Biogeografa 01 january 2007

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A new voice for biogeographers

In early 2006 Juan J. Morrone and Malte C. Ebach decided it was high time for a biogeographical society aimed at those academics, students and amateurs that are isolated from the publishing hubs of Europe and North America, where most of the learned “international” societies focus all their attention. After all, most biogeographical 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 at least Nature or Science. Imagine a western European university or a North American natural history museum without a functioning photocopier, a decent library or access to any online journal. Unfortunately for most Latin American institutions this is a reality. Without proper access to resources and the difficulty of publishing in 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 biogeographers through access to communication tools such as a multilingual list server (Biogeography Portal) and an electronic Bulletin. The Biogeography Portal, currently with over 300 members, is an active repository for forums and electronic documents such as pdfs. It has helped many to access the latest news and voice their opinions 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 article on Otto Kleinschmidt and his influence on early 20th century biogeography, a personal journey through Croizat’s life 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 French to 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. Ebach John R. Grehan Juan J. Morrone Co-editors, SEBA Bulletin, 2007.

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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 of zoogeography (and evolution), its basis founded on the dispersal of organisms over a static and unchanging globe (Croizat 1958, p. xi). The idea was perhaps first graphically captured by Ernst Haeckel’s diagram of wandering humans (Haeckel 1870, Taf. XV, see Williams 2006 [2007]), a visual image that remains in use today to illustrate the route humans wander from their place of origin (Kohn 2006, Shreeve 2006). The New York School’s primary developers, in chronological 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 (19051985)” – one might also add Jack Briggs (Briggs 1981, see Nelson & Ladiges 2001) and much of what goes under the heading of phylogeography (Avise 2000, Riddle & Hafner 2006 [2007], see Nelson 2004a, p. 134 and Heads 2005b, p. 681).

Ernst Haeckel’s (1834-1919) influence on the development of evolutionary theory was probably more profound than Darwin’s: Haeckel was “the chief source of the world’s knowledge of Darwinism” (Nordenskiöld 1936, p. 515). Among his many neologisms, Haeckel proposed chorology to mean the “…science of the geographic and topographic spread of organisms” (Haeckel 1868, pp. 286-289, translated): “I mean chorology, or the theory of the local distribution of organisms over the surface of the earth. By this I do not only mean the geographical distribution of animal and vegetable species over the different parts and provinces of the earth, over continents and islands, seas, and rivers, but also their topographical distribution in a vertical direction, their ascending to the heights of mountains, and their descending into the 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 speak of an independent science of chorology…” (Haeckel 1925, pp. 365-366). He continues: “The most important principle from which we must start in chorology, and of the truth on which we are convinced by due examination of the theory of selec-

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

Chorology was, then, the “science of migrations” (Haeckel 1904, p. 98, Third Table) from a single locality, an organism’s “central point of creation”. For the promotion and generalisation of the idea, William Diller Matthew’s Climate and evolution (1915), published while Haeckel was still alive, helped gain the idea respectability and influence.

In a recent history of panbiogeography, Heads noted that: “Matthew’s book [Matthew 1915] capitalized on the fascination of the American public with vertebrate palaeontology which had existed since the time of Thomas Jefferson’s exploits and became the most influential text in the history of biogeography. Its ideas were promulgated by Simpson, Mayr, Darlington and others, forming what Croizat (1958) labelled the ‘New York school of zoogeography’ (cf. Croizat 1984, Nelson and Ladiges 2001; for some reason Nelson and Ladiges did not include Mayr in the school)” (Heads 2005a, p. 89, 2005b, p. 63, 2005c, p. 681).

With respect to the omission of Ernst Mayr from the New York school, Nelson remarked that “Mayr does not derive from Matthew but rather from Stresemann” (Nelson pers. comm.). Croizat commented that “The latter [Mayr] in his turn, by grafting Stresemann upon Matthew, has greatly influenced North American, and world-wide, ‘biogeographic’ thinking” (Croizat 1964a, p. 179, fn 1).

Writing of early panbiogeographers, or perhaps ‘pre-panbiogeographers’, Croizat suggested that the ornithologist Otto Kleinschmidt (1870-1954) was a pioneer, if not creator of panbiogeography: “Suffice it here to conclude that Kleinschmidt is one of the first true biogeographers, to the extent at

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least that he correctly perceived that form-making is a process not to be dissociated from space and time. In sum, (pan)biogeography was not born in 1958 with me, but in 1897 with Kleinschmidt, whom everybody does 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 century ornithologists, including Erwin Stresemann (18891972). It would thus seem, potentially at least, that the biogeographic genesis of Mayr and Croizat might both lead back to Kleinschmidt.

Otto Kleinschmidt (1870-1954) and the ‘Formenkreis theory’ Otto Kleinschmidt was a leading ornithologist, protestant pastor, creationist and sometime Nazi sympathiser (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’s contribution to the species debate was to note that similar ‘forms’ (species) occur on separate and sometimes distant geographic regions. To explain this fact he developed the ‘Formenkreis theory’ to account for these closely related ‘forms’. While literally meaning ‘form-circles’, the allusion to a circle was coincidental as “the word ‘Kreis’ on the expression ‘Formenkreis’ means the area of distribution of a certain animal on the land or in the waters of the globe” (Kleinschmidt 1930, p. 27, see Kleinschmidt 1926). The many similar forms occurring within a certain ‘type’ would have a fixed suite of characters, detectable by examination.

