Biogeografía 1 January, 2007 Page 1 - John...

· Biogeografía 1 · January, 2007 · ·

· · Biogeografía 1 · January, 2007 ·

E D I T O R I A L

A new voice for biogeographers

In early 2006 Juan J. Morrone and Malte C. Ebach decided it was high time for a biogeographical society aimedat those academics, students and amateurs that are isolated from the publishing hubs of Europe and NorthAmerica, where most of the learned “international” societies focus all their attention. After all, most biogeo-graphical research is done by scientists living outside the USA or the EU.

Most western academics based in national institutions would find it unacceptable not to have access to atleast Nature or Science. Imagine a western European university or a North American natural history museumwithout a functioning photocopier, a decent library or access to any online journal. Unfortunately for mostLatin American institutions this is a reality. Without proper access to resources and the difficulty of publishingin a foreign language such as English, most non-European and non-US biogeographers remain unheard.

The Systematic and Evolutionary Biogeographical Association (SEBA) aims to provide a voice for biogeogra-phers through access to communication tools such as a multilingual list server (Biogeography Portal) and anelectronic Bulletin. The Biogeography Portal, currently with over 300 members, is an active repository for fo-rums and electronic documents such as pdfs. It has helped many to access the latest news and voice theiropinions on topics as diverse as South American endemism or the issue of “integrative biogeography”.

The SEBA Bulletin is also a repository for ideas, forums and reviews. In this first issue we have a lead articleon Otto Kleinschmidt and his influence on early 20th century biogeography, a personal journey through Croizat’slife in Venezuela, a biography of Julius Minding, and some book and software reviews.

We encourage biogeographers to submit their ideas, reviews and forum debates in English, Spanish or Frenchto future issues. Guidelines for authors and editorial information are on the last page of every issue.

We hope that you enjoy the first issue of the SEBA Bulletin!

Malte C. EbachJohn R. Grehan

Juan J. MorroneCo-editors, SEBA Bulletin, 2007.


L E A D A R T I C L E

Otto Kleinschmidt (1870-1954), biogeography and the‘origin’ of species: From Formenkreis to progression rule

BY DAVID M. WILLIAMS

Much has been written about the New York School ofzoogeography (and evolution), its basis founded onthe dispersal of organisms over a static and unchang-ing globe (Croizat 1958, p. xi). The idea was perhapsfirst graphically captured by Ernst Haeckel’s diagramof wandering humans (Haeckel 1870, Taf. XV, seeWilliams 2006 [2007]), a visual image that remains inuse today to illustrate the route humans wander fromtheir place of origin (Kohn 2006, Shreeve 2006). TheNew York School’s primary developers, in chronologi-cal sequence, are “W. D. Matthew (1871–1930), K. P.Schmidt (1890–1957), G. G. Simpson (1902–1984), P.J. Darlington, Jr (1904–1983) and G. S. Myers (1905-1985)” – one might also add Jack Briggs (Briggs 1981,see Nelson & Ladiges 2001) and much of what goesunder the heading of phylogeography (Avise 2000, Rid-dle & Hafner 2006 [2007], see Nelson 2004a, p. 134and Heads 2005b, p. 681).

Ernst Haeckel’s (1834-1919) influence on the de-velopment of evolutionary theory was probably moreprofound than Darwin’s: Haeckel was “the chief sourceof the world’s knowledge of Darwinism” (Nordenskiöld1936, p. 515). Among his many neologisms, Haeckelproposed chorology to mean the “…science of thegeographic and topographic spread of organisms”(Haeckel 1868, pp. 286-289, translated):

“I mean chorology, or the theory of the local distri-bution of organisms over the surface of the earth. Bythis I do not only mean the geographical distributionof animal and vegetable species over the different partsand provinces of the earth, over continents and is-lands, seas, and rivers, but also their topographicaldistribution in a vertical direction, their ascending tothe heights of mountains, and their descending intothe depths of the ocean” (Haeckel 1925, p. 364).

Haeckel credits Darwin with its instigation, as it is“…only since Darwin that we have been able to speakof an independent science of chorology…” (Haeckel1925, pp. 365-366). He continues:

“The most important principle from which we muststart in chorology, and of the truth on which we areconvinced by due examination of the theory of selec-

tion, is that, as a rule, every species has arisen onlyonce in the course of time and only in one place onthe earth – its so-called ‘centre of creation’ – by natu-ral selection…the distribution of the great majority ofanimals and vegetable species in regard to which thesingle origin of every species in a single locality, in itsso-called ‘central point of creation,’ can be consideredas tolerably certain” (Haeckel 1925, p. 367).

Chorology was, then, the “science of migrations”(Haeckel 1904, p. 98, Third Table) from a single local-ity, an organism’s “central point of creation”. For thepromotion and generalisation of the idea, William DillerMatthew’s Climate and evolution (1915), publishedwhile Haeckel was still alive, helped gain the idea re-spectability and influence.

In a recent history of panbiogeography, Headsnoted that:

“Matthew’s book [Matthew 1915] capitalized onthe fascination of the American public with vertebratepalaeontology which had existed since the time of Tho-mas Jefferson’s exploits and became the most influ-ential text in the history of biogeography. Its ideas werepromulgated by Simpson, Mayr, Darlington and oth-ers, forming what Croizat (1958) labelled the ‘NewYork school of zoogeography’ (cf. Croizat 1984, Nel-son and Ladiges 2001; for some reason Nelson andLadiges did not include Mayr in the school)” (Heads2005a, p. 89, 2005b, p. 63, 2005c, p. 681).

With respect to the omission of Ernst Mayr fromthe New York school, Nelson remarked that “Mayr doesnot derive from Matthew but rather from Stresemann”(Nelson pers. comm.). Croizat commented that “Thelatter [Mayr] in his turn, by grafting Stresemann uponMatthew, has greatly influenced North American, andworld-wide, ‘biogeographic’ thinking” (Croizat 1964a,p. 179, fn 1).

Writing of early panbiogeographers, or perhaps‘pre-panbiogeographers’, Croizat suggested that theornithologist Otto Kleinschmidt (1870-1954) was a pio-neer, if not creator of panbiogeography:

“Suffice it here to conclude that Kleinschmidt isone of the first true biogeographers, to the extent at


least that he correctly perceived that form-making is aprocess not to be dissociated from space and time. Insum, (pan)biogeography was not born in 1958 withme, but in 1897 with Kleinschmidt, whom everybodydoes follow today. Is this a droll state of affairs?”(Croizat 1964a, p. 182, see, for example, Croizat 1964b,p. 438, 1966, p. 92, 1977, p. 18).

Kleinschmidt was an influence on early 20th cen-tury ornithologists, including Erwin Stresemann (1889-1972). It would thus seem, potentially at least, thatthe biogeographic genesis of Mayr and Croizat mightboth lead back to Kleinschmidt.

Otto Kleinschmidt (1870-1954)and the ‘Formenkreis theory’

Otto Kleinschmidt was a leading ornithologist, prot-estant pastor, creationist and sometime Nazi sympa-thiser (Jordans & Peus 1950, Kleinschmidt 1950,Clancey 1950, Haffer 1997, pp. 791-802, Hoßfeld 2000,Pasternak 2002, Steigmann-Gall 2003, Junker 2004,p. 345; see Kleinschmidt 1933a, 1933b). Kleinschmidt’scontribution to the species debate was to note thatsimilar ‘forms’ (species) occur on separate and some-times distant geographic regions. To explain this facthe developed the ‘Formenkreis theory’ to account forthese closely related ‘forms’. While literally meaning‘form-circles’, the allusion to a circle was coincidentalas “the word ‘Kreis’ on the expression ‘Formenkreis’means the area of distribution of a certain animal onthe land or in the waters of the globe” (Kleinschmidt1930, p. 27, see Kleinschmidt 1926). The many similarforms occurring within a certain ‘type’ would have afixed suite of characters, detectable by examination.For Kleinschmidt, the Formenkreis suggested an un-changeable (but variable) entity, and hence allowedhim to support and promote a creationist view of theorigin of species – a viewpoint not unlike that of LouisAgassiz (see below).

Contemporary comment on Kleinschmidt is favour-able:

“I think there is little doubt that Kleinschmidt’sFormenkreis concept represents, as claimed by Mayr,“Systematics and the origin of species”, 1942, p. 112,‘one of the most productive working hypotheses oftaxonomy’, and while new work has necessitated theoverhaul of certain views expressed by Kleinschmidt(1900, 1930), the principles of the concept have beenwidely accepted in the scientific world and have wontheir author imperishable fame.” (Clancey 1950, p. 32)

Ernst Mayr and Erwin Stresemann

Haffer et al. noted that “After his high school exami-

nations (Abitur), Stresemann entered the University ofJena in 1908, where he took courses offered by ErnstHaeckel” (Haffer et al. 2000, pp. 399-400, see alsoHaffer 1997, p. 829). In 1931, Stresemann “declaredhistorical morphology as terminated, i.e. phylogeneticor systematic morphology…in the sense of ErnstHaeckel…and Max Fürbringer…” (Haffer et al. 2003:420). The meaning suggests that Stresemann relin-quished the pursuit of homology (the discovery of re-lationships among organisms, as in either “phylog-enetic or systematic morphology”) and embraced thebeginnings of what might be called the ‘genetic’ or‘populational’ point of view, the current evolutionaryZeitgeist.

Stresemann was said to have applied Kleinschmidt’sFormenkreis theory to his taxonomic work “withoutaccepting its underlying theoretical (typological) prem-ises” (Haffer et al. 2000, p. 410). Stresemann was alsosaid to have been an early proponent of the historical-dynamic method of zoogeographical analysis (Hafferet al. 2000, p. 414), of which an early version can befound in Stresemann & Grote (1929) but a more de-tailed explanation appeared in Stresemann (1939, seeVuilleumier 2005).

The historical-dynamic method of zoogeographicalanalysis

“…stressed the need to examine entire faunas in-cluding the dispersal capabilities and distributionalranges of each species as well as the ecological andgeological history of a particular region to understandthe zoogeographical history and differentiation of agroup of animals as a dynamic and continuing proc-ess” (Haffer et al. 2000, p. 414).

This ‘new’ method was contrasted with “the static,regional-geographical approach of earlier workers”(Haffer at al. 2000, p. 414). Later, Mayr (Mayr 1946, p.5) noted that the herpetologist Emmett Reid Dunn(1894-1956, see Schmidt 1957) “…was the pioneer ofthis concept [dynamic faunas]” (Dunn 1922). In Mayr’s1946 paper, he referred to two examples from Ger-man ornithologists, Stegmann (1938) and the previ-ously noted paper of Stresemann (1939) (Mayr 1946,p. 5, Mayr 1976 [1997], pp. 567-8). Yet, as noted above,the faunal approach, such as it was, reached back toHaeckel and is, in fact, a re-statement of chorology,the “…science of the geographic and topographicspread of organisms” (Williams 2006 [2007]).

Stresemann was professor to Ernst Mayr (Junker2003, Bock 2004). According to Mayr “…Virtually eve-rything in Mayr’s 1942 book was somewhat based onStresemann’s earlier publications” (Mayr 1999, p. 23,see Mayr 1997). According to Bock, Mayr’s “first andlast interest in ornithology is biogeography…”, wherehe applied “the (then) new ideas for analyzing the bio-geography of birds that were advocated by Stresemann(1939)” (Bock 2004, p. 645, 2005 p. 10, 1994 p. 291


and Vuilleumier 2005) – that is, “the historical-dynamicmethod of zoogeographical analysis” – chorology.

Species and their ‘parts’

The details and nomenclature for species and their sub-groups is unyieldingly complex, words and definitionstumbling from biological and philosophical journalsfor almost three decades (1930s to the 1960s) andbeyond (Wheeler & Meier 2000). In the run up to the‘modern synthesis’, Bernhard Rensch created two termsto represent different aspects of Kleinschmidt’sFormenkreis: Rassenkreis (circle of races) and Artenkreis(circle of species). Artenkreis relate to superspecies,the latter a creation of Mayr:

“Rensch [1928] has recently introduced into orni-thology the term “Artenkreis,” for a systematic unitthat was called “Formengruppe,” “Artengruppe,” etc.,by several earlier German authors. As there is just aboutas much difference between Artenkreis and species asthere is between species and subspecies, I propose forArtenkreis the more convenient term, superspecies. Idefine superspecies as a systematic unit containing geo-graphically representative species that have developedcharacters too distinct to permit the birds to be regardedas subspecies of one species” (Mayr 1931, p. 2).

Artenkreis relate to Huxley’s ‘geographical sub-genus’ (Huxley 1938, p. 255), while Rassenkreis (Rensch1929) relate to Huxley’s polytypic species (Huxley 1938,p. 255), “species that break up into subspecies” (Mayr1978, p. 86).

Mayr did, of course, modify his definition of super-species:

“A monophyletic group of entirely or essentiallyallopatric species that are too distinct to be includedin a single species” (Mayr 1963, p. 672, 1978, p. 86).Another composite, semispecies are “populationswhich have part way completed the process of spec-iation…” (Mayr 1963, p. 501), they are on the “bor-derline between subspecies and species” (Mayr 1978,p. 86, after Mayr 1940, p. 260), a definition differentfrom that in Mayr (1942, p. 165). For the 1942 semi-species – species comprising a superspecies – Amadonproposed instead allospecies (Amadon 1966, p. 245),hence a group of allospecies form a superspecies (=‘Artenkreis’). If not to complicate matters further, Mayrand Short (1970) introduced the ‘zoogeographic spe-cies’ for zoogeographic analysis:

“As the new polytypic species concept began toassert itself, a certain pessimism seemed to be associ-ated with it. It seemed as if each of the polytypic spe-cies (Rassenkreis) was as clearcut and as separated fromother species by bridgeless gaps as if it had come intobeing by a separate act of creation. And this is exactlythe conclusion drawn by men such as Kleinschmidt or

Goldschmidt. They claim that all the evidence forintergradation between species in the past was actu-ally based on cases of infraspecific variation, and, inall honesty, it must be admitted that this claim is largelyjustified” (Mayr, 1942: 114).

The ‘bridgeless gaps’ were Goldschmidt’s conclu-sion (Goldschmidt 1940, p. 310), the ‘separate act ofcreation’, Kleinschmidt’s (Kleinschmidt 1930) – andearlier, that of Agassiz.

More on Stresemann

Stresemann was also professor to Wilhelm Meise(1901-2002), who “was primarily a systematist study-ing problems of geographical variation, hybridisation,and speciation in birds” (Haffer, 2003, p. 117). Meisenot only worked and published with Mayr (Meise &Mayr 1930) but at the same time was mentoring the19 year old Willi Hennig (1913-1976), publishing twostudies on Rassenkreis of the reptile genus Draco, theonly lizard genus with the power of flight (Meise &Hennig 1932, 1935, Schmitt 2001, Haffer 2003).Hennig developed Rensch’s concept of Rassenkreis,creating the chorological method for determining char-acter polarity from geographical proximity, noting that“The use of the chorological method in zoology hasbecome known particularly through the books ofRensch” (Hennig 1966a, p. 133, Brundin 1966, see alsoHennig 1950, pp. 192-199 and Kiriakoff 1954), a viewthat led to Hennig’s ‘progression rule’ (Hennig 1960,1966b), the view that the direction of dispersal washypothesised as moving away from a particular cen-tre of origin, those places inhabited by the most basalor plesiomorphic species. Hennig’s ‘progression rule’was soon rejected (Nelson 1974, see Parenti 2006[2007]). Bernhard Rensch was also one of Stresemann’sstudents (Junker 2003).

