ones oe we, Biological Systematics The state of the art Alessandro Minelli Dipatinento a Bitoga Universi ql stud ct Padova Padova hay Bandon «Glasgow» New York Tyo - Maboure Madrasny npn a temdy Re, andn SE HN gem Le oy Ldn EAN a An era Netra Rd, eg, Sopra gn ie Moh St Ne Yor NYO, LA (Sepa tye Doma ingpn lawns SSiviay altaya, Sept smn aN Ala, Do ee, See Na hee poet os a CF a, eet emma ‘petite BOS la ‘Pelican Pe ro Fa, Corl _aweensans ‘Sgttonn ideo pape ese icles ered ee, ‘eon ana ci Coruna Real ctnd penne Peleearalt te wal ond ied oe SEgetnosiny sehlmatces eee ‘Seen aspera areas [Aen heb nto ton esi bey "Ss irgeCtehetnatoten eras Knees, od For Federica ‘and Francesca £0700 sock 92Contents ace Pore Proce PART ONE: Problems and Methods 2 Systane and dasesatins Systematics and wxoromy 12 Cisctenionvers syn 13 Biologia chsifeation [rm Andrea Cesapino tothe New Systematics 1.4. Bvolsonary syeematice 13 Novel tion 16, Hennig’ plylogenatic systematics 17 Contrasting systematic choos 11S Towards utara tem of ving organs 2 Some stupa cempaatvebiaoy 1 Carcass for cong tra 22 hance or oke 23 Honclogy 24 Homopiey 25 Carer ety 26 Monophyly, sant, plypyly 2 Deminingchnmce ory 2 Cladogm ond ese 25 Nuarial mtd fer the resntucton of phylogeny 20 Ancestors 21 Rd date amy 212 Grouping sod aking 233 Popeyes paon x BeSSeeRRRsessContents Contents_ix 3 Bishenkc and molcelarsystematie 32 oom 432 como 4 Thenpedes 4 Sputetonape 42 Thuonnle dey within th pies 6 ieee Spun 5 Resouces mia SD Buseresues 52 stn 53. Liat SM Naneadtre PARTTWO: Theta ofthe Ast 6 ete fe ie & Howmiy wis owe 65 Hew many pcs fe lobe discovered? Rue ae eee wo B ™ GRBEERERRSEE ERE E DURES 722 Deaterostomes, exchding chords 15 1723 Chordates excluding vetbrates 18 724 Vertratee, w 1725 Green pln, excading sngleperme 164 726 Angiospenms 1s. {8 Intros on the daily work of ystems: problems and tends ‘81 Speci groupes natal gouge 168 82 Genera 19 83 Spaces 1B 84 nfespeic taxa 1% 85 Choraces 100 56 From led work to monograph im. 9 The unequal distibation of txonomic diversity 81° Thevery lange panera 15, 92 Sine dstbations of higher xa Ey 10 Domesticated animals and cultivated plants AGI" Taxonomy and nomenchtuo of fomestcted animals 198 102. Taxonomy and nomenclature of clvatd plans a [PART THREE: Epilogue 11 Some dangerous rend anda hope forthe future a Appendices 1 Zoologia cocks an et as 2 Meh’ 0980) gona cession of ving organs as 3 Provisional esteation’ ofthe Pots according Cos 221 4 Phyle and csses ofthe Protos (Cvs Prot), according to Margulis et (1990) m 15 Man's (960 cnnsfeation of animale Ea 6 Nike's (1955) dasseation ofthe Asumala 25 17 Halo (1985 yee of Pathekmines 2s 8 Jmieson’s (S88 cyte af the Oigochata 2 9 Salvin-Plwen's (860) ciasiation ofthe Phylum Motusen 251 10 Hayat 89 ction of oops = TL Weygoldeand Paulus (1979) syst of tha Chalcot, Be ‘2 Shuler (190) system ofthe Celcrta a 13 Sehr (1985) cnseaton a the Crstace 26 1 Starobogetov (088) esto ofthe Crustacea a 15 Hennig (968) aystem of the inset 2 16 Henig’ (1985) sytem ofthe Choedata 2Canons 1 Ther pups ef Carta secodng to Nakon 196) 18 Rosi) casieatonofgathostome vertebrates 19 Carls (8 sstenton of vertebrates, ncading both ‘tt ig bee 28 aude an Len 98 dasieaton of iving bony fishes 2 Se and Aqui 1590 casliation of binds 2 Diener (0595 cessation of green plats 25 Dagens (6b chsseton ofthe flowering plen’s Rye three Sot ier enone @ 8 SeOBR as ag Foreword ‘Tosome pel ede of thisbook the description of Bola ye ‘sas enat ay eu uted sd aly wing For mot popes ‘ubjectve and unconstrained by aver ave Wil ae pac lye poem muy be italy conden cmpaion between Gece at ode tenon ea by my ft nc tt, On Sher hand moder Boge yc ~ partly pence nd hdiey = i oflred ws bet and ural governed by unreal ins: Tis implies thor clathextons of dferese poupe of ante, being he prose of tena, should be comparable respecte of stoi “Doughou this ook Mil justifies hie te by dav the theme threo dance ae i ct very una n Ch xpreins the ptr snd proces othe msc ria Spel sre boot ‘eithtrown groupe and adie a coments wha costes 1 spedes or genus or whether i sould be dant fo recognize sub- Spec, cata ce: Ontology recogni elope Sn ely Ed genue conn more nn 10 apes On the cer band come ‘cieotetits and bts work with goer wih over 50 spaces Tis ‘Symmetry may ref» ology tay expres» wong Praca, or imply an Metoal rat der ected genre Se ey ey een ane ey ‘our anya probins ef ageogrepty,etionton of rots nanos ‘elution fbodveriy oe aero xincin depend on band oo: omc esnaent fama om sh ooo Bao Bowen " 7 Ml surveys modem etumatis methods and uggs tat th te development of dads, compur ana new mokcdar toe, Boge! syteatc of he tne sadvncig rpdly = and wh ‘wuld dlags However nfs tended eny one pace Capt) erent en ond vied See sboe ht cree a pee, the prolone ohh hase end nel condades tite apes concep may ‘atbeappropte ra orprins. “As hop towards desing