Conservation evaluation and pli>!oge- I). P. sympathy with Borgess investigator, wc should not netic diversity. Biol. Conservation 61: 1-10. 1994. Phylogenetic pattern and the (juantifica- avoid burden of providing an intellectually . tlu* B lion of organismal biodiversity. Pliilos, Trans.. Ser. We sound and need satisfying solution. a systematic 345: 4,5-58. as well as an ecological chart of the biosphere (Eld- The Harris, S. 1979. information content of the ph\- J. & redge Salthe, 1984; Rieppel, 1988), and should logcnetic system. Syst. Zool. 28: 483^519. develop a systiMuatic plan to outwit as many im- ~ 1983. riie logical basis of [ilivh)genctic analysis. & 7—36 unk Ad- P[). in N. Platnick V. A. (editors). we I. minent laxic deaths as possible. In sliort, must I New vances Columbia Press, in Clatlistics 2. niv. I wt endeavor what believe be an appro- find to to York. priate scientific solution, which takes full account Forey, C. Humphries & R. Vatie-WViglil (Edi- P. L., J. I. of the principles and insights of systematics, in the tors). 1994. Systematics and Cons<'rvalion Evahialion. Oxford Lni\. 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