Thus, one might easily understand Stresemann’sdescendants to include some of the leading propo-nents of the ‘modern synthesis’ (Mayr, Rensch) as wellas, albeit indirectly through Wilhelm Meise, WilliHennig – even including the following generation ofphylogeographers (Avise 2000), providing an intellec-tual lineage from Formenkreis to phylogeography viathe progression rule – or from Otto Kleinschmidt toHennig, according to Croizat, the former “one of thefirst true biogeographers” (Croizat 1964), the latterirrelevant (Croizat 1978).

Discussion

Both Kleinschmidt and Hennig, and perhaps all theothers discussed above, understood species in thegeographical dimension by investigating their ‘origin’,


their emergence – so to speak – from a single centreof origin, “the single origin of every species in a singlelocality, in its so-called ‘central point of creation’”(Haeckel 1925, p. 367). Efforts towards further under-standing have been hampered, not so much by thecreativity of various commentators, but more because‘origins’ were the focus of attention.

The idea of a single origin for organisms can betraced to the Bible, a view given substance by Linnaeus(“If we trace back the multiplication of all plants andanimals…we must stop at one original pair of eachspecies”). Darwin found the idea of a single centre ofcreation captivating:

“Undoubtedly there are many cases of extreme dif-ficulty in understanding how the same species couldpossibly have migrated from some one point to theseveral distant and isolated points, where now found.Nevertheless the simplicity of the view that each spe-cies was first produced within a single region capti-vates the mind. He who rejects it, rejects the vera causaof ordinary generation with subsequent migration, andcalls in the agency of a miracle. It is universally admit-ted, that in most cases the area inhabited by a speciesis continuous; and when a plant or animal inhabits twopoints so distant from each other, or with an interval ofsuch a nature, that the space could not be easily passedover by migration, the fact is given as something re-markable and exceptional” (Darwin, 1859, p. 352).

Mayr once wrote of the Origin that “…Darwin’sargument was directed against authors like LouisAgassiz, who explained discontinuous distribution ofspecies by multiple independent creations” (Mayr1982, p. 619). Agassiz wrote on centres of origin andindependent creations:

“The greatest obstacles in the way of investigatingthe laws of the distribution of organized beings overthe surface of our globe, are to be traced to the viewsgenerally entertained about their origin. There is a pre-vailing opinion, which ascribes to all living beings uponearth one common centre of origin, from which it issupposed they, in the course of time, spread over widerand wider areas, till they finally came into their presentstate of distribution. And what gives this view a higherrecommendation in the opinion of most men is thecircumstance, that such a method of distribution isconsidered as revealed in our sacred writings” (Agassiz1850, p. 181).

Agassiz saw his ideas as in conflict with the “sa-cred writings”, rather than embracing them. Agassizsaw the ‘origin’ of the same species occurring manytimes at many places, in the cases of reported disjunc-tions – much as Kleinschmidt did (Klienschmidt 1897).Others, in the post-Darwinian, evolutionary literature,have seen something of value in Agassiz’ approach tobiogeography, if not his explanation. Léon Croizat(1894-1982, see Nelson 1973, Craw 1984a, b, Zunino

1992, Llorente et al. 2000, Morrone 2000, Colacino &Grehan 2003), for example, while rejecting Darwin’s“captivating idea”, did not cite Agassiz – but he didcite (and praise, see above) Kleinschmidt.

Heads (1985) and Grehan & Ainsworth (1985) of-fered a view that the problem of species origin residesin the “nature of ancestors” (Heads 1985), such thatancestors are ‘polytopic’ rather than ‘uniform’, in thesense of their characters, a view that relates to Croizat’spolytopism (= “multiple origins”, Heads 1985, p. 209,2005, p. 105, Grehan 2006, see Croizat 1971 andAubréville 1969, 1974, 1975a, 1975b). This view canbe contrasted with the monotopic ancestor, a posi-tion said to have been adopted by both Mayr andHennig (Heads 1985, but compare with Rensch 1934,p. 102, figure 22, reproduced herein as figure 1). Thus,for Heads (1985) and Grehan & Ainsworth (1985), dis-junct species may have arisen by some internal ortho-genetic process, giving the illusion that the ‘same’species occurs in different places, thereby offering anexplanation for disjunction and (possibly, we will neverknow) satisfying Agassiz’ problems of their origin (com-pare figure 1a with Croizat 1964, p. 783, fig. 84 andHaffer’s interpretation of Kleinschmidt’s Formenkreisin Haffer 1997, p. 797, fig. 47].

The idea relates to Kleinschmidt’s Formenkreis, inwhich “Each Formenkreis has probably an independ-ent center of origin, an independent period of originand an independent rate of development, or in a word,an independent existence” (from Kleinschmidt 1930,

Fig. 1. After Rensch 1934, p. 102, figure 22: “Schemader Rassenkreis-Spaltung in der Zeitfolge”.


p. 118, see Stresemann 1936, p. 155). Kleinschmidtremained a creationist, as did Agassiz.

When Ernst Mayr died in 2005 there was a veritableflood of obituaries, the good, the bad and the ugly. Attimes, referred to as the greatest evolutionary biologistsince Darwin, some of Mayr’s philosophical pronounce-ments have become somewhat doubtful. Polly Winsor,for example, has begun to unravel the ‘typology-essen-tialism-morphology’ (the ‘populational’ viewpoint) story,her account bringing doubt to the accuracy of certaincontributions of Arthur Cain (1958, 1959) and DavidHull (1965). Mayr’s history of essentialism “in its broadsweep across the history of systematics, this story isnot merely inaccurate in particulars, it is wrong andharmful in its basic message” (Winsor 2006, p. 3) thewhole forming a “miserable history of depreciatory com-ment without a particle of truth” (Nelson 2004b). Andso it may be with the ever changing species concepts,and their new and more exclusive categories.

One might follow the argument through the gen-erations, contrasting Agassiz with Darwin, Kleinsch-midt with Mayr, Croizat with Hennig, the phylogeog-raphers with…well, no opponent has yet emerged. Onthe other hand, it might be that there really is no pos-sibility of resolution to the problem, other than pro-posing new ways of discovering the ‘true’ nature ofancestors, determining its characters and proposing amechanism to explain how those characters change(alter, transform) into their descendants.

In 1820 Candolle wrote a significant essay on bio-geography suggesting that “All of the theory of geo-graphical botany rests on the particular idea one holdsabout the origin of living things and the permanenceof species” (Candolle 1820, p. 417, translation in Nel-son 1978, p. 285). Perhaps it is now timely to set asidethe ebb and flow of ‘the origin of living things’, puttingthe populational approach to one side, regardless ofthe contrasting ideas concerning polytopism (= “mul-tiple origins”) or monotopism (= single origin), andexamine the classificatory approach to taxa and theareas they occupy, a problem that has at the very leastpossibilities of solution (Nelson & Platnick 1981).

References

Agassiz L. 1850. Geographical distribution of animals. ChristianExaminer and Religious Miscellany 48: 181-204.Amadon D. 1966. The superspecies concept. Systematic Zoology15: 245-249.Aubréville A. 1969. A propos de l’ “Introduction raisonnée a labiogéographie de l’Afrique” de Léon Croizat. Adansonia 9: 489-496.Aubréville A. 1970. Vocabulaire de biogéographie appliquée auxrégions tropicales. Adansonia 10: 439-497.Aubréville A. 1974. Origins polytopiques des angiospermes tropi-cales: 2. Adansonia 14: 145-198.

Aubréville A. 1975a. The origin and history of the floras of tropi-cal Africa: Application of the theory of the polytopic origin of tropi-cal angiosperms. Adansonia 15: 31-56.Aubréville A. 1975b. Geophyletic studies on the Bombacaceae:The origin and history of the floras of tropical Africa. Applicationof the theory of the polytopic origin of tropical angiosperms. Adan-sonia 15: 57-64.Avise JC. 2000. Phylogeography: The history and formation of spe-cies. Harvard University Press, Cambridge, Mass; London.Bock WJ. 1994. Ernst Mayr, naturalist: His contributions to sys-tematics and evolution. Biology and Philosophy 9: 267-327.Bock WJ. 2004. Ernst Mayr at 100: A life inside and outside orni-thology. Auk 121: 637-651.Bock WJ. 2005.Ernst Mayr at 100: A life inside and outside orni-thology. Ornithological Monographs 58: 2-16.Briggs JC. 1981. Do centers of origin have a center? Paleobiology7: 305-307.Brundin L. 1966. Transantarctic relationships and their significance,as evidenced by the chironomid midges. Kungliga Svenska Vetens-kapsakademiens Handlingar 11: 1-472.Cain AJ. 1958. Logic and memory in Linnaeus’s system of taxonomy.Proceedings of the Linnaean Society of London 169: 144-163.Cain AJ. 1959. Deductive and inductive methods in post-Linnaean tax-onomy. Proceedings of the Linnaean Society of London 170: 185-217.Candolle AP de. 1820. Essai élémentaire de géographie botanique.F. G. Levrault, Strasburg.Clancey PA. 1950. Some appreciative remarks on the work of DrOtto Kleinschmidt by a British avian taxonomist. Syllegomena Biolo-gica. Festschrift zum 80. Geburtstage von Herrn Pastor Dr. Med. H.C.O.Kleinschmidt, Lutherstadt Wittenberg am 13. Dezember 1950 eds,A. von Jordans & F. Peus, pp. 31-34. Leipzig: Wittenberg.Colacino C, Grehan R. 2003. Ostracismo alle frontiere della biologiaevoluzionistica: Il caso Léon Croizat. Scienza e democrazia ed. byM.M. Capria, Liguori, Napoli.Craw RC. 1984a. Never a serious scientist: The life of Leon Croizat.Tuatara 27: 5-7.Craw RC. 1984b. Leon Croizat’s biogeographic work: A personalappreciation. Tuatara 27: 8-13.Croizat L. 1958. Panbiogeography or an introductory synthesis ofzoogeography, phytogeography, and geology; with notes on evo-lution, systematics, ecology, anthropology, etc. Vol. 1: The NewWorld; Vol 2: The Old World. Bound as 3 vols. i-xxxi, 2755 pp. Pub-lished by the author, Caracas.Croizat L. 1964a. Space, time, form: The biological synthesis. i-xix,881 pp. Published by the author, Caracas.Croizat L. 1964b. La distribution des Bombacacées misu au pointbiogéographique. Adansonia Ser II, 4: 427-455.Croizat L. 1966. L’Âge des angiosperms en general et de quelquesangiosperms en particulier. Adansonia ser. II, 6: 65-104.Croizat L. 1971. Polytopisme ou monotopisme? Le cas de Violaparvula Tin. et de plusieurs autres plantes et animaux. Boletin dela Sociedade Broteriana 45: 379-431.Croizat L. 1977. Carlos Darwin y sus teorías. Boletín de la Academiade Ciencias Físicas, Matemáticas y Naturales, Caracas 37: 15-90.Croizat L. 1978. Hennig 1966 entre Rosa 1891 y Løvtrup 1977:Medio siglo de “Sistemática Filogenética”. Boletín de la Academia


de Ciencias Físicas, Matemáticas y Naturales, Caracas 38: 59-147.Croizat L. 1981. Biogeography: Past, present and future. In Vica-riance biogeography: A critique. Nelson G, Rosen DE eds. pp. 510-523, Columbia University Press, New York.Croizat L. 1982. Vicariance/vicariism, panbiogeography, “vicariancebiogeography,” etc.: A clarification. Systematic Zoology 31: 291-304.Croizat L. 1984. Mayr vs Croizat: Croizat vs Mayr – an enquiry.Tuatara 27: 49-66.Croizat L, Nelson G, Rosen DE. 1974. Centers of origin and re-lated concepts. Systematic Zoology 23: 265-287.Darwin C. 1859. On the origin of species by means of natural se-lection, or, the preservation of favoured races in the struggle forlife. John Murray, London.Dunn ER. 1922. A suggestion to zoogeographers. Science 56: 336-338.Goldschmidt R. 1940. The material basis of evolution. Yale Uni-versity. Press, New Haven.Grehan JR. 2006. Non-Darwinian evolutionary mechanisms. In:Encyclopedia of Anthropology Birx JR ed. Sage Publications, Lon-don, pp. 1747-1750.Grehan JR, Ainsworth R. 1985.Orthogenesis and evolution. Sys-tematic Zoology 34: 174-192.Haeckel E. 1868. Natürliche Schöpfungsgeschichte. Gemeinver-ständliche wissenschaftliche Vorträge über die Entwicklungslehreim Allgemeinen und diejenige von Darwin, Goethe und Lamarckim Besonderen über die Anwendung derselben auf den Ursprungdes Menschen und andere damit zusammenhängende Grundfra-gen der Naturwissenschaft. Berlin, xvi, 568 Pp.Haeckel E. 1870. Natürliche Schöpfungsgeschichte. Gemeinver-ständliche wissenschaftliche Vorträge über die Entwicklungslehreim Allgemeinen und diejenige von Darwin, Goethe und Lamarckim Besonderen über die Anwendung derselben auf den Ursprungdes Menschen und andere damit zusammenhängende Grundfra-gen der Naturwissenschaft. Berlin, 688p. 2nd Edition.Haeckel E. 1904. The wonders of life: A popular study of biologi-cal philosophy. Watts, London. Supplementary volume to The rid-dle of the universe. Translated from: Die Lebenswunder, 1904.Haeckel E. 1925. The history of creation: or, The development ofthe earth…. Henry S. King, London.Haffer J. 1997. Reminiscences of Erwin Stresemann: Teacher andfriend. In “We must lead the way on new paths”, The work of Hartert,Stresemann, Ernst Mayr – International Ornithologists, ed. by J. Haffer,Ornithologen-Briefe des 20. Jahrhunderts. Ökologie der Vögel 19.Haffer J. 2003. Wilhelm Meise 1901-2002, ein führender Ornitho-loge Deutschlands im. 20. Jahrhundert. Verh. Naturwiss. Ver.Hamburg 40: 117-140.Haffer J, Rutschke E, Wunderlich K. 2000. Erwin Stresemann 1889-1972 – Leben und Werk eines Pioniers der wissenschaftlichenOrnithologie. Acta Historica Leopoldiana 34: 1-465.Heads MJ. 1985. On the nature of ancestors. Systematic Zoology34: 205-215.Heads MJ. 2005a. The history and philosophy of panbiogeography.Regionalización biogeográfica en Iberoamérica y tópicos afines.Llorente J, Morrone JJ eds. pp. 67-123. Universidad NacionalAutónoma de México, México, D.F.Heads MJ. 2005b. Dating nodes on molecular phylogenies: A cri-