the ot Bilge! Satematic, Mine (Pat bray then sation ofall sad ans point ‘Phere te moat gore ee of nership yatenatton arei Ferawort ‘ode found. Wh he lp of come 1450 references tie provides the eel ender witha wh Source of mater for discussion and fret, Lbvsigten, Pete Frey Natural History Museum Tendon UK Preface Bad taxonomy, of which there has been plenty, pests, Unite bad ‘hemistiy or bad physiology, of which there hae probably been equally 2s muct, it cannot be ignored; ft must be undone and ‘redone. Poor taxonomy isnot ony an il unto ise is contagious, ‘fen witha very lng incubation perod ..One assumes that when {experimental Biologists] stat thal they uéod 5 methanol, they were ‘ot using 6 mi of methanol and yet, ifthe experimental animal ‘wrongly identified, what are th grounds for such an assumption? DIK Me Kevan (1973, 1212) thi ook for flow profesional stage of development bar, fo my book hese won fran the place to Miles Ele’ Bvluionary Dyna Beg, 198, pl. ‘What prompted to wit ths pags, ws te Fein hal mal ete of olga temas eed 1 be posed one an eal whine page of ‘ne bak was propued io sie dp afesanet the aang of te asin coverage, eats tee are aa plenty of spsnlioed perma boo, ‘thre on can find out teat of pele tops. According i ot Fnened ss handbook ore reference Dok tr ber sa ese oad do ‘ics ope tht il rove of ore nts ce nt 1 flo tenis, ‘ral fo cer pop nth el of eutinary logy en ily a ge, Who ae inerested i geting a compres de of wit i o=cung toy ‘hin be oad eld of tema og. ate, bape at ame ptm lest wll provide a ring ad for ly techs deaing with syne ology nel cous. Bisogie!yeemates has ssconplished wemendoas development, tf ‘ioe arte or insu, a prclary nto a two ore eae, Tis oe part tena prope nto ass paral lec ology med ‘onpueeetniques, Molecular botogy ba opened anew dimetsion 1 the ‘ompunve study of vag gains whereas computers hve allowed teiv Pretece opt fet unt wagos ad iw iv A tes ogy soe) espe a “nr wo to ey peered ong te ny tof ee pepe aba sonal porn my ve, Be P- ‘steed eng se meat of log! yea ve Anew ee eee del open sce eal inc lig gen foe ‘cst sc ling a nga erie ‘oe rape emanate al np be pe ‘non ae peas not by hanes a fr even ok his steed ion, et aay tng ‘irons mon sn ter ete mah tee yu, oe sc ele Gn exe Dip, Reto fet) caret rome beg ch Hi Br Gin rte Pert Wi _ebitoms yyw eng. vce era Cnr ‘teig eos, Zaegi havent ep aioe ops bed ee coy Bd cn hav Hay ‘et he sort (amon nen easly HOS B13 ye magne pray pce erent {Ee 1982, Wins Bo pate ery oly aa ‘eset cin alan ene Se ea i abs ean wl ane SSirSe hime tel tinal eae om my eel! ecg, SLRs ely. Sul vs tp cod be del i DE es. 15, Boj SREM sous an tannic ace) contre senso Seeing png, PS WH Boe cereepod by dents of sells oF Soa eye gi Ret ns, i St pe mn pean al Sepa teat sw pps resect wel! Sec“ onesies rn se es ses PRA lsat esd as og sets tr en eB tne ww et ate vig fo oveccene the many puzze ‘ont, ‘Senemenoma ‘ese al tod me ‘Pars, Part One deals wit Issues, bo sSvettnegaa ib city wisn daca at Clap 2, ftiowed by a survey of concepts #8 Preface wv ‘oestoes inthe analysis of ptylogosi relonhips of ving oranians {Gaaper 2 Afri cell discussion (Chapter 3) i devoted to the aly evelping fel of moles tematic, wereasChuptr deen he ate neering species, sition snd infampecie tra, Ta at spar of sis rt dens ith th estou and med In systema, expeialy wih ate “eacenug Hirt sn menlare ‘Par Twos deed io asrvey of teats of ho Fit ow may spss go we know, and Bow ary spe do we eget disor (Chap 6), Wit Tows (Cher 7) is shor scout ofthe mor avant fate kets awlega nti he mor aa ofthe ving word (at sould ea eee ‘heh ook does nti at at comprehensive enya overage of all ‘jor ta sachs nth well known Syropus and Clsieaton of Lng Beige ‘ced by Sybil. ara (1982), Chapt soem cient expences Wah fly txombie work, ab wiaued by many ealegune Ive ierewed, on ‘ars sic secogaiten of genesis a sips, a cael teed indo! poe Theme apr a iy chs eect hn ad ‘rplzlnal improvements of jae waka wo ptr of Pat 79 se devted oles sul questons: oan ans of te ueeqadseibuton of ‘axonme diversty within te liven wee (Chega 9) al 1 te tomar ‘reatuet of domestic sia adclvated plans (Chap 10) at Thee is vty shor (Capi 11 only sn deals wit several problemas esti ish call ation, whe locking ovat fs. would ks slogan caomous debt te ny autos of he pat 1 yal asof te present, whose wots ave salted my view on Beopel ‘seas To member hem al would meceste duplicate teres ven a th eo of th ook. There, hee gute al themes of merous ‘ond and coleages who bate elped me de, Kindy aivering my spect ‘quest for infin fr this book BF Anietson, WR. aden L Ants), Argo, H- Auk Ase Biche, P. anti, DF Aas, B. Boos ‘Baldzee, EBs M, Baan, SCH, ret. Batol, Barsozs F Berit R Borla, RG, Bestel, M. Blond F. Boer, BL Bousol, H Brilowey, Ke Ber, Re Bek, §. Brookes Brow JD. Bure 1M, Capi, Carr F. Coan, D.C, Cally Swab E- Clonal, 