tique of molecular biogeography. Cladistics 21: 62-78.Heads MJ. 2005c. Towards a panbiogeography of the seas. Bio-logical Journal of the Linnean Society 84: 675-723.Hennig W. 1950. Grundzüge einer Theorie der phylogenetischenSystematik. Deutsche Zentralverlag, Berlin.Hennig W. 1960. Die Dipteren-Fauna von Neuseeland als systemat-isches und tiergeographisches Problem. Beitrage zur Entomologie10: 221–239.Hennig W. 1966a. Phylogenetic Systematics. University of IllinoisPress, Urbana [Reprinted in 1979, 1999].Hennig W. 1966b. The Diptera fauna of New Zealand as a problemin systematics and zoogeography. Pacific insects monograph 9,81pp [Translation of Hennig, 1960, with additional footnotes].Hoßfeld U. 2000. Formenkreislehre versus Darwinische Abst-ammungstheorie. Eine weltanschauliche-wissenschaftliche Kon-troverse zwischen Otto Kleinschmidt 1870-1954 und Victor Franz1883-1950. Anzeiger des Vereins Thüringer Ornithologen 4: 1-26.Hull DL. 1965. The effect of essentialism on taxonomy-two thou-sand years of stasis. British Journal for the Philosophy of Science15: 314-326, 16: 1-18.Huxley JS. 1938. Species formation and geographical isolation.The Proceedings of the Linnean Society of London 150th Session1937-38: 253-264.Jordans A, Peus F. 1950. Syllegomena Biologica. Festschrift zum 80.Geburtstage von Herrn Pastor Dr. Med. H.C. O. Kleinschmidt,Lutherstadt Wittenberg am 13. Dezember 1950. Leipzig: Wittenberg.Junker T. 2003. Ornithology and the genesis of the synthetic theoryof evolution. Avian Science 3: 65-73.Junker T. 2004. Die zweite Darwinische Revolution: Geschichtedes synthetischen Darwinismus in Deutschland 1924 bis 1950.Marburg: Basilisken-Presse.Kiriakoff S. 1954 Chorologie et systématique phylogénétique. Bulle-tin et Annales de la Société Royale Belge d’Entomologie 90 :185-198.Kleinschmidt A. 1950. Leben und Werk. Syllegomena Biologica.Festschrift zum 80. Geburtstage von Herrn Pastor Dr. Med. H.C. O.Kleinschmidt, Lutherstadt Wittenberg am 13. Dezember 1950 eds,A. von Jordans & F. Peus pp. 1-31. Leipzig: Wittenberg.Kleinschmidt O. 1897. [No title]. Journal für Ornithologie 45: 518-519.Kleinschmidt O. 1926. Der weitere Ausbau der Formenkreislehre.Journal für Ornithologie 74: 405-408.Kleinschmidt O. 1930. The Formenkreis theory and the progressof the organic world: A re-casting of the theory of descent andrace-study to prepare the way for a harmonious conception of theuniversal reality. London, H.F. & G. Witherby.Kleinschmidt O. 1933a. Kurzgefaßte deutsche Rassenkunde.Armanen-Verlag, Leipzig.Kleinschmidt O. 1933b. Blut und Rasse. Die Stellung des evange-lischen Christen zu den Forderungen der Eugenik. Unter Zugrun-delegung eines am 18. April 1933 auf der zweiten Konferenz evan-gelischer Akademiker in Hannover gehaltenen Vortrags. VerlagMartin Warneck, Berlin.Kohn M. 2006. Made in Savannahstan. New Scientist 19 (2558):34-39.Llorente J, Morrone JJ, Bueno A, Pérez R, Viloria Á, Espinosa D.2000. Historia del desarrollo y la recepción de las ideas pan-biogeográficas de Léon Croizat. Revista de la Academia Colombiana


de Ciencias 2493: 549-577.Matthew WD. 1915. Climate and evolution. Annals of the NewYork Academy of Sciences 24: 171-318.Mayr E. 1931. Birds collected during the Whitney South Sea Expe-dition, XII. Notes on Halcyon chloris and some of its subspecies.American Museum Novitates 469: 1-10.Mayr E. 1940. Speciation phenomenon in birds. American Natu-ralist 74: 249-278.Mayr E. 1942. Systematics and the origin of species. ColumbiaUniversity Press.Mayr E. 1946. History of the North American bird fauna. WilsonBulletin 58: 3-41 [reprinted and modified in Mayr 1976 [1997].Evolution and the diversity of life. The Belknap Press of HarvardUniversity. pp. 565-588].Mayr E. 1963. Animal species and evolution. Belknap Press ofHarvard University Press, Cambridge, Mass.Mayr E. 1976 [1997]. Evolution and the diversity of life: Selectedessays. Belknap Press of Harvard University Press, Cambridge, Mass.Mayr E. 1978. Origin and history of some terms in systematic andevolutionary biology. Systematic Zoology 27: 83-88.Mayr E. 1997. Reminiscences of Erwin Stresemann: Teacher andFriend. In “We must lead the way on new paths”, The work ofHartert, Stresemann, Ernst Mayr - International Ornithologists.Haffer J. ed. Ornithologen-Briefe des 20. Jahrhunderts. Ökologieder Vögel 19: 848-855.Mayr E. 1999. Thoughts on the evolutionary synthesis in Germany.In die Entstehung der Synthetischen Theorie: Beiträge zur Geschichteder Evolutionsbiologie in Deutschland 1930-1950. Junker T, EngelsEM. pp. 19-30 Verlag für Wissenschaft und Bildung, Berlin.Mayr E. 1982. Review of Nelson G. and D.E. Rosen 1981 eds.Vicariance biogeography: A critique. Columbia University Press NewYork. The Auk 99: 618-620.Mayr E, Short LL. 1970. Species taxa of North American birds. Acontribution to comparative systematics. Publications of the NutallOrnithological Club 9: 1-127.Meise W, Mayr E. 1930. Theoretisches zur Geschichte des Vogel-zuges. Der Vogelzug 1: 149-172.Meise W, Hennig W. 1932. Die Schlangengattung Dendrophis.Zoologische Anzeige 99: 273-297.Meise W, Hennig W. 1935. Zur Kenntnis von Dendrophis undChrysopelea. Zoologische Anzeige 109: 138-150.Morrone JJ. 2000. El tiempo de Darwin y el espacio de Croizat:Rupturas epistémicas en los estudios evolutivos. Ciencia 51: 39-46.Nelson G. 1973. Comments on Leon Croizat’s biogeography. Sys-tematic Zoology 22: 312-320.Nelson G. 1974. Historical biogeography: An alternative formali-zation. Systematic Zoology 23: 555-558.Nelson G. 1978. From Candolle to Croizat: Comments on the his-tory of biogeography. Journal of the History of Biology 11: 269-305.Nelson G. 2004a. Cladistics: Its arrested development. In Milestonesin systematics. Williams DM, Forey PL eds. pp. 127-147. CRC PressFlorida.Nelson G. 2004b. [Untitled-Debate on Phylocode]. http://www.systass.org/events_archive/phylocode-debate.html.Nelson G, Ladiges PY. 2001. Gondwana vicariance biogeographyand the New York School revisited. Australian Journal of Botany

49: 389-409.Nelson G, Platnick NI. 1981. Systematics and biogeography:Cladistics and vicariance. Columbia University Press, New York.Nordenskiöld E. 1936. The history of biology: A survey. Translatedfrom the Swedish by Leonard Bucknall Eyre. New ed. Tudor, New York.Parenti L. 2006 [2007]. Common cause and historical biogeogra-phy. Biogeography in a changing world. Ebach MC, Tangeny RSeds. Pp. 61-82. CRC Press, Boca Raton.Pasternak P. 2002. Themata Leucoreana. 177 Jahre, ZwischenUniversitätsschließung und Gründung der Stiftung Leucorea: Wis-senschaft und Höhere Bildung in Wittenberg 1817-1994. Elbe-Dru-ckerei Wittenberg GmbH.Rensch B. 1928. Grenzfälle von Rasse und Art. Journal für Orni-thologie 76: 222-231.Rensch B. 1929. Das Prinzip geographischer Rassenkreise und dasProblem der Artbildung. Berlin.Rensch B. 1934. Kurze Anweisung für zoologisch-systematischeStudien. Akademische Verlagsgesellschaft, Leipzig.Riddle BR, Hafner DJ. 2006 [2007]. Phylogeorgaphy in historical bio-geography: Investigating the biogeographic histories of populations,species, and young biotas. Biogeography in a changing world. EbachMC, Tangeny RS eds . pp. 161-176. CRC Press, Boca Raton.Schmidt KP. 1957. Emmett Reid Dunn, 1894-1956. Copeia 1957: 75-77.Schmitt M. 2001. Willi Hennig 1913-1976. Darwin & Co.: EineGeschichte der Biologie in Portraits II. Jahn I, Schmitt M eds. pp.316-343, 541-546. C.H. Beck, München.Shreeve J. 2006. The greatest journey. National Geographic 2093:60-69.Stegmann B. 1938. Principes généraux des subdivisions ornithogeo-graphiques de la region paléarctique. In Faune de l’URSS, n.s. 19Oiseaux 12.Steigmann-Gall R. 2003. The Holy Reich: Nazi conceptions of Chris-tianity 1919-1945. Cambridge University Press.Stresemann E. 1936. The Formenkreis-theory. Auk 53:150-158.Stresemann E. 1939. Die Vögel von Celebes. Zoogeographie. Jour-nal für Ornithologie 87 312-425.Stresemann E, Grote H. 1929. Verbreitung und Gliederung afri-kanischer Formenkreise. Verh. 6th Int. Ornith. Kongr. Kopenhagen1926: 358-374.Vuilleumier F. 2005. Ernst Mayr’s biogeography: A lifetime of study.Ornithological Monographs 58: 58-72.Wheeler QD, Meier R. 2000. Species concepts and phylogenetictheory: A debate. New York: Columbia University Press.Williams DM. 2006 [2007]. Ernst Haeckel and Louis Agassiz: Treesthat bite and their geographical dimension. Biogeography in achanging world. Ebach MC, Tangeny RS eds. pp 1-59. CRC Press,Boca Raton.Winsor MP. 2006. Linnaeus’s biology was not essentialist. Annalsof the Missouri Botanical Garden 93: 2-7.Zunino M. 1992. Per rileggere Croizat. Biogeographia 16: 11-23.

David M. WilliamsDepartment of Botany, The Natural History Museum,Cromwell Road, London SW7 5BD, United [email protected]


F O C U S A R T I C L E

Journey to Coro

BY JOHN R. GREHAN

My entry into biogeography came about in a ratherunpredictable way. Biogeography was the last thingon my mind during the early 1980’s. But I then learnedabout the biogeographic interests of Robin Craw andMichael Heads and the rest is history.

By early 1988 New Zealand was in the grip of the1987 stock market crash and there was a correspond-ing crash in government support for science research.The entire New Zealand scientific establishment wasin turmoil as positions were slashed and opportuni-ties lost. This was no time to begin one’s scientificcareer. Having finished my PhD (on the evolutionarybiology of the ghost moth Aenetus virescens) I de-cided to make a seven day visit to Coro (Venezuela),where Croizat lived and worked for the last years ofhis life before passing away in 1982. Michael Headshad earlier visited Coro to study Léon Croizat’s life’swork as a guest of his wife Catalina Croizat.

In 1976 Léon and Catalina Croizat moved to Corofrom Caracas and became the first directors of the JardínBotánico Xerófito, now named after Croizat. Catalina(see figure 1) kindly agreed to my visit and I found my-self traveling to Venezuela in July 1988. At that time Ihad never traveled beyond the South Pacific (Australia,New Caledonia, and Fiji). Since there was to be an En-tomology Congress at the University of British Colum-bia that included a symposium on primitive moths, itseemed like a good opportunity to combine this withmy visit to Coro. I flew to Canada via the US and spentthe next five days in Vancouver. From there I was joinedby Diane, a friend from New Zealand, and the two ofus flew down to Caracas via Chicago and Miami. Nei-ther of us spoke Spanish so it was by a bit of luck andperseverance that we managed to transfer ourselves tothe domestic terminal and locate our flight to Coro.The next leg of the journey was an hour’s flight alongthe Caribbean coast where we saw a dry, desert-likeregion of low scattered vegetation, farmland, and sanddunes. We arrived at a small airport not much unlikethat of a rural city in New Zealand, with its open land-scape and small terminal.

After disembarking our next challenge was to findour way to Catalina’s house. We had her address, butknew nothing about local transportation. But as we

were waiting for our luggage we noticed a small, eld-erly woman smiling at us and we wondered if shemight be Catalina. After a tentative enquiry we foundour assumption to be correct. I think we stood outfrom the usual crowd. I had sent Catalina our travelschedule but we did not know if it was received (mailwas then sometimes unpredictable). As she drove usto her home we noticed that she drove quite slowlyand with seeming disregard of other vehicles; how-ever, we were later told that everyone in the town knewand liked her and made allowances for her driving.

The Croizats’ house was enclosed by a high walltipped with broken glass. This was a common defenseagainst forced entry in Coro. I was to learn that secu-rity was a major challenge due to the great disparityin economic prosperity in the region. After Catalinaendured the experience of having a burglar enter andthreaten her, the University arranged for a night secu-rity guard. Her house was surrounded by a wealth ofeuphorbias and other strange (for a home grown NewZealander) tropical plants. At the back of the housewas a guest house where we stayed. The center of thehouse comprised a large, open lounge with a highceiling and large windows at the top. The walls werelined with paintings, including some of Croizat (Fig.2). Each morning Catalina, with help from a maid, pre-pared a breakfast of what were to us exotic fruits anddrinks and other food. It seemed that we never hadthe same thing twice. It also seemed as if we couldnever eat enough to satisfy what Catalina thought weshould have!

My purpose in coming to Coro was to see the bo-tanical gardens, see where Croizat worked, and see someof his works I had not previously accessed. Croizat’s of-fice was left much as it was when he died, and onecould almost feel a presence in the room’s quiet soli-tude. Unfortunately, I never took a photo of this space,but I remember the walls being lined with an array offascinating books and publications. He even had a wellworn, and annotated, copy of Brundin’s TransantarcticRelationships. Looking back I realize that there was muchI overlooked – probably a combination of inexperience,culture shock, and jet lag; however, I felt I was in myelement and in the midst of a unique experience.


Fig. 1. Catalina Croizat in her office at the Jardín.

Fig. 2. Artistic caricature of Croizat by Celedonio Otaño.

During our stay we met a number of kind and sup-portive people, including Dr. Rodolfo Bastidas, an en-tomologist at the Coro campus of the Universidad Fran-cisco de Miranda and chair of the Friends of the Jardínwhose members worked to maintain funding and othersupport for the botanical garden operations. Bastidaskindly took Diane and me on a day-long trip to thecoastal ranges behind Coro to see the local region andpeople. We drove through a region of low, dry veg-etation with small scattered towns and houses. As wedrove higher into the hills we entered an area of cloudforest that was maintained largely by atmosphericmoisture. In some ways the vegetation was not muchunlike parts of Australia with widely spaced trees, lackof undergrowth, and dry leaf litter. I looked for signsof ghost moth damage (ghost moths are my primaryentomological interest) but without success. As wewere driving back down towards Coro the roadzigzagged back and forth with extremely tight turnsdecorated by many crosses and small chapels. Catalinalater muttered that people too often drove down thehill while drunk and failed to make the turns.

Visiting the Jardín was one of the highlights of ourjourney. I had no idea what to expect as I had not seenany pictures or detailed descriptions. We drove out alittle way on the main highway between Coro andCaracas. Located on the strip of land between the high-way and the Caribbean coast, the Jardín entrance wasfrom the highway through a high-walled façade withthe name Jardín Botánico Xerófito “Dr. Léon Croizat”on the front and offices behind (Fig. 3). As we enteredwe found ourselves at a central pathway and low flow-ing shrubs. Beyond, we could see sand dunes obscur-ing the beach beyond. Other paths led between dif-ferent focal areas with mesh-in enclosures and steppedterraces within. Apart from the euphorbias (Fig. 4) andcacti there were some surprises such as two large speci-mens of Welwitschia mirabilis (Fig. 5) that are nativeto the desserts of southern Africa. I had heard aboutthese plants and their intriguing evolutionary positionso I found their presence in the Jardín to be very excit-ing. I understand that these two plants were amongthe largest specimens in the New World. There wasalso the strange looking baobab tree (a group nativeto Africa, Madagascar, and northern Australia), a spe-cies I had never before other than in books.

Daily operations were supported by a small staffthat tended the plants. As I walked around the JardínI could see signs of the challenges faced by Catalinaand supporters of the gardens. Although much of thegarden was intact I could see signs of deterioration.The enclosures were in a state of disrepair and theplants needed attention. Many other plants were stillawaiting transfer from the nursery into the garden.Catalina said the company contracted to work theJardín was often not paid, and the barrier along the


center of the highway prevented regular use by visitorssuch as school groups and university students. Pottedplants were available for sale, but the money did notgo to support the Jardín. She said there were only about10 people working the Jardín when 40 were needed.