410. Carle, Cane, BS. Ganson, M, Durer, G,Dalge,D. Decal, DAL Divs, W Doth. Dine. RP Be, K Fach, Frans yen PA. Fpl el Gap, Gr, GAP. Gs, Gitso, A, loan Sk, F. Gin, Gold, H. Goulet, WF. Gra R. Grol. Hoey CL. nse $8 Has, Rl Ha, Hels, LD, Monson YL. Hon. Mavi, Glee LAS, ohne, S18. Jones,Z. Katy DH, Kanan, ‘era, RE Kut LA, Kori, H-Xoyana, WP, Kestesen, BL La Gre, Niteving KG. Mens F. Magen, 3 Manes, BR Madi, A. Mei CD. Micherer, K Misa, ©. Meg Le Mou, L Manas ES. Nien, Pi Nimis B, Nordenstam, M, Oni P. Onoda, O.Oss, PPro, P asain Taree, B Palle, 1. Pegs, Jai, LP, Petiplene, MN. Pian, Wit, Pigvn, REG. Pedi Seenlil, DIG. Folch, R Pog Vwi Pelace Pahl Ragu, KRG, Rel Cle ES, Rose, R Row okay, 8. Rat A, Ratner, O.A Scr, 8. Seba, M. Sar, SC Schl GD. Shi HK, Senne, ML Sets, RT. Shah, R. Sil, MR Si 1A Sher, A. San, CX Siiber, M, Soline, We Ste 1. So Suns. Sys. Tar, LTavr M. Thay, LE. Tha. “Tong 6 Undriood CU, 0. Viggia A, Vig Tagian V Vos, NP Wagt, R Woles,E, Weber Wilby, Yamatai Te Zane 4 Zl ne a Stas Provo deat were very erful reeds evercveyetily CasHampars an Ptr Foey, wh ese) ms tee a, Witte hap fl te peopl ts boa could nove ave ten wit. Fonee yar not esposile forthe views hee ‘pee inte as My et towards ny ded cllegues at ie Natl ‘isty Musum aio) is een ge, bowever,teianing with Lane? Mol gag titer okay mt es years ag and ee it Pa ary kindy wing he ewer a nl io snova Casi Neonat del Ricerche od are, gant fom whch have alowed m= hese er and th ti tof Capa il oe pene ap re eee ily mu a fe lal ein of hes A. Mina Padova, gus 192 PART ONE Problems and Methods «th tatty fon sen ase avatae of atest some eo iy Sasa eng stb te bakin sane O° sun aytoms tsar ante poorest oct ree ‘vatags ool seater lazy ern tb clasiton al prove luca elorg ron ‘Sth otter bat assets fa fo advance Boogal Peey PR Stren (1986, 9.329)Systems and classifications 1 Ifthe goal of systematics isto dapict relationships accurately, then ‘any traction that intereres with this goa shoud be abandoned, de Quran M3, Donoghue (1988, 9.32) [LL SYSTEMATICS AND TAXONOMY ‘There o gene agreement tthe defines of jst, and of eat ‘ot, ach a tone, ptm and clusion, Capron | hse and reed was eyo page of ur ie, vena 184, do ‘sis amend teed ae of tems, ethoeand tse He ied his ‘el revo of th etre y banning any hope of estring te es ‘hat he slowed to be thir xg, inte ese, Posibly tn ae wort ty, tect ofthe despre inflections, ch at ‘Sinprn's (1961) well-known oes "Sytem ithe ii ny of he Kinds ad ves Fegan nd of sy atl slanstige mar er (Simpea, 1618.7» "Texcoomy ite theres of atts, nclading Hs es, nie, oceaes, sd les! (Simpson, 1961p) “Poclgica elscton ete ering of minal no gro4p or 8) 08 te ‘ase of hei relationships ats of socaos by cong, inary, orb (Sinpsa, 1961, 9) ‘iat Simpson refered ow systematics coud dsb ty compat ‘ve Woigy ton 1990, Wiley, 18H; Rlppel 98; se also Brooks ane MeLennn, 1991 Harvey sod Page, 198, The fd of sysmati oceney Uedtood Is nanos (Wie, 181), albugh ted only to prodteing ‘usifcatons and enueaon faye Sul more anfroat was Sion’ “kine xenomy, altho ary zbos se with Me eopsiing th lobes bac ot smean. Acrorancs wi apse’ elon, web ‘ped hori or teelgi probes he ete of tenon,4 Systems an classifications ‘cps gay see he prt anc of totes, Inde, Bisby (080, abbots aoe Reps (585) nl Hevheworl sn Disy (1985) rea ‘he pres of cto at noone fran sete comping Sse Aeals ronenctze, cscs, aniseed’ wheres esters ‘ke nanoy pis e legal inaaos = evn ystems und genet, ylgeny and evoiony sss, boggy and synecsogy' (Hews ‘a and Bt, 198, 9), However, a Havhavorh an Bisby (198, 10) ste wk te leeingy wile sony of elds cure ivestgned by ‘cronio o sae cssiter aongemets the seat fn pace tetvos eacnony ané ssa at deta above hat csne eae ive rc Biy' (98) per reviews fata dfs of yt See eto ho Ax (56, 1B, ep yaar ad toca ‘mony new men para lope ost (me's lt profes ‘csr fe Dents fom hoses kes sa ‘ttn niu ivy Cain (984, Seges (ely Dane a Howe (ED Maye (95 19), Hen (10h Dads Seat Se {72s Ro 88 Baer 7 Hage so Cav (90), Gran C980 HET Pe 940, yee ou), Wa ie (HE) Malwa USE, Rap ERY, Sa 50 le si (90, Stes (HD a Nye EA 12 cusssiteanion vans system Ace pay cgay SAG ey ont yea li = age oh coco of clasifean, We sl wicley Ste List cpus cncon oe pecan eet wet hr en sd sist he eration of in ves sae, ange of a (984 tte orig of ene Into eee ore mbt 58 Cer and hic 380d Ons HOHE, ale appre he (28 98 Mn (Dia my trad ee ee clog clssifeaon (auch a species, gener xd Sete hate te ea moore, AS ns ay. rr, chs, yam and Kingdon) oh Biological classifications 5 Mh e nda a, cng @ et ve mci er om ‘Accordingly, an, dg anf ose ae exes pce, Resse an Plone tn exelent Eshinndsnae ad Moca ate expt ‘A clasieaton is heehicl srnganest of ajc wid cass too tect ao place in these cls eee ty he deo mae ses at ‘he eg Fropets ofthe se i sen, avi ne re Pam ‘yet fa spose els cle Homa spent At high lel fy sch Caan o Rowe sal aclas, the emborof wch eegoe ares ‘sch ak Canis, Ros ia chs, te member of ih specs, Seem sso regard apse teases, the ms f wh inves ‘sas bt hs pots rch spats (Chap), By cons ih scien, ysam a eeseaton ofthe ving work ‘8 wines struct i the reo sme ral poss ugh wich oe ‘eof he system a elated. Orpen eid xagh to ifr peo ese: nerireding aad cmon drcen Inte cen oe tee 18 pie tron to Went pees oh nite ate ead 8 wholes, aber thn cles: Chap 4). Conon desert usa Recent and extn ving beings to ingle, rie of ch ve ore a ‘ou phylogunt system. In is vio, nit are te batches fhe Dbloseneic uve, rater han HlSilly ranged tbe fan lonnng ‘as ving beings thy arth ato abl mre Ieee etd ‘iin ner pars e sane wile Stageaig assem, aa thn asicatn te or ingen oaaae sli reerc, my sound uss, lv ove ested ne a Sept an unconenitonl, but Ei, view wienever we develop bee! ‘sents within plognae framework 13 BIOLOGICAL CLASSIFICATIONS, FROM ANDREA CESALTINO ‘TOTHE NEW SYSTEMATICS ‘Toca means aging objects ino groups. Line clas of nas ‘plans washer, 8 ae all asians in zologial and borcl ‘enooks A enon epllypreesppned at uel race by the provisos of tema! Cote of Nomen, wits sone pesible ‘sean at thence ee (Chap), ‘npn, Merrie chien col be evloped eer ina eed 42g onward) or a ascecng (ava) way. Downward clasfeton vat erly Ae Cie De et, by pig Aan ‘nn iets othe nie of plane pis known i. A ‘hase wo sc te tad) ping oe rel ets sede. THe proce she epee unl sat {icy oval of desianon has bn reached. Thi ened asda deat Seaton ey, wh eauples of nntasing ra ae presi or coe, lowing "eogaon of easly smi eof pecs. Asan open al f= i itieaton,downnerd clic very wetland ily el, How6. Salome and cleslfeations ‘iio i mated i. Sn sd cemepion oe {Pape vel bergen pce wie ekey ded pe oe ‘iow nian he onshore ttn coe “sg itr pel cso eh ay (19) ay av ben he Actuary ke Spas ea pi Bata, 10) at senna ptng maternal peso egy tre attr eh aig, re, lt) A ‘cg aod wa aes oe bya wierd pate cae oF States eee flowing Copan’ obey Ray's ence an Scr iin pe eg ope pope goal a. ee foalson cnt perros notary ens ‘We ling npr clo klstnsy ey enny tected ets heel tre wich ows eteion egy mae le ai Meron, ee cr or rong ‘tt i inchs wwe source ct he, ‘Er enn any, eer rng te eid ak, Th esol lt tnt of se omer ee inl eg an ces te i 985 a Ki CR api suf ey of "ee ten oped by Linea cnn of ction sc, Theis een of Ga led nl ‘ioc dip The sa stated fae eens Sect srr sted pile fr sity coe Cv 8.63): ne diated tng leas rlin,whera ost te septs Soe hee mci ihe ee inl vi whl fet, son compe ty plas. = (Me way, Cover cel opie iin eal goo weuble tsi es hye, te Sei oe At Mote nd Verba With SSUES title es dani by ttt mology The Siacmara ny tee li es faite th none ese eee ‘erie twee ea dear, 8 yee onal pheamena seh ab et Src pee Ome dec one) ep 9 ete enerrttion of spe asd dy dt pci seo ely cx ae “ene systema, Rouplyepsiog, ep Numerical taxonomy _7 Simites may be uaentond spot of rane in he plage we, ‘ving tgs ean be cased so geen, sii ai oon, esting some ar offs, rom seh ecm ner sgn ions, ‘Soon are pubiation of Darin (189) Oi ef Spec, walk, esa tact, dawns of phyogesti tes by Est Hecke! append (Ga Hhectl, 1865, 187). The way sere be ope fe heart Pe {oundson aod an opel efrmuion of oll csi Int, tte entusiastebepings ld down by Hackl fed to tevolsionize ut Selene Its 9% el ever sae ltr ta th Druin an Halon erage na welded genera entensvel recedes 14 EVOLUTIONARY SYSTEMATICS (Ove of the mont problnaic areas rived around te concep) apexes. rein of dent sere ton, ser aswell npaleonogy nd posi, prone th developer of ws urea known a "The New ‘Systematic he mar aleve of which ce mired a book of te same ‘amo (Hey, 1940) ae wel ain Rasch (1829, Maye (12,1963, Sebbins (4880) and Simpron (1961. Se ako Maye and Provine (1980) fra hs ‘ecoute. Later developments of hese debts on he sels, ening mani ‘Mound the r-aled bao! psi eee vcore Caper. ora log time, tees detlopmens Ie ha Myr (1982) cals mio. ‘wxonay, te sleoe of pects, an que deena am be couse at rmcrtecanny, the sinc of cision, Tere conmen rng from eit tat supp taxa Were maul ene, vague sitet ht = openers the mor marl How do we eet logs? "ow do we represent itn 9 fone esfeon? For may year, he mest ‘emproenive hortal ent on compare tan Renae, 1352), a8 ‘ella the ober wea textbooks elope ystenatis (6 Simp, 1845, 196i; Map, Linsey ané Ug, 195% Baye, 1968) dg not adtnce ‘ttl the theo tckgonné of sso, using te 194s ae 150, sents plied rnd few ting es ‘pzcly Ema tye and George Guard Singson, Lae, wih be dvlopet ‘of iret approaches to balgaleyemat, i ecane asim ety 4 Schl of Evolutionary Syezraescomssponting 10 the views of Mast ‘Sinpson an oer, We ave ge bt thse elt opel (0 Mayr, ‘Whas od has nied gener often nd eon logs ‘Atte sare me one cana simply grr th concep! ogres of east sds, we cance be simply ep wie Be prog of he Bol iontry Symes sho Su ee rent apache inet Sau espeily rearing maroon. 