Dr. Bastida also took us on a tour of the scientificinstitutions in Coro, including the University campus,and the Centro de Investigaciones de Ecología y Zo-nas Áridas de la UNEFM. Through the support of Centerdirector Dr. Miriam Díaz I was invited to give an eveningpresentation on Croizat’s theories to a small audience.Although I was dealing with a subject that I was stillworking to understand, I had a few slides with meand I was able to give a brief outline of the principlesof biogeography, symmetry, and evolution. Eventhough I could only communicate in English, the au-dience was forgiving and the slides at least made itpossible to convey the main points. Afterwards Catalinasaid the presentation agreed very well with her under-standing of Croizat’s work. This was most reassuring asI was somewhat nervous about speaking on this topicwith such a knowledgeable person in the audience.

Our time in Coro came to conclusion all too quickly.At the very end Catalina agreed to tell me about her-self. I asked to interview her when I arrived, but shehad put it off. She now said that rather than an inter-view she would tell me her story in her own words.She said she was born in Hungary. Her father was alawyer while her mother an intellectual and philoso-pher closely attuned to the importance of generalknowledge. She attended a public school and thenconvent school where she did very well. She attendedthe Université de Paris Sorbonne, first with financialsupport from the convent and then through her aca-demic performance. She learned several languages andthen taught French and German in Hungary. She mar-ried a man from the family of her stepfather. As theSecond World War started to unfold, she and her hus-band decided that they needed to leave Europe andVenezuela seemed to be a very good place to go. Herhusband traveled to Venezuela first and she was thento follow. She had just boarded a ship sailing fromGenova (Genoa) to Venezuela when Italy entered thewar. The ship was attacked and sank. She and othersurvivors were rescued by another ship and returnedto Genova where she came down with pleurisy andbecame very ill. She was still very sick when sheboarded another ship that took her to Venezuela.

In Venezuela Catalina first worked as a cook andthen started a career in child psychology, having stud-ied psychology in Italy. In Caracas she established thefirst institute for this discipline. While returning froma visit to France, Catalina met Croizat. She describedhim as a person with a broad knowledge and interest- quite unlike other botanists who she generally foundto be dull and uninteresting. She later amicably di- Fig. 5. Welwitschia mirabilis in the Jardín (1988).

Fig. 3. Entrance to Jardín Botánico Xerófito “Dr. LéonCroizat” (1988).

Fig. 4. Euphorbia millii of Madagascar in the Jardín(1988).


Fig. 7. New plaque dedicated to the work of Léon Croizat.

Fig. 6. Dedication of new headstone for Catalina andLéon Croizat (2002) (María Elvira Gómez, rector of theUNEFM, and Mario Zunino, unveiling the plaque).

vorced from her husband as they had been separatedfor some time. She married Croizat and suggested thathe leave his university position because there was aproblem created by a demand that his articles beauthored by another person. Catalina proposed thatshe support them through the development of a land-scape gardening business. She made several trips toCalifornia to learn the practice. Having worked withvarious families in Caracas through her psychologyinstitute she now had contacts for establishing herlandscaping business. Catalina said she was initiallyreluctant to move to Coro, but she moved because ofCroizat. But now she liked Coro even though it is asmall town with limited commodities.

As we were to leave at the end of the week, Catalinaproposed coming with us to Caracas. It was to beCroizat’s birthday and she would rather be out of thetown (I had totally overlooked the significance of the

dates I was visiting, which shows that I was a bit clue-less about such things at that time). We stayed with aHungarian friend of Catalina for a couple of nightswhich gave us the opportunity to see some of Caracas(including a very nice restoration of a house belong-ing to Simón Bolívar or one of his colleagues, and avery nice bookstore where we were able to buy somenatural history books) before catching a plane back tothe US and then to New Zealand. Since that time Iwas unable to keep up regular contact with Catalina.She passed away on July 29, 1997 and is buried alongwith Croizat. Last year it was reported in the newspa-per El Nacional that the Jardín finally fell into suchneglect that numerous plant collections from all overthe world were lost, including the Welwitschia speci-mens. A recovery has been proposed. In February 2002there was an exhibition of Croizat’s works, and a pub-lic event where presentations were made by Dr. José


Vicente Scorza (Universidad de Los Andes), Dr. MarioZunino (Università di Urbino “Carlo Bo”, Italy) and Dr.Ángel Viloria (Instituto Venezolano de InvestigacionesCientíficas). A new gravesite (Fig. 6) and plaque (Fig.7) were also dedicated to Croizat with support fromthe Società Italiana de Biogeografia.

The journey to Coro gave me a unique insight andunderstanding of the intellectual and practical chal-lenges faced by Croizat in the development of his radi-cal ideas on biogeography and evolution. Looking backat what he accomplished for panbiogeography, andthe prominence that later came to his work, I am stillamazed. For the most part, his leading contemporar-ies in evolutionary theory were established within theinstitutions of science and had resources that Croizatcould only dream about. But through his extensivecorrespondence, Croizat was in surprisingly close con-tact with many of the influential biologists of his time.That Catalina was also pivotal in changing the courseof evolutionary biology through her total commitmentto Croizat’s work is something I found beyond ques-tion. For me, the impact of their commitment extendswell beyond their lifetimes in both space and time –as so it should.

Acknowledgements

I am most appreciative to Jürg De Marmels, ÁngelViloria, and Mario Zunino for feedback on the draftarticle, and also to Ángel Viloria for illustrations of thededication.

John GrehanBuffalo Museum of Science, 1020 Humboldt Park-way, Buffalo, NY 14211-1193, [email protected]


Una perspectivalatinoamericana

de la biogeografía(1a. edición), 2003,

Morrone JJ, LlorenteBousquets J, eds.

Universidad NacionalAutónoma de México,

México, 307 pp.ISBN 970-32-0498-8.

MXN $168 (USD 15.30).

Since the beginning of the 20th century, biogeogra-phy has been shaped by various “schools”. The dis-persalist “First New York”, the vicariance “Second NewYork”, and the panbiogeographic “New Zealand”schools have each had their periods of prominence,and continue to generate both light and considerableheat among historical biogeographers. With the dawnof the 21st century, a new school has appeared that isgiving a fresh perspective and direction to biogeogr-aphic research. Members of this school are from LatinAmerica, and the focus of their research is the Neo-tropics and Mexico. Rather than promoting particularmethodologies, the “Argentine-Mexican School”(named for the two countries where biogeographicworkers have been the most active) is pragmatic, di-verse, and focused on areas and biotas of the Neo-tropics that have been rather neglected by First Worldbiogeographers.

In fact, South America has had an active cadre ofnative biogeographers for almost as long as the UnitedStates. For those wanting to learn more about the“Argentine-Mexican” biogeography school, and to readabout recent developments in Neotropical biogeogra-phy, Una perspectiva latinoamericana de la biogeogra-fía is an excellent introduction (with three articles inEnglish, one in Portuguese, and 30 in Spanish, somereading knowledge of the latter is required). This vol-ume is one of a series of books focusing on Latin Ameri-can biogeography published since 2001 by the Uni-

B O O K R E V I E W

Una perspectiva latinoamericana de la biogeografía

BY R. WILLS FLOWERS

versidad Nacional Autónoma de México. All were writ-ten primarily for a Latin American audience, with thegoal of both publishing contemporary works andmaking some of the classic English biogeographicalstudies accessible to a Spanish-speaking audience.

Una perspectiva latinoamericana de la biogeografíais divided into three sections: history; theories, meth-ods and concepts; and case studies. The historical pa-pers include two essays on the works of Charles Lyelland Alfred Russel Wallace; both are intended to makethese classic authors accessible to Spanish speakersbut may be of limited interest to English speakers, whocan readily find both the original works and innumer-able English commentaries. Of greater interest are es-says on Oswaldo Reig and Raúl Ringuelet, two Argen-tine biogeographers. Reig worked with South Ameri-can mammals and Ringuelet pioneered the study ofbiogeographic subregions in the Southern Cone basedon his studies of freshwater crustaceans and leeches.Another essay evaluates the impact of Léon Croizaton Latin American biogeographers; this essay includesas appendices Spanish translations of two of Croizat’slater, hard-to-find, and relatively short publications.

The second section on theories, methods, and con-cepts covers topics from tracks to parsimony analysisof endemism to phylogeography. Most of the articlesare intended to bring Spanish speakers up to speedon recent developments in the United States and Eu-rope; thus, readers outside Latin America may find thissection of limited value since much of the methodol-ogy has already been expounded and extensively dis-cussed in English. I found one short article in Englishby Nelson and Ladiges on geographic paralogy worth-while because it actually includes a definition of geo-graphic paralogy, something that is unaccountablylacking in many English articles on the subject (includ-ing the otherwise seminal Historical biogeography[Crisci et al. 2003]). In their article on biogeographicaltransitions, Ruggiero and Ezcurra provide a revealinggraph of past climactic events, which shows thatglaciations in the Southern Hemisphere began in theEocene and, with a slight letup in the Miocene, havecontinued to the present. (Yet First World biogeogr-aphers continue to obsess over whether the much more


recent Pleistocene glaciations were what shaped thebiota of the Amazon.) In their article on phylogeogr-aphy, Lanteri and Confalonieri cite two interesting casesin the Curculionidae (Coleoptera). One study showedthat the boll weevil (Anthonomus grandis) in Argen-tina comprises two distinct populations: one identicalto pest populations in the southern United States andone ancestral population related to Mexican boll wee-vils. In the other study of the genus Galapaganus, mito-chondrial DNA shows that the age of the species onthe Galapagos Islands predates the oldest extant island.

The third section on case studies was for me themost interesting part of this volume. While vertebratesare not ignored, most of the articles featured inverte-brate or plant groups as subjects for analysis. There isa definite emphasis on the Mexican Transition Zoneand on biogeographic subregions of Argentina andsurrounding countries. Some of these studies (e.g.,Morrone’s article on trichodactyline decapods) areupdates or continuations of earlier studies publishedin English (e.g., Morrone & Lopretto 1994). Many au-thors follow the recommendations of Crisci et al. (2003)in their selections of methods of analysis. A numberof papers begin with track analysis followed by areacladograms and parsimony analysis of endemicity.Some notable contributions include Kohlmann andWilkinson’s delineation, using distributions of dungbeetles, of a biogeographic barrier in Costa Rica alongthe Río Grande de Tárcoles at the foot of the Talamancamountain range. Interestingly, Croizat (1976) repeat-edly cites nearby Punta Mala as a major biogeographicnode. As an entomologist, I found the articles on tropi-cal Andean Satyridae (Lepidoptera) by Viloria, a world-wide review of scorpion biogeography by Lourenço, areview of Neotropical Muscidae (Diptera) and a biogeo-graphic study of the arthropods of the precordilleraof Argentina by Roig-Juñent, Flores, and Mattoni ofparticular interest. All except the last are in English.

My most serious complaint about this book is itssmall type size and sans serif font, which makes read-ing difficult except in very well-lit conditions. Some ofthe distribution maps would be more legible in a largersize, but in some cases this is a result of reproductionof small originals. However, for anyone with an inter-mediate reading knowledge of Spanish (and a goodreading glass), this volume offers a wealth ofbiogeographic information from a mega-diverse partof the world.

References

Crisci JV, Katinas L, Posadas P. 2003. Historical biogeography: Anintroduction. Cambridge: Harvard University Press.Croizat L. 1976. Biogeografía analítica y sintética (‘panbiogeogra-fía’) de las Américas. Caracas: Biblioteca de la Academia de Cien-

cias Físicas, Matemáticas y Naturales.Morrone JJ, Lopretto EC. 1994. Distributional patterns of fresh-water Decapoda (Crustacea: Malacostraca) in southern SouthAmerica: A panbiogeographical approach. Journal of Biogeogra-phy 21: 97-109.

R. Wills FlowersCenter for Biological Control, Florida A&M Univer-sity, Tallahassee, FL 32307, [email protected]



Amazonian relationships:An example of a complex area and a complex problem

BY IVONNE J. GARZÓN-ORDUÑA AND DANIEL RAFAEL MIRANDA-ESQUIVEL

Amazonia is a rich and diverse ecosystem. It comprisesthe Amazon basin and the lowlands of southern Co-lombia, southern Venezuela, eastern Ecuador, north-ern and eastern Peru, eastern Bolivia, northern Brazil,and the Guiana. Different approaches (Cabrera & Willink1973, Müller 1973) have been used to identify its lim-its. However, the concern for its biogeographic historyis recent (Amorim 2001). Just a few years ago the firstquantitative approaches using phylogenetic hypotheseswere generated. Cracraft & Prum’s (1988) study gavethe first steps to resolve the question about Amazonia’sorigin, but Amorim’s (2001) work was the first to chal-lenge the traditional view of Amazonia as a single unit(Cabrera & Willink 1973, Hooghiemstra 1997).

The earliest attempts to reconstruct the Amazo-nian biogeographic history were non-quantitative ap-proaches based on narrative descriptions of thoseevents that contributed to today’s diversity of species.These are known as the Amazonian diversification hy-potheses (Wallace 1852, Haffer 1969, for a revisionsee Haffer 1997 and Nores 2000). Although these ap-proximations were very useful at that time becausethey opened the question about Amazonian diversity,since the development of quantitative biogeographyapproaches they have been severely criticized for twomain reasons. First, because they are based on explicitor implicit a priori assumptions (Amorim 2001), andsecondly because their statements could not be em-pirically tested (Cracraft & Prum 1988, Patton & da Sil-va 1998, Alexio 2002). Under an explicit biogeographicframework the first cladistic and quantitative analyzesbegan with Prum (1988) and Cracraft & Prum (1988).Although Cracraft & Prum (1988) did not explicitlydescribe the methodology used to obtain their areacladogram, Brooks Parsimony Analysis (BPA) was themost frequent method used. After them, da Silva &Oren (1996), Bates et al. (1998), Ron (2000), and Ra-cheli & Racheli (2003, 2004) used a modification ofPAE (Cracraft, 1991) to generate their hypotheses ofarea relationships. Amorim (2001) and Hall & Harvey(2002) have proposed the latest and most recent arearelationship hypotheses using phylogenetic and dis-

tributional data, using “cladistic biogeography” andBPA, respectively.

In spite of all these efforts, the origin and thebiogeographic history of the Amazonian diversity isstill in discussion. This is evident from the amazingnumber of papers dealing with the subject (Peres &Terborgh 1995, Haffer 1997, Hooghiemstra 1997, Ma-rroig & Cerqueira 1997, Colinvaux et al. 2000, Bates &Demos 2001, Rose & Grainger 2003). From thebiogeographic framework, however, these area rela-tionship hypotheses have failed to document explicithistorical events, like those events resulting fromcladogenetic events, since seven of the 16 area clad-ograms were developed without phylogenetic infor-mation of the Amazonian taxa (Table 1). For example,Racheli (2004) has criticized the use of the Matrix Par-simony Representation technique used by Hall & Harvey(2002) to produce their combined area cladogram,questioning the use of the competing topologies assource data. Additionally most of these hypotheseshave been formulated from a single taxonomic group.It is obvious that the history of the area had to haveaffected all the Amazonian taxa that coexisted overthe region, producing their speciation, extinction, ordispersion pattern(s).

The diversification hypotheses:The non-quantitative approaches

There have been proposed more than 16 diversifica-tion hypotheses (Nores 2000), these hypotheses for-mulated a causation perspective for the Amazoniandiversity - lacking any approach to quantitative analy-sis and therefore these hypotheses have had a strongnarrative tradition. Nores (2000) presented an exten-sive review of these hypotheses. One of the first tohighlight the diversity present in the Amazonian for-ests was Alfred Wallace (1852), after his voyage alongthe Amazonian Basin. His observations lead him topropose the first diversification hypothesis to theAmazonia, known now as “The Riverine Hypothesis”.


Table 1. Main hypotheses of area relationship proposed for Amazonia. For each pattern the author’s name, taxa used, method andexplanation suggested are shown. Be, Belem; BPA, Brooks parsimony analysis; CAT, Caatinga; CB, “cladistic biogeography”, no methodspecified; GU, Guiana; IM, Imeri; IN, Inambari; IV, implicit vicariance; MRP, matrix parsimony representation; NP, Napo; PA, Para; PAE,parsimony analysis of endemicity; PH, phylogeography; RO, Rondonia; SD, Serra do Mar.

Author Taxa Method Area cladogram Process invoked

Prum (1988) Birds BPA ((GU,(PA, BE)),(RO, IM, IN, NP)) IV*Prum (1988) Birds BPA (GU,((RO,(BE,PA)),(IM,(IN,NP)))) IV*Cracraft & Prum (1988) Birds BPA (SD,((CH,CA),(GU,((SD,BE_PA),(IM,(IN,NP)))))) Vicarianceda Silva & Oren (1996) Primates PAE (GU, (PA, BE)), ((RO, IN1), (IM, IN2, NP)) VicarianceBates et al. (1998) Birds PAE (GU, (IM, (IN, NP)), (BE, (PA1, (PA2, (RO)))) IV*Patton et al. (2000) Rodents and marsupials BPA (GU, (IM, (IN, NP))), (RO, PA, BE) VicarianceRon (2000) Squamata PAE ((PA,(RO,(BE,GU))),(IN,NP)) VicarianceRon (2000) Anura PAE (BE,(RO,(GU,(IN,NP)))) VicarianceAmorin (2001) Insects and primates CB (((IM,(GU,BE)),(NP,IN)),(PA,(SD,CAT))) VicarianceHall & Harvey (2002) Butterflies PH (GU,((RO,(BE,PA)),(IM,(IN,NP)))) VicarianceHall & Harvey (2002) Vertebrates and butterflies MRP ((RO,(BE,PA)),(GU,(IM,(IN,NP)))) VicarianceMarks (2002) Birds PH ((GU,(IN,(NP,(CH,(CA,IM))))),(RO,(PA,SD))) none explicit**Marks (2002) Birds PH (RO,((GU,(IN,(NP,(CH,(CA,IM))))),(PA,SD))) none explicit**Racheli & Racheli (2003) Birds PAE ((IM,(NP,IN)),(GU,((BE,PA2),(PA1,RO)))) IV*Racheli & Racheli (2004) Butterflies PAE ((GU,(BE,(PA2,(RO,PA1)))),(IM,(IN,NP))) IV*Racheli & Racheli (2004) Butterflies PAE (GU,((BE,(PA2,(RO,PA1))),(IM,(IN,NP)))) IV*

* We assume implicit vicariance since the hypothesis was produced using pattern methods that recognize only vicariance events.** Marks (2002) found low genetic divergence between Imeri and Central America populations product basically of dispersal, but thereis no explicit reference about the process involved.

The principal argument is that the current biodiversityis the result of the vicariant events depicted from theformation of the Amazon River and its tributaries. Allof the diversification hypotheses have presented avicariance perspective of the Amazonian history; clearlythe speciation is given by allopatric events caused bya variety of barriers to gene flow: rivers, lagoons, eco-logical gradients, or the sea level. Marroig & Cerqueira(1997) offered some of the phylogenetic and biologi-cal implications of the statements proposed by themost important diversification hypotheses: thePleistocene refuge hypothesis (Haffer 1969), theriverine hypothesis (Wallace 1852, Sick 1967), and thegradient hypothesis (Endler 1977).

The quantitative approaches

Quantitative analyses gained ground during the lastfew years since the impact and acceptance of thephylogeography research program (Lougheed et al.1999, Ditchfield 2000, Patton et al. 2000, Marks et al.2002, Costa 2003). Population genetics analyses, un-doubtedly, have contributed to the “Amazonian cause”,for example, testing some predictions of the riverine di-versification hypothesis (Patton & da Silva 1998,Lougheed et al. 1999, Alexio 2002). Nevertheless, most

of these analyses have been developed to answer thespeciation history of one or two particular taxonomicgroups and not to explore the relationship betweenthe areas.

Under an explicit biogeography framework, thearea relationship for the Amazonia has been recon-structed eleven times (Table 1), but since some authorspresented more than one, there are 16 proposed areacladograms. All these biogeographic studies stated thatthe history of Amazonia was driven mainly by vicar-iance events that have produced a hierarchical pat-tern (Moritz et al. 2000, Ron 2000, Amorim 2001, Bates2001, Hall & Harvey 2002). Ron (2000) and Hall &Harvey (2002), based on the congruence between thehistory of the areas for taxonomically distant groups(butterflies, primates, birds, anurans, and lizards), pre-sented a strong common history of vicariant isolationevents proposing as the first event the disjunctionbetween Guiana and the Atlantic Brazilian Forest. Thesestudies also have shown some persistent relationships,for example a consistent Inambari-Napo group. TheGuiana position, however, continues to be uncertain,its relationship with the so-called upper (Imeri andNapo) and lower (Inambari, Rondonia, Para, andBelem) Amazonia is inconsistent among the hypoth-eses and sometimes appears to be an independentunit. Additionally, it is still unclear what barriers were


responsible for the proposed vicariant patterns (Ron2000, Hall & Harvey 2002) and the timing of the diver-sification. The controversy continues at this point, andseveral hypotheses have been proposed to explain thatvicariant pattern (the diversification hypothesis) andalthough the Pleistocene Refuges model (Haffer 1969)has generated discussion, this is still accepted for somegroups of animals and within the neontologists, toexplain the diversification patterns (Brower 1994, Hall& Harvey 2002). One major problem that involves allof the above mentioned hypotheses of area relation-ship is that they were formulated to account for eventsof the late Pleistocene period, leaving out earlier peri-ods (Cretaceous or late Tertiary) which molecularphylogenies of frogs, lizards, birds, and mammals sug-gest to be a critical time for the main events ofspeciation (Moritz et al. 2000, Ron 2000).

The most important contributions about the ori-gin of the Amazonian biota probably have come fromrecent phylogeographic studies (Patton et al. 2000,Marks et al. 2002, Costa 2003). Marks et al. (2002),using the diversification pattern of Glyphorynchus spi-rurus, showed that some Amazonian haplotypes (Imeripopulation) are more closely related to Central Americaand Chocó populations than they are to other Amazo-nian taxa (Napo populations), a pattern that they docu-mented to be common within the Amazonian birds(Marks et al. 2002, p. 164). Costa’s (2003) analysis ofsmall mammals in Amazonia and the Brazilian Atlan-tic forest showed that these two regions are not ex-clusive in their fauna composition, and that the ge-netic similarity exhibited between the mammals in thetwo areas is often higher than the similarity withineach of the areas. Although others works had alreadyachieved a similar conclusion (Cracraft & Prum 1988,Amorim 2001), it is interesting to see how this resultrepeats when new taxa are studied, and how wrongwas the idea of two different units (provinces or do-mains) that now appear to be a single biogeographicunit. Another conclusion produced by Costa’s (2003)work, which must be tested with strictly historicalbiogeographic approaches, predicts that a general areacladogram for the history of the Neotropical area isnot likely and that a single vicariance model will notexplain the speciation events. Although these worksare focused on birds and mammals, they have pro-duced new insights about the origin of the Amazo-nian biota and have placed new challenges to the bio-geography research program. Given this scenario if wewant to understand the biotic diversity of theAmazonia, and the reasons to explain why it supportsthat diversity we have to search for a reconstructionof the Amazonian history using all the distributionaland phylogenetic information available, using an ex-plicit cladistic method to account all the possible eventsthat have taken place in Amazonia.

Acknowledgments

We are deeply thankful to Isabel Sanmartín, LeonoraCosta, Juan J. Morrone, Jim Patton, Paula Posadas,Carla Penz, André Freitas, Keith Brown Jr., SalvadorArias, Mario Quijano, and Claude Gascon for their con-structive comments. The first author is much indebtedto Isabel Sanmartín for her constant incentive andencouragement. All the members of the Laboratoriode Sistemática & Biogeografía are acknowledged fortheir individual efforts. Malte Ebach helped us as Edi-tor. The financial support of the Vicerectoría deInvestigaciones y extensión, UIS, is kindly acknowl-edged.

References

Aleixo A. 2002. Molecular systematics and the role of the “Varzea”-“Terra-Firme” ecotone in the diversification of Xiphorhynchuswoodcreepers Aves: Dendrocolaptidae. The Auk 119: 621-640.Amorim DS. 2001. Dos Amazonias. Introducción a la biogeografíaen Latinoamérica: Teorías, conceptos y aplicaciones. Llorente -Bousquets J, Morrone JJ eds. Pp. 245-255. Facultad de Ciencias,UNAM, Mexico, D.F.Bates JM. 2001. Avian diversification in Amazonia: Evidence forhistorical complexity and a vicariance model from a basic patternof diversification. Diversidade Biológica e cultural da Amazonia.Viera I, D’Incao MA, Cardoso da Silva JM, Oren D eds. Pp. 119-138.Museu Paraense Emilio Goeldi, Belém, Pará, Brazil.Bates JM, Hackett SJ, Cracraft J. 1998. Area-relationships in theneotropical lowlands: An hypothesis based on raw distributions ofPasserine birds. Journal of Biogeography 25: 783–793.Bates JM, Demos T. 2001. Do we need to devalue Amazonia andother large tropical forest? Diversity and Distributions 7: 249-255.Brower AVZ. 1994. Rapid morphological radiation and convergenceamong races of the butterfly Heliconius erato inferred from pat-terns of mitochondrial DNA evolution. Proceedings of the NationalAcademy of Sciences 91: 6491-6495.Cabrera AL, Willink A. 1973. Biogeografía de América Latina.Monografías de la O.E.A. Serie Biología. Nro 13. Washington, D.C.Colinvaux PA, de Oliveira PE, Bush MB. 2000. Amazonian andNeotropical plant communities on glacial time-scales: The failureof the aridity and refuge hypotheses. Quaternary Science Reviews19: 141-169.Costa, LP. 2003. The historical bridge between the Amazon andthe Atlantic forest of Brazil: A study of molecular phylogeographywith small mammals. Journal of Biogeography 30: 71-86.Cracraft J. 1991. Patterns of diversification within continentalbiotas: Hierarchical congruence among the areas of endemism ofAustralian vertebrates. Australian Systematic Botany 4: 211-227.Cracraft J, Prum RO. 1988. Patterns and processes of diversifica-tion: Speciation and historical congruence in some Neotropicalbirds. Evolution 42: 603-620.Ditchfield AD. 2000. The comparative phylogeography of Neotro-pical mammals: Patterns of intraspecific mitochondrial DNA varia-


tion among bats contrasted to nonvolant small mammals. Mo-lecular Ecology 9: 1307-1318.Endler JA. 1977. Geographic variation, speciation and clines.Princeton University Press. Princeton.Haffer J. 1969. Speciation in Amazonian forest birds. Science 165:131–137.Haffer J. 1997. Alternative models of vertebrate speciation inAmazonia: An overview. Biodiversity and Conservation 6: 451-476.Hall JPG, Harvey DJ. 2002. The phylogeography of Amazonia re-visited: New evidence form Riodinid butterflies. Evolution. 56: 1489-1497.Hooghiemstra H. 1997. Tropical rain forest versus savanna: Twosides of a precious medal? A comment. The Hague, pp. 31-43. TheNetherlands Organization for Scientific Research.Lougheed SC, Gascon C, Jones DA, Bogart JP, Boag PT. 1999.Ridges and rivers: A test of competing hypotheses of Amazoniandiversification using a dart-poison frog Epidobates femoralis. Pro-ceedings of the Royal Society London 266: 829-1835.Marks BD, Hackett SJ, Caparella AP. 2002. Historical relationshipamong Neotropical lowland forest areas of endemism as deter-mined by mitochondrial DNA sequence variation within the Wedge-billed Woodcreeper Aves: Dendrocolaptidae: Glyphorynchusspirurus. Molecular Phylogenetics and Evolution 24: 153-167.Marroig G, Cerqueira R. 1997. Plio-pleistocene south Americanhistory and the Amazon lagoon hypothesis: A piece in the puzzleof Amazonian diversification. Journal of Comparative Biology 2:103-119.Moritz C, Patton JL, Schneider CJ, Smith TB. 2000 Diversificationof rainforest faunas: An integrated molecular approach. AnnualReview of Ecology and Systematics 31: 533-563.Müller P. 1973. The dispersal centers of vertebrates in theNeotropical realm. Biogeographica 2: 10-198.Nores M. 2000. Species richness in the Amazonian bird fauna froman evolutionary perspective. Royal Australian Ornithologists Union100: 419-430.Patton JL, da Silva MNF. 1998. Rivers, refuges and ridges: Thegeography of speciation of Amazonian animals. Endless form:Modes and mechanisms of speciation. Howard D, Berlocher S eds.Pp. 202-213. Oxford University Press, Oxford.Patton JL, da Silva MNF, Malcolm JR. 2000. Mammals of the RioJuruá and the evolutionary and ecological diversification ofAmazonia. Bulletin of the American Museum of Natural History244: 1-306.Peres CA, Terborgh JW. 1995. Amazonian nature reserves: An analy-sis of the defensibility status of existing conservation units anddesign criteria for the future. Conservation Biology 9: 34-46.Prum RO. 1988. Historical relationships among avian forest areasof endemism in the Neotropics. Acta XIX Congressus InternationalisOrnithologici. Pp. 2563–2572. Ouellet H de. XIX Internat. Ottawa.Racheli L. 2004 The nigthmare of the combination: Comments onmatrix representation with parsimony and its first application inbiogeography. Cladistics 20: 208-211.Racheli L, Racheli T. 2003. Historical relationship of Amazonianareas of endemism based on raw distributions of parrots Psittacidae.Tropical Zoology 16: 33-46.Racheli L, Racheli T. 2004. Patterns of Amazonian area relation-

ships based on raw distributions of papilionid butterflies Lepidop-tera: Papilionidae. Biological Journal of the Linnean Society 82:345-357.Ron S. 2000. Biogeographic area relationships of lowland Neo-tropical rainforest based on raw distributions of vertebrate groups.Biological Journal of the Linnean Society 71: 379-402.Rose S, Grainger A. 2003. Multivariate mapping of spatial varia-tion in biodiversity in Peruvian Amazonia. Diversity and Distribu-tions 9: 237-249.Sick H. 1967. Rios e enchentes na Amazonica como obstaculo paraa avifauna. Atlas do Simposio sobre a biota Amazonica. Lent H ed.Pp. 495-520. Conselho Nacional de Pesquisas. Rio de Janeiro.da Silva JMC, Oren DC. 1996. Application of parsimony analysisof endemicity in Amazonian biogeography: an example with pri-mates. Biological Journal of the Linnean Society 39: 427–437.Wallace AR. 1852. On the monkeys of the Amazon. Proceedingsof the Zoological Society of London 20: 107-110.

Ivonne J. Garzón-OrduñaDepartment of Biological Sciences, University ofNew Orleans, 2000 Lakeshore drive, New Orleans,LA, 70148, USA

Daniel Rafael Miranda-EsquivelLaboratorio de Sistemática & Biogeografía, Escuelade Biología, Universidad Industrial de Santander,A. A. 678 Bucaramanga, Colombia.


B O O K R E V I E W

Ghosts of Gondwana

BY JOHN R. GREHAN

Ghosts of Gondwana,2006, Gibbs GW. Craig

Potton Publishing,Nelson, New Zealand.

232 pp. ISBN 13: 978-1-

877333-48-4.NZD 49.99 (USD 34.50).