1.5 NUMERICAL TAXONOMY ‘ts compen eff develop ew dene pata apres ‘alos sycnns was aoe by Reba Sa we PELA Sh8_Syetums and casicatons ecto oe ance gmetad Eee rears eaetraanaeee vere ee aan ene nce ony ace ie aoe aaron mma anaemenr con’ Tnmionneniined mreegesctot Tc et ae ait ra setitneelccatemeas meee seipinunet emmneome Sie alt ee ete ta Scenes SSE asc anacn wifiraimcsasova rege mt a Sn Siseirr aes ompcaeemesamrens Sein unacraatemrenmesee iekena algo an ete hoaae eae citomitiran ate ae deen Frenette rane sinha seeape ome ee Soca ameoren esr = ‘is ron wih mas eb ns a otching Jagan ent be developed by hid. Sen ope pom ne “tls ty snd ach Sra fete Te cin ot oad non ae iN Salona ote, wa era Wee tt ae Lemos mivLocenetesysrmuaries sat Sots wok romped ei oe 2 tae debs one foundations ihe pce va tegen en oe ny Darna eee seme BED, Contrasting systematic schools_9 Hennig sressed the obelve mod for a though and comiseat us of legen information in ting Woe casio, Heder aoe sie rules for rerouting pylogecy ar wel ee ning paopeone ‘ees in cst scheme. 1.7 CONTRASTING SYSTEMATICSCHOOLS oni’ wosk ine polation rom Ono se, Neon 1571972, 1970) Brotin (197 and Paton C1982 er gle act gg ‘ary tee of te cnneny a oh pyr pl roe ‘secon wih te anos of talon! spwoxtes, Oe sta ‘eed ns prac or arto pone sonton {Breworng of cong an pce 2. moe feral ere Howe ‘Ser te mes xp bd eaten eps emis Gs eages e itu, y May C979 nd by Chi ON); te Hens 370 ay © gd et een Neale for etnsrecing pings bt oteion 1h the pyogenes information (the Beach ere of) sod be ‘ned een recon en gn Th ec we ‘od win mah empass in atexboky Mae ard Ak (199, Ths, ben tea an 1, he andes of on ant te she pip and mets of oma Wake pose! ace eat Aas af ts cfc al bezel srk Rens "inte Sr Ci Slt Sat Keil syn Nays el Seon’ velo Seni, The coe ‘lo ese tra has tere ie ment fr camp Ch 582 ‘tte ts rave ty eed eae te theo pn changing ete. Te Snes een ‘are ‘tn bon nny cee in eget tray party ‘n Syetematie Zoology (Hall, 198). Tae new ideas lo led to the foundation of new ‘certo psn 47, i: Name Toney Cle Wat ‘irene Koes fens et te oy Spe ly; ay bere eal apres Eve, vere Teepe adele! pooped oe ae Wi ni See fom i Ion Ths pop au ered jou ldo, foi ES. Depts cnet ten pied ore ate, coos aya sal tee ops vet evil an wt it, ‘Wn cnc for nal te sealed pn csi’ eo hte paso Sree pec pen cote wl, ‘ews eft one oso ey ren coi sein ae © eed ner {Segre (Cgrns) fon he of ‘ang npn ar sty Pern cs ave Ong eri cigs, a giving Wal te en a cl eo sage aoe ate Teale Be ews were dee happy fen cee nets RE10 _Systoms and classeations (G90, py scgyevoed sca ya epson hat waaes ‘hereon mensip ia oj pylogeads elerti, toss) Soenlogy aso cemen nsession estan “Te sy efhettee ca scoot dann mow cies, hres he dae eta dito eatnny seat lily. Hove igi dia oie ae ulin ee oman to sl sadn sige oa ge coal the opponents vaunay sy st npr, a fen rxoed in ity aeptvetea, a deat eigenen of Saal nd Sneath xe sty Pyare chien edeeud by Hemlg. Wit he passage of me, ani’ spzions fre recomssion of plogesy have propessively becroe ncaa wis ie nd agg fh berets of the evar onary systema sci, Hove, soa cote emi, or exo, ‘sping te wre fis sald pray una (een 2.9), which tee by itt ey Flows of vlna temas Rada tines oss be srsed ox many ceasons Hower Sve oft ane pees of erent scans tae beg to ako ege is et muy sexe opto Mayr and Alok (55) fave we ot fink to clarity he afrences beets eons anes sya to cosas ay expe it erent ec fhe fs appch Aa ine ote mst ep e estates vo cho five wi ms onplaged 6 Ne ‘Sy te cova is ace ae dein Capers, anna eft cenit clogs (rang deans depicing she bao sel hr etal et). Te weer, se scion 23, en homology), hess yniponaplic (Le shared d= renames, ose anes ‘nis level sees or er teatro) rc, ppc ance SS PR ‘Accor, ts ots {a2 Als tile) ‘Towards a natural system of ving organisms 11 “he dus for the essing specnntion is poi by wiles ison of eres, such a Git ocarecs, opal range or exnton a xe (May fale o gst). Paton and Sei (187) and Si ad Pate (1988) hve clegaly emonsbaed the exrenslydangeoos ares driving fom ‘ema lnhing ppl tain sve buon. apy xt sium to be extine whe thy ae sued by amor advned bony which eae spat of the ame non My pute exons we ths wthing tlethan "tazonomiexinesns ‘Within cladises, thre Isto ererion ake than clive insivenes, for esting the an of aon whereas evealonuyaysenais works wii [Refined Linaen) ranking rarer next ranked eorlng Tae dope of diference exhib eapes tthe 15 TOWARDS A NATURAL SYSTEM OF LIVING ORGANISMS {wood wheter tr athig nr nan ‘rel sini” tte te {en ora jem econ he ie verity ging ‘Scan wie tow oe wer To ac ¢ ‘en scoping every eng nt may wen ging ‘ivi ‘Giales ky tunnel of erblg At esuune beach Mowe IRby ons unveralhaguge. hs psy not by hare a se Sitolng senate fhe ut ae tn south prog 1 ‘Sco nage owing Jo Ray sed cms oe Wikis wen ‘ela wed spt ging elnino l pose cores {Si ing Howes, Ry au he pl sn ot be Re te Foss