George Gibbs faced no small challenge in writingGhosts of Gondwana. New Zealand has long been amajor center of attention for global biogeographicstudies, and its origins and evolution were subject tointense debate over the last quarter century. Ghostsof Gondwana attempts to portray the origins and evo-lution of organisms in the context of New Zealand’slinear temporal sequence, so that different groups arediscussed in each chapter or section according to theirorigin before or at various times following New Zea-land’s geological isolation according to fossil, molecu-lar, or dispersal criteria. In Ghosts of Gondwana Gibbsattempts to grapple with new biogeographic researchpresent a balanced view on the historical argumentsover the origins and evolution of the fauna and flora.As noted in the introduction, this is not just anotherbook describing the plants and animals of New Zea-land, but a selection of examples that best illustratethe unique features of New Zealand life by drawingon biological and geological information to trace theirhistory. In many respects Ghosts of Gondwana is verysuccessful in its exploration of a broad range of issuesand examples in a coherent and sequential picture oftraditional theories about New Zealand’s biogeo-graphic origins and evolution and some of the centralissues of contention. The issues are arranged in fivesections. Section I emphasizes New Zealand’s ende-mism, section II looks at the historical issues for bio-geography, section III looks at various explanationsfor New Zealand’s life, section IV examines specific

groups or ecologies within New Zealand, and sectionV provides some final thoughts on the problem of dis-tinguishing vicariance and dispersal. Each of the ma-jor issues and prominent examples are complementedby extensive illustrations, beginning with a geologicaltimescale that includes an overview of dominant life-forms and some major historical geological events, andfollowed by various focal inserts summarizing the per-tinent geographic, systematic and biological featuresor individual organisms or various groups.

Ghosts of Gondwana represents a significant sign-post in the history of biogeographic and evolutionaryresearch in New Zealand. During the mid 20th centuryan almost universally accepted history of the New Zea-land biota was established by Sir Charles Fleming, whoused geological stratigraphy to construct a center oforigin and dispersal model that divided organisms intoancient endemics (later to become Gondwana ele-ments) and later colonists from various centers of ori-gin or derived from New Zealand itself as a center oforigin within the region. Only with rare exceptions,such as the pioneering New Zealand palynologist LucyCranwell (1962, 1964), was there any early willing-ness to consider geological influences in the evolutionof New Zealand plants and animals. The scientific ex-ploration of New Zealand’s evolution is characterizedin Ghosts of Gondwana as a search for criteria to dis-tinguish New Zealand’s original biota before separa-tion from the hypothetical super-continent of Gond-wana, and those that subsequently dispersed to orfrom New Zealand after its geological and geographicisolation about 80 million yeas ago.

The division of New Zealand’s biota into ancientendemics and later dispersal was established as thefoundation for New Zealand’s evolutionary history bySir Charles Fleming. With popular acceptance of con-tinental drift and plate tectonics the ancient endemicsbecame the icons or New Zealand’s original Gondwanainheritance. But all the elements of the biota contin-ued to be seen as the product of dispersal either be-fore or after isolation and arriving from various centersof origin in the local region or other parts of the globe.Life was seen as being continually on the move as vari-ous newly evolved groups moved along one or other


dispersal pathway to sometimes finally even end evenup at the most isolated locations such as New Zea-land. It was not until the 1980’s that Fleming’s evolu-tionary model was challenged by local support for LéonCroizat’s panbiogeography. It would be an understate-ment to say that this was received with strong oppo-sition from the scientific establishment. George Gibbsfound himself in the middle of the debate through hisrole as the graduate advisor for Robin Craw, Ian Hen-derson, and John Grehan, who were finding panbio-geography to provide a much more scientifically de-fensible approach to New Zealand’s biological evolu-tion. While taking a critically cautious path in the faceof some very radical propositions about New Zealandhistory, George Gibbs also found the opportunity toapply panbiogeographic methods to his interest in theprimitive micropterigid moths, and even explore thepossibility their biogeographic origin was linked to aPacific geo-history (Gibbs, 1983) and that their NewZealand distributions may reflect patterns dating backto the Cretaceous (Gibbs, 1990). Following a very suc-cessful symposium on panbiogeographic and non-pan-biogeographic perspectives in 1989 (Matthews 1990),the panbiogeographic synthesis failed to receive thenecessary support from New Zealand’s research andfunding institutions to continue as an active researchprogram and there has now been no endemic panbio-geographic research program for many years.

Through Ghosts of Gondwana, one is in a positionto see how various animal and plant groups are inter-preted through theoretical interpretations of fossil andmolecular evidence. The principle deficiency of thisapproach lies in what is left out. The panbiogeographicanalyses of New Zealand are presented as a largelyhistorical event as Ghosts of Gondwana looks to mo-lecular systematics as the fuel for the “latest revolu-tion in biogeographic thinking” that can provide a di-rect measure of time in biogeography, whereas Croi-zat’s panbiogeography could only analyze space. Thedevelopment of routine methods of sequencing theDNA molecule which “contains information on theevolutionary history of every living organism” is char-acterized as the most exciting advance in recent years,and molecular technology is presented as a biogeo-graphical revolution generating “new” interpretationsof how and where the present fauna and flora reachedNew Zealand. In one sense these characterizations aretrue, but constructing a phylogenetic tree requiresappropriate concepts of character homology to sepa-rate primitive and derived character states involvingonly four base combinations that provide no trace ofindividual replacements at each site. Ghosts of Gond-wana does not show any empirical evidence for mo-lecular similarity being necessarily any more preciseabout phylogenetic relationships than morphology soit is not possible to discern whether the molecular

emphasis and historical relegation of panbiogeographyis scientifically justified.

Throughout Ghosts of Gondwana the molecular ap-proach is used to generate divergence estimates fortaxa where fossils are not available. Even though thefirst fossil appearances are acknowledged to be po-tentially be unreliable and that molecular biology onlygives “approximate hypothetical answers” these con-cerns are overruled by “a cascade of studies that offervery persuasive arguments for efficacy for trans-oce-anic dispersal”. Molecular claims for over-water dis-persal when divergence estimates post-date earliergeological events are accepted at face value, eventhough molecular divergences are in reality minimalestimates calibrated by fossils representing minimalages of fossilization. It is noted that a discrepancy of10 million years either way may not matter much, butif the discrepancy between age of origin and age offossilization may be many times greater, moleculardivergence cannot be assumed to falsify earlier geo-logical events (Heads 2005). A few examples will serveto illustrate the perspective presented in Ghosts ofGondwana and illustrate the methodological and con-ceptual boundaries of what is in many respects themost comprehensive popular book on New Zealand’sorigin and evolution in thirty years.

Chapter II presents recent DNA research to sug-gest ancestors of the short-tailed bat, Mystacina tu-berculata, were separated from ancestors of noctilio-noid bats in South America about 54 Ma, after whichthey became widespread across Antarctica and Aus-tralia before they “probably reached New Zealand byflying across the Tasman sea”. The empirical evidencefor this South American center of origin and the dis-persal story involving Antarctica and Australia is notidentified. The molecular center of origin story ignoresthe biogeographic pattern that connects New Zealandand Australia (through fossil representatives) directlywith northern South America and Central America di-rectly over the Pacific (Fig. 1). This is a pattern foundin many other animal and plant groups that do notconform to the expectations of a Gondwanic origin,but predicts a Pacific geohistory responsible for thistype of biogeographic pattern as well as the origin ofallocthonous terranes around the Pacific margins (Crawet al. 1999).

The origin of southern beeches (Nothofagus) inNew Zealand through geological isolation is describedas a theory long supported by the strong belief thatthe trees are incapable of long-distance dispersal.“Modern” molecular studies (Knapp et al. 2005) aresaid to now suggest that probably all of New Zealand’smodern Nothofagus species arrived from Australia, andreplaced the previous species within the last 35 mil-lion years. Knapp et al. (2005) claim to have “unequivo-cal molecular clock evidence” that trans-Tasman Sea


distributions can only be explained by long-distancedispersal and that the absence of Lophozonia andFuscospora pollen types in the Cretaceous of New Zea-land is evidence for Tertiary dispersals to New Zealand.The dates were calibrated by fossil pollen divergenceof for subgenera Lophozonia, Nothofagus, and Fus-cospora at 75 million years ago along with a “mini-mum” divergence between N. cunninghammii and N.moorei based on “intermediate” fossils and others that“closely resembled” N. moorei. There is no indicationof why the 75 million year fossil record should repre-sent the upper limits of this group. A calibration at 65million years for sister taxa between New Zealand andAustralia produced “unrealistic age estimates for thebasal nodes”, which might actually suggest that themolecular clock model is itself unrealistic. This prob-lem aside, the dispersal speculations would have beenenhanced by reference to the vicariant main massingsof the four Nothofagus subgenera (Heads 2006). Thesepatterns (Fig. 2) are concordant with the distributionsof each subgenus having their origins in a process ofvicariant differentiation well into the Mesozoic, andthe patterns may well represent an independent ‘test’of the molecular clock theories.

Dispersal directionality is also a common featureof molecular biogeography. For example, the basal po-sition of the New Zealand kauri (Agathis astralis) isseen as evidence that its relatives dispersed out of NewZealand. Similarly, Metrosideros has been “shown” byWright et al. (2000) to have dispersed out of New Zea-land to reach a number of Pacific islands as far awayas Hawaii. In neither case is this dispersal demon-strated. Rather, each represents an application ofHennig’s progression rule that assumes derived taxarepresent more recent dispersal events so the distri-bution of primitive and derived groups tracks the pathof dispersal. This purely theoretical Darwinian modelof evolution ignores the possibility that the same pat-

Fig. 1. Biogeography of the New Zealand short-tailedbat and its fossil relative in Australia as red polygons,and their probably nearest relatives the South Ameri-can Mormoopidae and Noctilionidae (blue) (Kennedyet al. 1999, Koopman 1984).

Fig. 2. Biogeography of Nothofagus main massings.Large text – extant species, small text – fossil pollen-leaves-cupules (from Heads 2006).

tern may arise from sequential differentiation of awidespread ancestor. This is particularly pertinent forMetrosideros, which comprises three largely vicariant(exception of a node at Fiji) taxa involving New Zea-land and the Pacific (Fig. 3). If these tracks are theresult of three serendipitous chance dispersal events,one is confronted with explaining just how each man-age not only to jump across the ocean, but to do so insuch a way as to end up on only a few islands (andnot other islands or mainland areas) without overlap-ping each other than one case in Fiji, and also track


the same path found in other organisms such as theTahiti-Hawaii connection that is also found in somePacific shorefishes (Heads 1983) or find their limits onBonin Island – another classic biogeographic bound-ary in the Pacific. Gibbs’ suggestion of a very recentorigin for one lineage scarcely varies by more than onenucleotide may be true or it shows that for this line-age there has not been much in the way of moleculardifferentiation (i.e. effectively this means the ‘clock’slowed down or is in stasis).

The treatment of Hebe biogeography provides aglobal example of the sometimes too limited scope ofcomparison between different current biogeographicframeworks and their bearing on the origin of the NewZealand biota in Ghosts of Gondwana. A molecularphylogeny by Wagstaff et al. (2002) is said to showthat the Hebe ancestor reached New Zealand about10 million years ago. Their molecular phylogeny is prob-lematic in that it does not always correspond to pat-terns of relationship derived from morphological analy-sis, yet they presume that the molecular results aresomehow more correct. They note that the earliestfossil appearance is Pliocene for Hebe, mid-Miocenefor Scrophulariaceae, and mid-Eocene for Lamiales, andadmit that their divergence estimates are “crude” andthat older fossils could be found in the future. Butthey then claim that it would be inconsistent with thecurrent fossil record to assume a Gondwana or earlierdivergence even though the earliest divergence of Hebeis not constrained by the current fossil record otherthan a pre-existing fiat on the part of the authors. Gibbscites the study by Wagstaff et al. (2002) as evidencefor dispersal away from a New Zealand center of ori-

gin, but again this dispersal is read into the relation-ships based on Hennig’s progression rule – a purelytheoretical construction based on one of the manyconflicting theoretical criteria for defining the theo-retical center of origin. Whatever position one mighttake on the molecular study, the Hebe example wouldhave been greatly enhanced by referencing and illus-trating the study by Heads (1994) showing the nodalposition of New Zealand with respect the four sub-genera (Fig. 4), the similarity of Alpine fault discon-nections shared between Hebe, Leonohebe, Nothofa-gus, and Coprosma as a tectonic correlation poten-tially falsifying 10 million year molecular divergenceas an underestimate.

Spatial analysis, acknowledged to be so importantin Croizat’s panbiogeography, represents the majorabsent methodology in Ghosts of Gondwana, wherespatial correlation is briefly illustrated for the alpinefault translocation. Fossils from the mid-Miocene LakeManuherikia of inland Otago are cited as a evidencefor reconstructing the forest communities that grewaround its shores. This example could have been greatlyenhanced by reference to the living ring species for-mations in Otago that are spatially correlated with theshores of this progressively declining lake and illus-trating the integrated biogeography of fossil and liv-ing members of the New Zealand biota (Heads 1990).Further geological correlations may have enhanced thebook by illustrating the extensive biogeographic arcsand their correlation with zones of tectonic activitysuch as plate and terrane margins, fracture zones, andbelts of granitic intrusion as well as the role of uplifttransforming mid-Tertiary lowland-coastal communi-ties (Craw 1989, Heads 1990).

It is somewhat surprising to see that biogeographicmaps are noticeably absent from a book with bioge-ography as its central theme. The global context ofNew Zealand is represented by several geohistoricalmaps, three geographic maps, and one showing thedistribution of tuataras and their fossil relatives. WithinNew Zealand there are two maps illustrating the al-

Fig. 3. Vicariant ranges of three Metrosideros lineageswithin the subgenus Metrosideros (lineages 1, 2, and3 + M. robusta/M.bartletti from Wright et al. (2000,Fig. 1).

Fig. 4. Biogeography of Hebe complex: blue – Hebe,yellow – Chionohebe, red – Leonohebe, crimson –Parahebe (from Heads 1994).


pine fault and the southern North Island disjunctions,and beech. In place of comparative biogeographicmaps, Ghosts of Gondwana looks to theories from his-torical geology, ecology, and phylogeny to obtain an-swers on the origins of New Zealand’s terrestrial plantsand animals. Since New Zealand biogeography hasbeen so intensively studied by both Darwinian and non-Darwinian research programs, the inclusion of biogeo-graphic maps could help the reader evaluate compet-ing theories about the origin of New Zealand’s plantsand animals. For example, the map illustrating the lo-cation of fossil tuatara relatives (Sphenodontia), in allcontinents except Antarctica does not show that theclosest relatives appear to be found only in the Jurassicand early Cretaceous of Mexico and the western UnitedStates (Fig. 5). This is a pattern again clearly incongru-ent with the Wegenerian model of Gondwana requir-ing a permanent Pacific Ocean.

According to Ghosts of Gondwana, the apparentinability of the giant carnivorous land-snails (Rhytidae)to travel over oceans along with their presence on partsof Gondwana (Africa, Madagascar, eastern Australia)suggest they could represent a Gondwana founder inNew Zealand. This is contrasted with their presencealso on “oceanic islands” such as the Seychelles, Indo-nesia and Melanesia and this is said to imply “a capac-ity for trans-oceanic dispersal” (Fig. 6). The alternativepossibility, that such “oceanic” islands are either notoceanic in origin, or they inherited a non-oceanic com-ponent from former non-oceanic landscapes is notconsidered. These possibilities, demonstrated for theGalapagos and Fiji (Grehan 2000, Heads 2006), sug-gest the label ‘oceanic’ is not necessarily informativefor biogeographic analysis. The real question, perhapsnever really addressed in this book, is why shouldGondwanic landsnails, distributed over former partsof Gondwana as have a completely different biogeo-graphic orientation (Indian Ocean) from the Pacific ori-ented tuatara, frogs and southern beeches that arealso distributed over former parts of Gondwana andsupposedly Gondwanan in origin? This kind of ques-tion is extensively addressed in panbiogeographic ap-proaches to New Zealand, so their omission from thisbook is problematic for achieving a balanced overviewof its biogeography and evolution.

A biogeographic correlation between New Zea-land’s biota and its geological history is addressed byGhosts of Gondwana in general reference to Wegener’smodel of Gondwana as a unified geological entity. Theexistence of accreted Pacific terranes is acknowledgedalong with the possibility of former Pacific land, buttheir biogeographic significance is discounted by the“view of tectonic geologists” that such land wouldhave been “insignificant, perhaps in the form of is-land arcs” and larger (how much larger?) terrestrialmodels are ridiculed by “mainstream geologists”. These

opinions raise the very important question of whethersuch island arcs (or any other mobile units such asmicro-continents), however small, were also necessar-ily biogeographically insignificant. Ghosts of Gond-wana presents a very different view on the role of ge-ology in determining biogeographic interpretations tothat of Gibbs (1983) where it was the evidence of bio-logical distributions represented by the Micropterigidaethat were seen to suggest that both New Zealand andNew Caledonia are composite areas comprising ele-ments from the Pacific as well as from Australia (Gond-wana). Gibbs (1983) noted that the New Zealandmicropterigids cannot be interpreted on the assump-tion that the present land mass is nothing more thana fragment of eastern Australia, yet in Ghosts of Gond-wana that is what New Zealand has largely become interms of its biogeographic evolution. In view of theextensive and diverse Pacific based distributions (in-cluding that of the Micropterigidae), the historical roleof the Pacific in the ‘Gondwana’ biota may be far lessresolved than implied by Ghosts of Gondwana. Be-cause Wegener’s model of Gondwana (that proposesa permanent Pacific ocean) is accepted as an empiri-cal reality, any phylogenetic relationship linking NewZealand closer to Australia than South America is nec-

Fig. 5. Biogeography of tuatara and closest fossil rela-tives (from Reynoso and Clark 1998).

Fig. 6. Biogeography of Rhytidae land snails (fromEmberton 1990, Van Bruggen 1998).


essarily classified as over water dispersal because theclassical breakup sequence requires New Zealand tohave separated earlier than Australia and SouthAmerica. The alternative possibility, that trans-Tasmanpatterns are derived from a Pacific accretion pre-dat-ing New Zealand’s geological isolation remains a criti-cally important biogeographic question to be ad-dressed in the popular literature.

According to Ghost of Gondwana, molecular stud-ies “can tell us the location of ancestral groups, whenthe founders arrived, and how rapidly they diversified”.But this is not quite the whole story. The location ofancestral groups is not derived from molecules, but atheoretical speculation, proposed by Darwin, thatvicariant distributions evolve from geographically nar-row centers from which they disperse according totheir individual means of dispersal. The theorizedcenter of origin is located according to one or moresometimes contradictory theoretical criteria. Molecu-lar studies usually use the location of the basal line-age, the oldest fossil, or area of greatest taxonomicdiversity to denote the theorized center of origin. Noneof these criteria empirically demonstrate anything otherthan records of past or present distribution. The timingof ‘founders’ is also problematic as it represents a theo-retical extrapolation of the fossil record. Since fossilsrepresent minimal age of fossilization, any clock-likemodel can also only predict minimal dates of divergence.In practice molecular dispersalists treat the fossil recordas more or less the actual history of a group and mo-lecular divergences are therefore said to falsify earlierevents if such events predate the divergence estimate.Such claims are methodologically invalid; for all thatthey are extremely popular (Heads 2005).

Molecular dispersalism relies on theory (e.g. mod-els of molecular evolution, molecular substitution, andfossil calibration), whereas panbiogeography relies onthe facts of distribution that are accepted by everyone,including molecular biogeographers. While Ghosts ofGondwana treats the molecular model as an independ-ent ‘test’ (of what, is not quite clear), so too is thespatial analysis that has generated the facts of NewZealand’s biogeographic structure (its composite bio-geography, its correlation with accretion and platetectonics, parallel arcs, etc.). Ghosts of Gondwana’scharacterization of panbiogeography as a spatial analy-sis lacking the element of time is also incorrect sincethe method provides a temporal estimate through thespatial correlation of distributions and tectonic struc-ture. Geological correlations sometimes predict agesolder than the oldest fossil and so provide a potentialfalsifier of the minimal estimates predicted throughfossils and the molecular-fossil clock.

With its emphasis on centers of origin and disper-sal, classification of organisms into supposed ancientelements, dispersal over the sea according to molecu-

lar clock speculations, and double invasions, Ghostsof Gondwana seems somewhat reminiscent of SirCharles Fleming’s (1979) Geological history of NewZealand and its life dressed up in molecular technol-ogy. Since there is such an overwhelming focus onmolecular clock theory as both a conceptual and tech-nical advance over panbiogeography and other bio-geographic approaches, the book seems to inevitablyfalls short of the author’s goal of providing an overallbalanced description of current biogeographic researchon New Zealand’s evolution. But these concerns aside,the book does stand out as a unique and significantcontribution to the popularization of New Zealand bio-geography and evolution. Ghosts of Gondwana cov-ers a tremendous range of examples and stands out instark contrast to other major works supporting a tra-ditionalist view on New Zealand evolution by at leastacknowledging the existence of an alternative. Becauseso many organisms are discussed, and so many ques-tions are raised, it will make an excellent student text-book (indeed it is already being used for the courseNew Zealand Flora & Fauna at Victoria University ofWellington) if students are encouraged to explore thebiogeographic literature in general. Only in this waywill the author’s goals, and real potential and value ofGhosts of Gondwana, be achieved.

References

Cranwell LM. 1962. Endemism and isolation in the Three Kings Is-lands, New Zealand - with notes on pollen and spore types of theendemics. Records of the Auckland Institute and Museum 5: 215-232.Cranwell LM. 1964. Ancient Pacific floras. University of HawaiiPress, Honolulu.Craw RC. 1989. Continuing the synthesis between panbiog-eography, phylogenetic systematics and geology as illustrated byempirical studies on the biogeography of New Zealand and Chat-ham Islands. Systematic Zoology 37: 291-310.Craw RC, Grehan JR, Heads MJ. 1999. Panbiogeography: Track-ing the history of life. Oxford University Press, New York.Emberton KC. 1990. Acavid land snails of Madagascar : Subgenericrevision based on published data (Gastropoda: Pulmonata: Stylo-matophora). Proceedings of the Academy of Natural Sciences ofPhiladelphia 142: 101-117.Fleming CA. 1979. The geological history of New Zealand and itslife. Auckland University Press/Oxford University Press.Gibbs GW. 1983. Evolution of the Micropterigidae (Lepidoptera)in the SW Pacific. GeoJournal 7: 505-510.Gibbs GW. 1990. Local of global? Biogeography of some primitiveLepidoptera in New Zealand. New Zealand Journal of Zoology 16:689-698.Grehan JR. 2001. Biogeography and evolution of the Galapagos:Integration of the biological and geological evidence. BiologicalJournal of the Linnean Society 74: 267-287.Heads MJ. 1983. Pacific plate biogeography, with special refer-


ence to shorefishes. Journal of Biogeography 10: 543-548.Heads MJ. 1994. Biogeography and evolution in the Hebe com-plex (Scrophulariaceae): Leonohebe and Chionohebe. Candollea 49:81-119.Heads MJ. 2005. Dating nodes on molecular phylogenies: A cri-tique of molecular biogeography. Cladistics 21: 62-78.Heads MJ. 2006. Panbiogeography of Nothofagus (Nothofag-aceae): Analysis of the main species massings. Journal of Biogeog-raphy 33: 1066-1075.Heads MJ. 2006. Seed plants of Fiji: An ecological analysis. Bio-logical Journal of the Linnean Society 89: 407-431.Kennedy M, Patterson AM, Morales JC, Parsons S, Winnington AP,Spencer HG. 1999. The long and short of it: Branch lengths andthe problem of placing the New Zealand short-tailed bat, Mystacina.Molecular Phylogenetics and Evolution 13: 405-416.Knapp M, Stöckler K, Havell D, Delsuc F, Sebastiani F, Lockhart PJ.2005. Relaxed molecular clock provides evidence for long-distance dis-persal of Nothofagus (southern beech). PLoS Biology 3: 38-43.Koopman KF. 1984. Orders and families of recent mammals of theworld. John Wiley & Sons, Inc. New York.Matthews C. 1990. Panbiogeography special issue. New ZealandJournal of Zoology 16: 1-815.Reynoso VH, Clark JM. 1998. A dwarf Sphenodontian from theJurassic La Boca Formation of Mexico. Journal of Vertebrate Paleont-ology 18: 333-339.Van Bruggen AC. 1998 Gondwanaland connections in the terrestrialmolluscs of Africa and Australia. Journal of the Malacological Societyof Australia 4: 215-222.Wagstaff SJ, Bayly MJ, Garnock-Jones PJ, Albach DC. 2002. Clas-sification, origin, and diversification of the New Zealand hebes(Scrophulariaceae). Annals of the Missouri Botanical Garden 89:38–63.Wright SD, Yong CG, Dawson JW, Whittaker DJ, Gardner RC.2000. Riding The Ice Age El Nino? Pacific biogeography and evolu-tion of Metrosideros subg. Metrosideros (Myrtaceae) inferred fromnrDNA. Proceedings of the National Academy of Science, USA 97(8):4118-4123.

John GrehanBuffalo Museum of Science, 1020 Humboldt Park-way, Buffalo, NY 14211-1193, [email protected]



Who is Carl Julius August von Minding (1808–1850)?:Some data on the history of biogeography

BY ALEXANDER I. KAFANOV

The paper presents a concise overview of the life andbiogeographic ideas of the German physician, writer,and natural philosopher Julius Minding (November 8,1808 - September 7, 1850). His book entitled “Überdie geographische Verteilung der Säugethiere” (Mind-ing 1829) played a rather important role in the devel-opment of regional approach of zoogeography.

A small book by Julius Minding of only 104 pagesentitled “On geographic subdivision of mammals”(Minding 1829) (Fig. 1) is deposited in the library ofthe Zoological Institute of the Russian Academy ofSciences in St. Petersburg. Strangely enough, this bookis not listed in data bases of the largest libraries of theworld (for instance, The Library of Congress, The Brit-ish Library, etc.) though it result Engelmann (1846)and Agassiz (1852). Neither is it listed by Wood (1931).Carus (1872) mentions Minding only in passing, whenhe cites another book (Minding 1832). Hofsten (1916)writes that Minding explains an origin of the disjunctfaunas by the action of “creative force” [Schöpfung-skraft] “…which aspires everywhere to formation ofsimilar organisms which however there are only analo-gous and not absolutely similar essences under influ-ence of local conditions in the removed districts” [“J.Minding… nahm seine Zuflucht zu einer “Schöpfung-skraft”, die überall zur Bildung gleichartiger Organis-men strebt, von den lokalen Verhältnissen aber derartbeeinflußt wird, daß in entfernten Gegenden nuranaloge und nicht ganz gleiche Geschöpte entstehen”](Hofsten 1916, p. 81). To Russian-speaking readers,Minding is known through the work by Menzbir (1882),which briefly summarizes the key ideas of the book inquestion.

The major part of Minding’s book (Minding 1829,pp. 46-104) borrows extensive “Tables of geographi-cal subdivision of mammals” [“Tabellen über geogra-phische Vertheilung der Säugethiere”] (Fig. 2). In ta-bles on the basis of added data (Illiger 1812) Mindingpresents a list of 1230 Mammalia species for the basicregions of the Earth (Europe, northern and southernAsia, Africa, North and South America, Australia).

In an introductory part of the book Minding’s bio-

geographical views are presented. He considers thattaking into account physical geography alone is notsufficient for “natural” delimitation of continents. Ofequal importance is geographical distribution of or-ganisms, which closely interact with environmentalconditions. For instance, the Sahara and Libian deserts,rather than the Mediterranean Sea, should be consid-ered as the border between Europe and Africa [“AndernTheils möchen auch die alten Annahmen der Scheidezwischen Europa und Afrika verschiedene geogra-phische und geognostische Ansichten gegen sich haben,da Binnenmeere, wie das Mittelländische, wirklich mehralsgrosse Flussbetten, deren Gebiet sich nach beidenSeiten ihres Laufes erstreckt, zu betrachten wären; eineAnnahme, in deren Folge die eigentliche Grenzezwischen Europa und Afrika durch die saharische undlibische Wüste gebildet würde”] (Minding 1829, p. 5).

On the basis of the subdivision of the Earth intothe Eastern and Western hemispheres (rather than intothe Northern and Southern, as generally acceptednowadays), Minding proposed the following originaldivision of the Earth’s surface: the Northeastern land[Festland, the firm ground] (30-78° N, 8-190° E) in-cluding Europe and North Asia; the Northwestern land(to the north of 23° N and 190–360° E) including NorthAmerica; the Southwestern land (23-54° S, 250-344°E) including South America; and the Southeastern land(30-47° S, 1-250° E) including Africa, South Asia, andAustralia. The “Festlands” established by Minding, al-though Minding himself does nor use the term “re-gion”, obviously correspond to faunistic regions.

Minding did not consider rivers, narrow straits, etc.as barriers to species distribution, but referred to themas conditions (like climate and soil) promoting expan-sion of species. [“Flüsse sind so wenig als schmaleMeerengen für Thierscheiden anzusehen; im Gegen-theil pflegen sich verschiedene Arten gern längs ihremLaufe zu beiden Seiten aufzuhalten, was wohl in derGleichmässigkeit des Bodens und Climas seinen Grundhaben mag” (Minding 1829, p. 11)].

Minding is consistent in his belief in the perma-nency of continents and oceans, rejecting the concept


of multiple “land bridges”. “The assumption that landmasses were connected in the past, although exten-sively argued for, especially by earlier authors, seemsto be so highly conjectural and inadequate, that nofurther arguments are needed to disprove the state-ment that such bridges may have ever existed. – Amore precise comparison suggests that such assump-tions are needless; clear differences between distinctland masses are as ancient as the contemporary pat-tern of the Earth’s surface, which has been affectedonly by recent revolutionary changes”. [“Die Annahmeverbindender, untergegangener Länder, zu der so Viele,insbesondere in frühener Zeiten ihre Zuflucht nehmenzu müssen glaubten, ist so hypothetisch und zugleichso unzureichend, dass es keiner Beweise wider dieMöglichkeit solcher Existenzen bedarf. – Eine genauereVergleichung führt auch zugleich auf das Unnöthigederselben hin, indem die strengen Verschiedenheitengetrennter Länder mehr auf eine Trennung hindeuten,die so alt ist, als die jetzige Gestalt der Erdoberfläche,denn auf eine, erst durch spätere Revolutionen aufge-hobene Verbindung”] (Minding 1829, p. 11).

Minding suggests that there might have been twoor more centers of Creation for each species. He con-siders it a special proof of harmony in nature, which

Fig. 2. Pages 92-92 of Minding’s (1829) book with “Tabellen über geographische Vertheilung der Säugethiere”.

Fig. 1. Title page of Minding’s (1829) book.


simultaneously introduces the same living beings inregions with similar environmental conditions. Onpages 27 to 37 of the book, the author treats at lengthwhat is presently termed as geographic vicariance:since the nature tends to introduce more and moreperfect forms, fully interchangeable, if not identical,forms must appear in different regions with similarenvironmental conditions. However, not all species andgenera have vicarious forms, because, environmentalconditions are never completely identical.

Unlike Buffon (1776), Minding suggested that fau-nas on different continents were not of the same age.On each continent, faunas comprise a variety of forms,which were introduced at various times in such a waythat depending on the environmental conditions ei-ther similar or different forms populated different con-tinents at any given period (Minding 1829, p. 11).