ryt ie, ng rh We wa wong oN {yond ea haa nPop! ange Rave, 92) {edo mach he prion te, Oo aay at, ice Adon ed ‘orto he wig ot Fee nage owe ong eh Foleo (emg te em mt sya (Nest etsy he emcee ng aid {Uae (2) Nal tenella dl fst io G's ei asin gun, C) Nol brian en ety saan hte) Nal use on dese, OGD, oe ol Gur (40) cont a tral ens Lean ms, The feo mans oe, b) rg te cae liony of ‘Soc and ete, ng rane x mode ol of Ov eh a ‘acs nr, T we a wh et Dero eo bn No wonder th ene ei ls ‘toa apie ml rey ese tee ovate with "ont pope. “ae pt av wn ce Ral (0 rs of bop een ote oy bon sin 1 osc el cst Frctons though be gos, nthe ve bk, hat ‘neered ais12 _Syetems and cessifeations itr ie a gs Shicceweainr tery Ehyecep ime eaerisay tenet! SDE een ategncaes SEES os Semen SoS ia Reames Earl rarer hn py at ty Seca ceo a sian terre Farkas tata iSgutneamala escape te Semaine catia ciara tale cecnetetacevaaneNarecae rneiaau deat Cae Sita ocd erect sora cap fiona as wit te earmgecs Semin ta ea (piensa Seterowhadanaa srs et a a oe Sis stn ees Saeed crc eset eae ae eer cer rocranay seetia sare Si erent cy eet oe Senescence irae rime hn seers Sreeaetan cca eos =e oan ats eStats anne Siemens ater ee clara tame: net vaccn D pat eee seer tly eal heelys ‘species of | ier wien ta So ema om tee imei) Towards a natural system of hing rganioms 18 4 natal na soold be menophyels? Petar pabuns ae of «me Sperone or intr, hoa lees of eed anos a De "ecogelae and sumed wit esse? Narang il poste monoplane "sk of pediing ltr of ane whic ny sises som Kido peal fever "for nomen rect the Lense eas mens ih e cently ny moron ronal Coes of Neeser Ax (198,57 ove ges ‘olla th om gee nae and “pee nn! ‘ete, and elcng therm with ame and “ocd mae’, repent Bt tebe fi Sree ong Zante eemamsnnr ne ntsc es amo ont motel Siren dui rater og SSSR helene SSRI See es mtn tne SER eigen tenes te Sener af Bano ten testo ea "a nym ete seo sci sEees SEER etn SSSI ae et ch ce Sool acpi spat ie rc ESepeton du ens clecinm tmet tn Seeley atte tno plo, iinet pe atest cnn eee escent Soucy ey Sepa eine it coir ey en aes en esters sy ig Specie cera i Sor i om np Ro Ct ccc os €9 Shiai ot Stead ae hearer fr hen ree: ew ence oe yc ry olen en arr Sete a etna ny cy ES Sg eaten serene erin coe Slens temo ovate Senrhsmatrae ise eae Dhylatic, of section 2.6) phylogenetic us“Tosa op Teuton af ena wi chasifeae ane of eet and psf tts we have need fo te pe-eveainary pel a hat ‘ean no oad, Tins of eats now a in he fe ae fount festa th bien of serie! reconston nd historia repeat: the remesturon an} epresentton of ney ps (OH, 192, 7.136, Some steps in comparative biology 2 A natural system is something we discover, not something we create MIT. bisa 1967, p30) 21. CHARACTERS AS ‘SYMPTOMS’ FOR RECOGNIZING TAXA, “Tpsteologlaly a mun sysem hse n where see viene ot nes te Onopcly, aaa sstem i based upon te obec really ‘hie it conespands” Asoringy, “cen clasifatin i xed yon an Uectaing ofthe eeteclaied and my race we eae ein. ‘eaal (Boh gostaes from Gas, 187, 182 In ts sense, care ‘Soul be reared a eympon’ of nse cer We a ying 0 eover ‘sndio tani ino tes (or csifenton). ‘Chater at theta of on Invests sult ofthe dv of living rpms cts ‘robin s wheter the econsuctn of gogey depends on psig ase ot Eyres stout how evelsion was, f 5 comin by ptm cai r= ‘iy her bald nly bs onion sr oo ts const by "ety ae ‘rar anaes lags and eset “Cobos thai fe lnonships npr) en be ed spr fhe cna ott preitns of ‘lutionay pate temming fem teases of De evolinary process” (Breage oo Cara, 1980, 0.325) “An idoyh usin of ie ev woud be ot of plc a ls Bok ute ‘ot lation aot simple tt hat evo has oxcored ad tat ‘rains threfore ined ins mre oes ete way so eno I ‘nat gat sry pages, nur clsifnion wa pylogeete sytem, rsutlon dee not nly ean sping Hepes (Chdopoes, pean. als ‘eas adaption, snd consti. When dei which ste spine and ‘hl ir ndvanced botnet abeones or pretence of wings, we emt ‘one cuter wih pes enls of resect and abwnes i a carer ‘ai, We ao eed koow somal aor the acon vale ofthe wig,18_Scmestepsin comparative biology lo pile pve sgiane of sabre, Sil nev shinny ing te deo of thew ep co et sn ons ef wing ontogeny wih eee veep ‘es Of an, i infrmton i ldom tial, be tl 3 ‘meth enocis tar endeavours in econtctigpse, seaman gat tel no af Soa ater (6 a ‘8 svchost beatae OO 22 GHARACtERS oR cHorcE Saas ie a ce and related mates; tet eum, by Win C98 Ghee 860) ad Eo Sea fe aoopencaly fle wands ee coms tik dng bre ale ene Steve ref ewer Hove aero apenas ty Pct on Riggs tai ero rin i by operationally defining a cnc st Cried intt eae ot aan ne mo a Coens, 2 eatin by explaining tha eam, Ye et ®# nophsine (Ma, 19S 201) ae He ate 1am eet ten ee Pot ese (8 "ee see Rn ‘hace (88 pat "ances ce at scm ye enn ran sy ates seit Spey en Be wr arta aoe an ere eae (Bes, he impos ietMation about their inteclationships, a 20% ‘Seer etter dram ge ot mee rte ta tes atta