Summarizing what has been said above, one clearlysees that Minding’s (1829) book is the second great-est, after C. Illiger’s works (1811, 1812), contributionto the regional approach in zoogeography and there-fore is worth referring to as a classical treatise on bio-geography. Surprisingly, personal information aboutJ. Minding is hardly available. Some data about himcan be found in the free Wikipedia encyclopaedia(http://de.wikipedia.org/wiki/Julius_Minding), which,without mentioning the book (Minding 1829) in ques-tion, says the following:

“Julius Minding was born on November 8, 1808 inBreslau, Germany (Wroclaw, as it now called, Poland).In 1833 he graduated from Berlin University with hisdoctor of medicine degree [Dr. Med.] after submittinghis thesis ‘De vitæ functionum perturbationibusnotiones generales. Dissertatio inauguralis pathologica,etc.’ At that time he entered private practice and si-multaneously started his literary work (for instance,he translated ‘Frithjofs-Saga’ by Esaias Tegnér fromSwedish into German). Later, he involved himself intoa risky commercial venture. Being on the verge of bank-ruptcy he moved to New York in 1850. There he andhis friend established a medical bureau. This venturealso failed. On September 1, 1850, J. Minding com-mitted suicide” (http://de.wikipedia.org/wiki/Julius_Minding).

Since J. Minding is undoubtedly an influential fig-ure in biogeography, further studies of his biographyand development of his expertise are needed. Of par-ticular interest is the origin of his biogeographic views.

Acknowledgements

I am grateful to Drs. Igor Smirnov and Sophia Ste-panjants (both at the Zoological Institute, RussianAcademy of Sciences) for providing much literature.This paper is partly supported by the grant of the Far

Eastern Branch, Russian Academy of Sciences.

References

Agassiz JLR. 1852. Bibliographia zoologiæ et geologiæ. A generalcatalogue of all books, tracts, and memoirs on zoology and geol-ogy. Vol. 3. London: Print. for the Ray Soc.Buffon GLL. Comte de. 1776. Histoire naturelle, générale et parti-culière, etc. Supplément. T. 3 : …servant de suite à l’histoire desanimaux quadripèdes. Paris, De l’Impr. Royale.Carus JV. 1872. Geschichte der Zoologie bis auf Joh. Müller undCharl. Darwin. München, R. Oldenbourg.Engelmann W. 1846. Bibliotheca historico-naturalis. Verzeichnissder Bücher über Naturgeschichte, welche in Deutschland,Scandinavien, Holland, England, Frankreich, Italien und Spanien inden Jahren 1700-1846 erschienen sind. Bd 1. Bücherkunde.Hülfsmittel. Allgemeine Schriften. Vergleichende Abatomie undPhysiologie. Zoologie. Palaeontologie. Mit einem Namen- undSachregister. Leipzig: W. Engelmann.Illiger JKW. 1811. Prodromus systematis mammalium et avium;additis terminis zoographicis utriusque classis, eorumque versionegermanica. Berolini: C. Salfeld.Illiger JKW. 1812. Überblick der Säugethiere nach ihrer Vertheilungüber die Welttheile. Abhandlungen der Königliche Akademie derWissenschaften zu Berlin. Physische Klasse für 1804-1811: 39-160.Menzbir MA. 1882. Ornithological geography of European Rus-sia. Pt. 1. Moscow: Imperial Moscow University [in Russian].Minding J. 1829. Über die geographische Verteilung derSäugethiere. Berlin: Anslinsche Buchhandlung.Minding J. 1832. Lehrbuch der Naturgeschichte der Fische. Berlin:A. Rücker.Wood CA. 1931. An introduction to the literature of vertebratezoology: Based chiefly on the titles in the Blacker Library of Zool-ogy and other libraries of McGill University, Montreal. Oxford: Ox-ford University Press.

Alexander I. KafanovA.V. Zhirmunsky Institute of Marine Biology, FarEastern Branch, Russian Academy of Sciences,Vladivostok, 690041, [email protected]


S O F T W A R E R E V I E W

En la actualidad, la geoinformática, el creciente de-sarrollo de algoritmos o secuencias automatizadasde instrucciones y la tendencia a incorporar tecno-logías modernas, como los Sistemas de InformaciónGeográfica (SIG), han motivado a que los biólogos seenfrenten con el problema de representar en un mapalos patrones espacio-temporales de la biodiversidad(Morrone 2000, Tikunov 2002). Uno de los métodosque involucra principios, técnicas y tecnologías a ni-vel geográfico e informático es el método de la pan-biogeografía. De acuerdo con Grehan (2001), la prác-tica del método de la panbiogeografía en lugarescomo América Latina, el suroeste del Pacífico y Euro-pa se ha extendido desde la década de los 1990’s, nosolo porque algunos autores, como Morrone y Crisci,han adoptado el método panbiogeográfico para in-tegrarlo con los análisis biogeográficos cladísticos,sino también porque el método panbiogeográficoinvolucra el aspecto geográfico dentro del análisis dela evolución, parámetro que en gran parte de los aná-lisis cladísticos se reduce a las ramas terminales delos cladogramas, construidos a partir de análisis en-tre caracteres biológicos.

Por los motivos anteriores se planteó el reto deformalizar una aproximación al método de la panbio-geografía desde un punto de vista informático, desa-rrollando una herramienta para crear trazos indi-viduales, en lenguaje de programación AVENUE® paraArcView GIS 3.2® (ESRI 1996).

Antecedentes

El uso en la biología de algoritmos como modelos depredicción de patrones espaciales de la biodiversidad,ha adquirido gran importancia en los últimos años paraprobar hipótesis biogeográficas, mejorar atlas florís-ticos y faunísticos, y definir áreas prioritarias para laconservación (Margules y Austin 1994, Hausser 1995,Kienast et al. 1996, 1998, Mourell y Ezcurra 1996). Lamayoría de estos modelos se basan en el análisis de larelación especies-ambiente, asunto central de la bio-

Una herramienta automatizadapara realizar análisis panbiogeográficos

geografía y la ecología (McArthur 1972, Myers y Giller1988, Espinosa et al. 1993, Arita y Rodríguez 2002,Llorente et al. 2002), que permiten predecir la organi-zación u ordenamiento espacial de los individuos yayuda a tener mayores parámetros para el conocimien-to sobre la biodiversidad. Los ejemplos más importan-tes del uso de algoritmos en la biología están relacio-nados principalmente con los modelos de predicciónde los patrones espaciales, que involucran la instru-mentación de los Sistemas de Información Geográfica(SIG) y el uso y desarrollo de algoritmos genéticos comoel COVER (Moeur 1985), BIOCLIM (Nix 1986), GARP(Genetic Algorithm for Rule-set Prediction) (Stockwelly Noble 1991, Stockwell 1999, Gouch y Rushton 2000,Murguía y Rojas 2001, Navarro et al. 2003) y GAP Ana-lysis (Scout et al. 1993), entre otros.

El problema del árbol de extensión mínima

La base que fundamenta la instrumentación del méto-do panbiogeográfico es el trazo individual, que extra-polado a un plano matemático es un conjunto mínimode enlaces de N que conectan todos los nodos en A y porlo tanto al menos un árbol de expansión puede ser en-contrado en un grafo G. El árbol mínimo de expansióndenotado por T* es un árbol de expansión cuyo pesototal de todo los enlaces es mínimo o “árbol de exten-sión mínima” (Campos 2002, Milková 2004). Es decir:

∑=Ee

ijt

ij

wT

εmin*

Formalmente, un grafo G consiste en dos conjun-tos finitos N y A. N es el conjunto de elementos delgrafo, también denominado vértices o nodos. A es elconjunto de arcos, que son las conexiones que se en-cargan de relacionar los nodos para formar el grafo.Los arcos también son llamados aristas o líneas. Losnodos suelen usarse para representar objetos y los ar-cos para representar la relación entre ellos. Cada enla-

POR CAMILO ANDRÉS ROJAS PARRA


ce tiene un número positivo real asociado denotadopor W que representa la distancia, el costo o un valorcualquiera de datos dado (Milková 2004).

Luego, para que un árbol de expansión sea míni-mo debe cumplir una de tales condiciones:

ser un grafo de G conectado con n –1 enlaces.ser un subgrafo de G sin ciclos con n –1 enlaces.

Algunos algoritmos de tiempo polinomial, que seresuelven acotados por un polinomio, se desarrolla-ron para solucionar el problema del árbol de exten-sión mínima entre los que se encuentran los algoritmosde Prim, Kruskal, Dijisktra y Solin (Milková 2004).

El algoritmo de Prim

El algoritmo de Prim es una de las soluciones de tiem-po polinomial desarrolladas para solucionar el proble-ma del árbol de extensión mínima (Fig. 1). Las especi-ficaciones para programar el algoritmo son de lasmenos complejas. Supongamos un grafo G = (N,A),con los arcos etiquetados con su costa. El algoritmode Prim va construyendo un subconjunto de nodos U,aumentándolo hasta que contenga todos los nodosde N. Inicialmente, U contiene un único nodo de N,que puede ser cualquiera. En cada paso se localiza elarco más corto (u, n) tal que u∈ U y n∈ N, repitiendoel proceso hasta que U = N.

Algoritmo Prim (G: Grafo; VAR T; Conjunto de Arcos);

VariablesU: Conjunto de arcosu, n: arcosInicio

T:={}U:={Cualquier n∈ N}MIENTRAS U <> N HACER

Sea (u,n) el arco de costo mínimo tal que u∈ Uy n∈N.

T: = T {(u,n)}U:=U ∪ {(n)}

FIN MIENTRASFin Prim

Para encontrar fácilmente el arco de menor costoentre U y N-U, se mantienen dos matrices, MAS_CERCANO[i], que devuelve el nodo en U más cercano a i enN-U y MENOR_COSTE [i] que devuelve el coste del arco(i, MAS_CERCANO[i]). En cada paso, se escoge k talque k N-U es el nodo más cercano a U y se actuali-zan ambas matrices teniendo en cuenta que ahorak∈U. Para evitar que k vuelva a ser seleccionado, seasigna MENOR_COSTE[k]= .

Fig. 1. EL algoritmo de PRIM. Representación de lassoluciones de tiempo polinomial desarrolladas parasolucionar el problema del árbol de extensión mínima.


Conclusiones

La herramienta de trabajo Trazos 2004 ofrece unainterfaz gráfica al usuario, dentro de la plataformaArcView 3x, que permite llevar un control del procesoy desarrollo de las tres etapas básicas del método dela panbiogeografía, a través de temas espacialesgráficos referidos a un sistema de coordenadas, demanera sencilla. El hecho de catalogar como herra-mienta el desarrollo aquí presentado significa que sufunción es la de ser una extensión, un módulo adicionalde nuestras capacidades, que facilita realizar una tareaespecífica dentro del método de la panbiogeografía,de manera automatizada pero bajo la supervisión ycriterio del investigador.

La automatización computadorizada del métodopanbiogeográfico, a través de Trazos 2004, agiliza eldelineado gráfico de los trazos, independientementede la complejidad espacial que tenga la nube de puntosque representan las localidades del taxon o grupo detaxones que se estén investigando. A pesar de que nose trata de un programa computadorizado orientadoa su comercialización, permite a diferentes grupos deinvestigación continuar con el desarrollo de otra seriede adiciones y aplicaciones informáticas que ayuden amejorar la herramienta y el método de la panbio-geografía en sí mismo. Para el método de la panbio-geografía no se había desarrollado una herramientaautomatizada como la que se presenta en estainvestigación. La herramienta se encuentra en procesode desarrollo y mejoramiento.

En la actualidad, se están desarrollando nuevosmétodos biogeográficos para obtener modelos depredicción de los patrones espaciales de las especies,que involucran el uso de la panbiogeografía y otrasáreas de estudio como los sistemas, la estadística y lasmatemáticas. Estas aproximaciones hacen que Trazos2004 sea una herramienta sujeta a crecimiento sinpresentar dificultades para la incorporación de nuevosmétodos.

Referencias

Arita H, Rodríguez P. 2002. Ecología geográfica y macroecología.En: Llorente J, Morrone JJ (eds.). Introducción a la biogeografía enLatinoamérica: Teorías, conceptos, métodos y aplicaciones. Las Pren-sas de Ciencias, Facultad de Ciencias, UNAM, México, D.F.Espinosa DN, Llorente J. 1993. Fundamentos de biogeografíasfilogenéticas. Coordinación de Servicios Editoriales, Facultad deCiencias, UNAM, México, D.F.Gough MC, Rushton SP. 2000. The application of GIS-modellingto mustelid landscape ecology. Mammal Review 30 (3-4): 197-216.Grehan JR. 2001. Panbiogeography from tracks to ocean basins:Evolving perspectives. Journal of Biogeography 28: 413-429.Kienast F. 1998. Towards a socially accepted, sustainable manage-

ment of agricultural landscapes. En: Schellnhuber, H. M. y V. Wenzel(ed.). Earth system analysis: Integrating science for sustainability.Llorente J, Papavero N, Bueno A. 2002. Síntesis histórica de labiogeografía. en: Llorente J, Morrone JJ (eds.). Introducción a labiogeografía en Latinoamérica: Teorías, conceptos, métodos y apli-caciones. Las Prensas de Ciencias, Facultad de Ciencias, UNAM,México, D.F.Margules CR, Austin MP. 1994. Biological models for monitoringspecies decline: The construction and use of data bases. Philosophi-cal Transactions of the Royal Society of London, B 344: 69-75.McArthur RH. 1972. Geographical ecology: Patterns in the distri-bution of species. Harper and Row, New York.Moeur M. 1985. COVER: A User’s Guide to the CANOPY and SHRUBSExtensions of the Stand Prognosis Model. General Technical Re-port INT-190.Ogden UT. U.S. Department of Agriculture, Forest Service, Inter-mountain Research Station.Morrone JJ. 2000. Atlas biogeográficos. PrIBES. http://entomologia.rediris.es/pribes.Mourell C, Ezcurra E. 1996. Species richness of Argentine cacti: Atest of biogeographic hypotheses. Journal of Vegetation Science,7: 667-680.Murguía M, Rojas F. 2001. Biogeografía cuantitativa. En: LlorenteJ, Morrone JJ (eds.). Introducción a la biogeografía en Latinoa-mérica: Teorías, conceptos, métodos y aplicaciones. Las Prensas deCiencias, Facultad de Ciencias, UNAM, México, D.F.Myers AA, Giller PS (eds.). 1988. Analytical biogeography: Anintegrated approach to the study of animal and plant distribution.Chapman and Hall, Londres.Navarro AG, Peterson AT, Nakazawa J, Liebig-Fossas UEI. 2003.Colecciones biológicas, modelaje de nichos ecológicos y los estu-dios de la biodiversidad. En: Morrone JJ, Llorente J (eds.). Una pers-pectiva latinoamericana de la biogeografía. Las Prensas de Cien-cias, Facultad de Ciencias, UNAM, México, D.F.Nix HA. 1986. A biogeographic analysis of Australian elapid snakes.En: Atlas of elapid snakes of Australia. Longmore R (ed.), AustralianGovernment Publishing Service, Cambridge.Stockwell DRB. 1999. Genetic Algorithms II: Species DistributionModelling. En: Fielding AH (ed.). Machine learning methods forecological applications. Kluwer Academic Publishers. Cambridge,Massachussets, pp. 124-144.Stockwell DRB, Noble IR. 1991. Induction of sets of rules fromanimal distribution data: A robust and informative method of dataanalysis. Mathematics and Computers in Simulation 32: 249-254.Tikunov VS. 2002. La revolución de la información en geografía.Traducido del ruso. http://www.unesco.org/issj/rics150/tikunov150.htm.

Camilo Andrés Rojas Parra377 Placid Lake Drive, Sanford, 32773 Florida, [email protected]


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