blog caste of ue i ayn I # Dior eiteia of choice for den ‘rena ee Dinesh any a Lo and enn gene = ‘Sethe a ct ied ty fe oye er er tee in folly scoepable es ‘lin te ey Sto pts pe vere ) extend thst connaoos variables camot pvt ter systematats, such ss Donoghee ens ect a ia on ch ee they te aa es Characters or choice _17 “These quslons fad os to the poten of eiacacer weighing Soul al erates te re the wae eal Inova asym analyae? I at bow dierent ave they tobe waged? “citing was contrary fo pote pnp, eeu seemed ateduce 3 ‘sbjectn is into an bie tecnigue, but odny the son of wiping shy bt in owe ares ene forearms by Fre (19, 1983), Neff (986), ‘Whose (140), Ste (186, ryt (1989 a Albert nd ihe (192), in ginis, one cou ferent bw weighting prion sné weighing « aserir. Weighing @ prion meme tat we deve ak dite of ‘erent aa before galego, ate, before king th mats where Tey ae ease. An exreme instance of weighing a prio ie etna uglet of iene fn We soul sl senior weighing df bo ‘hte we kao, fo example ts ites pel of aven molec change (ior rotate teeters ener, bse J Wel, rhe arta ie. pene of sre characte Ges nde hee les nomav, ace lor wah. Weighing «pater means weighing scoring the dibton of ra. ‘es within th daa mais For example lowering the weight othe careers "ht prove bees normative with avn Slow, or example See of ‘ml independent oeeuenss in dierent espe Ste tie of weighing @ ‘ose is ealy vena in xno sas, wheres te etna snd sili ofa pio weighing sl debt Tus vou of sone wel feat de mgng pcos erate now ne ey we eet ‘oder ose Mutat dependiace vera indepedeee of canctrs shoul be of aor eon ‘xm forall sudous, whatrer plore ulock, bt were very seldom Informed as there of fa. Only na er cae Go we ow he gente ‘ano of the chaaees wo std ad the plies of pene iad ‘Untortnatey same edunaney i genealy icl oavod in iting characters, ning ngsabty of campy eve el se pnts © “pint ineedepensnc. ry de nd oS the at hi ey nee a na strongly subj selves. Tis ay be ny of daring eves dl geabeto the, whch don nat gee with ir tout peyogets. omeve. its widely beled th sony else hears vary to cay o ‘en mus wef pyogenic somata, Inpaizal hy ae poe al. ‘ei. This was andere by Cross (198, p18) a ‘specily tre le ths ngiogems, invite evolinay tans etree differnt eclogi Sos eoql ini. Te carats ted sigs the fale ad ones ‘tangirpema ain ge mene hogs wc re cal ela aap ‘and srl vals hypo iy, pc plyealy, empty, fd apewiy: pocpy snd syea pesto; number parte ach nthe presen or tere of endear sper nese he euros development of theses and eee ta, sh lie Insc ed tan pnts ii 1967, 911) ane tt dapve carters18. Same steps incomparatve biology ‘oy wi Hey (1958 dca paces, the prod of anagem ik ‘eolon saat}. wieeas manatative chutes deny cas Ct ‘cts fe paloma te. doe! by eadogeneis). The ence \commguas fs mioton ca ferent kins of chara pal nea mf te wierd use of alos leetopboree) Sa ‘Buoys wie sen 24 Things ar ote simp, howe fe Eee p85) pits ou aoa i ae elatvely es 0 ea fins eof nal ence maple eter 082) [Btalcyos Tc cum oun ht ey ae nse eb nal est {GEM paz ier tn pave aowtege Tee question ade et hd ens fen baby, uch as cosa set Fe peta tet view, We canot ca wth these tenes 2 Setowerstes Sey Hoon frat ge! Caper he 23 wowotoer oy oy ea ; 2 npc cnt cpa ls mea electors maim han cor ese fase etartan rcanegn YS oF Seca ncsses may suggest someting le encom SS Weshoall not ge ht the cone of al thn chun 2 wold ot ett stk fo Over 1) hag oe RH fey. Delle discussions of te ene yn ae Haven ae ten et Some suds, howene, iting eepinent e yc Yailey of homology. In tis respect 2° ee ne TY Gon nd Fa (1983) Ty #2 hn cue na sae a feed dapat tof hel owy he whol ind Bt fea (Gaon, 1980, Honeve, San SSS eet ret adn a Fa ‘krone tpn lt MY coulé no sus tim per at Rasta antl iaemne e tan ‘Rae by rae Yip accept a catia depres of COMP asi a ag nan tet ete el sain emmaa (86) Ta ep of ceeopmessl C AB in dees level at whic ‘be rasolved. We Sista cine cee ‘Aled agen ae vs sto eaabhanneeepecc bons Ink of ng gh cthDe by Sar oe, protease soos me cmp h wructralcomespondcae is. SERIES Se Stor ene eM ey may be dropped we shld “ecm! eo Homology 18 “The frat question shoul be, wha seal compared? Evry ompusion, bit Intaat of bameloy oer caoot en wat an na wlinges compare vo objets edge and Cra 198 Rep, 985). Why ae me [ntti lscusig te posible homology oewesn he ving of id nd be forearm of man, wile worever ey to campars th we i we xual fn of Gish? Ou primi hypoese of Bonony ae ganerly ede by ‘ssn sinriy, neadigerous Knowledge abut te ernest een eda ts vrins relates, lung ote ypethse of aac tae ‘hom previous ss ofthese erin. The oust of evo wrk vl song att the meanings of new sates adhe vay f {ngimens Siri iota eto orhomelgy evr cn nates te todo diitned eomputson, Theor bomoloy willbe ought 'eaveen ‘Compal’ ebjers Inher anne ames fh n he <ésnon of borlogy given by NeGon (10, p78) "exe aed fo be nomopos my be sted, Simp tbe cmp ad wien compote a) ‘ie exter in an hype of cea condone Bu sects ston {st th brain of cu efoto creme ti is fou inven oF Brnolois “Te concept of honey dovlopd In onpraive momo. Beto Being sve is dem ne by Oven (1843), ad ast snapped nema fess consis way hy sever! eomprinvesnatomty expel by Edoane ‘Gotioy Sint Hine (1850, According te Goofy, he bevirng ely ot ‘nial forms eoacas the tue uy of set pln th existence of mF ‘Slows compre thes ts of afer nial Every bane in 2g onesondence witha giea bone aoa, na money or inst Mery comes prndoows cn be easd beyond the lt of tnonomic clas, fr exon Imam! with sh ad og wth ar. eatoy ae ted oie the Socal coresponicaces between animale belonging 1 deel phy (r ‘erhmaserets ows the term clea by Cave) In Gat’ eid tie conespeeing pars irdeto were elledslogus (x et) Aliesetomasing of htm ns) ‘Wen intedcing the tm borolgy, Onca ssa retined Gotoy’ concep of alent (niogue) Sat atte on al pee eel fox eset) fort ident ueoaitelprietGeviee by he Fach ant Joa hed! perspective, we en ender this poosopie! sit tovadsigtts. Ath same in, must eet ovoid i in ‘when tang" es cone it md lngusge a ing is so ‘ose within an evltonay foes “Tho no doubt tan else homology fen more rein wo te opin compared are disuly ele Hwee, cud guesien whe ‘esto be stl andr jst aaendn he concept of hogy Men erpacing vo vr closely ened rps I it eit peak of he logy baweenconpanble uruurs of two individ belong ote sane secs Ax (1968, forex, rues tha stat Fremont {peak of bomsogytateon pasa th sae ein? Taste esos at~ 20_Some stops in comparative bilagy ‘ove i dire ways by les sudo, [dtd the wont compebene ‘octet of honcogy. allowing irate nd nna comparisons (inal, 19t; el aa Pe, 159; Hares, 1990 Sere observed (82, ppt) tthe ifs prs of an anal are the repton of chor, AY apy semauzd by Riegel BBS, pl), Sees sei, pron revs stat by Boren (1751, Vig Aap CTG6) ad uni (106, They heaped, respectively, te egal of the eo sis he pk a hele fe body as vere: he equals of te tee ad poste apenas ofa Yenshete tie post eaonhip {etven he Bae fhe sl a he verbena envelopes y Gosek ‘Toecauaise teen sane realy eng pote Sy hs ‘stmt bun called honeromyo al honsony. ‘at Velen (198,07 overs hat te Boma osu win ven ‘iit we tii ily geet ee gms oan en ‘Yom oF: eh ofa mma On th saat at haope wie ala oul is if ody he mole ects tw Bal to ‘et ood lls These holga otecng coves in ems wiseaging cones haley my ts moe of teat ie a lows t cone umole een hghr past coca abe, ‘uss tnd ronan hse rps costing of modus (nec ary Yao {ess 21 mcessy meen it nolons of homens ‘6 etc te sar (De Pine 1593 Al Homology 21 ‘oo often simple egeencs snlaiesterveenmraolcule of moe or fs dsl reledcxgnins ae caledhameloges, dep the ong ets ised agaist his race by a auorsave roof cecal blogs Rese erat 98. ‘Ths is probably te best placer nuodue Wagner’ (198908) dino of src tod tiolapal eps of homely. Hil cones (eof me ie conce forte ysteutis) develo by refeene oe al ety of dese fom commen anes, whereas olga sone dvsep by reference to woe Iechanism, whch may be developmen ahr han 1 oes. stall concette om thos aspects of homology tht are of concen to 2 ‘stati eging wit Te detionsof therm,Lfllov bee Paterson's (€b82) elegant sualistan of enol, eveutonay, pent, cists od "alta intone, “Lett camp the tel defn given by Oven 188, p34), szzartng to which a homologue "te sme ep ner tery any fom fe, ‘in’, wit May's (98, p45) evoatenary dfn: hemolayes fxs (or sels of etre in vo o moe eguans we hes at on be acd bck tt sme feat rst) nbs commonaestor fhe organs accra, 'o Ax (198, p16, athr’s tana) Maye ein i tho ictal and logeliy widest poo means [of homology within ie conten of eter tionary theory’ Boek's (1877 pt) din Ceres (or ondions of {ear Into ore eran ae Hemelagos i ey stm pyogenes fea the same fue (othe sue coon ofthe fests) te tne amc sce of hse rans) aay look Weta be ot beet ‘he qulicaion “immed common tees” elled into aubeantie Ue reson Foran example of phsntcdeniton of homology. Jatine (1967, 130) sted tn Homeloy is ating mae than "the wen between pats hich ‘copy eorejonang relate postions nconpre suse he et ot ‘wo orgmie wheres ‘atria ion (ecru sya by Paterson (982) is th oF Blckweler (67 p16 "Carn sts me bloga, ch as he Wingo td te fee of he, an th se man, ad ss ‘be compared fly wi cach ate” Nobing a ah re Lane lo rvie some else deiltous Dilfoent characters ht ‘be rguded ss ussfrmaion of he same pal cht ae general ale ‘omeloprs" (oni, 1968, p38. ’Homaogy tte ely of he manly of ‘acu’ Hes ma Bawa 977, 0.) th last few yuu, ose debs on Hamokey hve developed see olor and lit oneps, wees peo a uiliarn compte les pond forthory-ined semaine Lt us ao ook mae dsl ae liste of tamolog a deeaped win aise, begining with Henng'sPlyogeneie Syma (1969) Beanie introsuce te ow widely appl! datncten beeen plsomarphy and epee ‘morph. We cn dete he plesomorpe dhe apomurple chs est