For Kleinschmidt, the Formenkreis suggested an unchangeable (but variable) entity, and hence allowed him to support and promote a creationist view of the origin of species – a viewpoint not unlike that of Louis Agassiz (see below).

Contemporary comment on Kleinschmidt is favourable: “I think there is little doubt that Kleinschmidt’s Formenkreis concept represents, as claimed by Mayr, “Systematics and the origin of species”, 1942, p. 112, ‘one of the most productive working hypotheses of taxonomy’, and while new work has necessitated the overhaul of certain views expressed by Kleinschmidt (1900, 1930), the principles of the concept have been widely accepted in the scientific world and have won their author imperishable fame.” (Clancey 1950, p. 32)

Ernst Mayr and Erwin Stresemann Haffer et al. noted that “After his high school exami-

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nations (Abitur), Stresemann entered the University of Jena in 1908, where he took courses offered by Ernst Haeckel” (Haffer et al. 2000, pp. 399-400, see also Haffer 1997, p. 829). In 1931, Stresemann “declared historical morphology as terminated, i.e. phylogenetic or systematic morphology…in the sense of Ernst Haeckel…and Max Fürbringer…” (Haffer et al. 2003: 420). The meaning suggests that Stresemann relinquished the pursuit of homology (the discovery of relationships among organisms, as in either “phylogenetic or systematic morphology”) and embraced the beginnings of what might be called the ‘genetic’ or ‘populational’ point of view, the current evolutionary Zeitgeist.

Stresemann was said to have applied Kleinschmidt’s Formenkreis theory to his taxonomic work “without accepting its underlying theoretical (typological) premises” (Haffer et al. 2000, p. 410). Stresemann was also said to have been an early proponent of the historicaldynamic method of zoogeographical analysis (Haffer et al. 2000, p. 414), of which an early version can be found in Stresemann & Grote (1929) but a more detailed explanation appeared in Stresemann (1939, see Vuilleumier 2005).

The historical-dynamic method of zoogeographical analysis “…stressed the need to examine entire faunas including the dispersal capabilities and distributional ranges of each species as well as the ecological and geological history of a particular region to understand the zoogeographical history and differentiation of a group of animals as a dynamic and continuing process” (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 of this concept [dynamic faunas]” (Dunn 1922). In Mayr’s 1946 paper, he referred to two examples from German ornithologists, Stegmann (1938) and the previously 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 to Haeckel and is, in fact, a re-statement of chorology, the “…science of the geographic and topographic spread of organisms” (Williams 2006 [2007]).

Stresemann was professor to Ernst Mayr (Junker 2003, Bock 2004). According to Mayr “…Virtually everything in Mayr’s 1942 book was somewhat based on Stresemann’s earlier publications” (Mayr 1999, p. 23, see Mayr 1997). According to Bock, Mayr’s “first and last interest in ornithology is biogeography…”, where he applied “the (then) new ideas for analyzing the biogeography 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-dynamic method of zoogeographical analysis” – chorology.

Species and their ‘parts’ The details and nomenclature for species and their subgroups is unyieldingly complex, words and definitions tumbling from biological and philosophical journals for almost three decades (1930s to the 1960s) and beyond (Wheeler & Meier 2000). In the run up to the ‘modern synthesis’, Bernhard Rensch created two terms to represent different aspects of Kleinschmidt’s Formenkreis: 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 ornithology the term “Artenkreis,” for a systematic unit that was called “Formengruppe,” “Artengruppe,” etc., by several earlier German authors. As there is just about as much difference between Artenkreis and species as there is between species and subspecies, I propose for Artenkreis the more convenient term, superspecies. I define superspecies as a systematic unit containing geographically representative species that have developed characters too distinct to permit the birds to be regarded as subspecies of one species” (Mayr 1931, p. 2).

Artenkreis relate to Huxley’s ‘geographical subgenus’ (Huxley 1938, p. 255), while Rassenkreis (Rensch 1929) relate to Huxley’s polytypic species (Huxley 1938, p. 255), “species that break up into subspecies” (Mayr 1978, p. 86).

Mayr did, of course, modify his definition of superspecies: “A monophyletic group of entirely or essentially allopatric species that are too distinct to be included in a single species” (Mayr 1963, p. 672, 1978, p. 86).

Another composite, semispecies are “populations which have part way completed the process of speciation…” (Mayr 1963, p. 501), they are on the “borderline between subspecies and species” (Mayr 1978, p. 86, after Mayr 1940, p. 260), a definition different from that in Mayr (1942, p. 165). For the 1942 semispecies – species comprising a superspecies – Amadon proposed instead allospecies (Amadon 1966, p. 245), hence a group of allospecies form a superspecies (= ‘Artenkreis’). If not to complicate matters further, Mayr and Short (1970) introduced the ‘zoogeographic species’ for zoogeographic analysis: “As the new polytypic species concept began to assert itself, a certain pessimism seemed to be associated with it. It seemed as if each of the polytypic species (Rassenkreis) was as clearcut and as separated from other species by bridgeless gaps as if it had come into being by a separate act of creation. And this is exactly the conclusion drawn by men such as Kleinschmidt or

Goldschmidt. They claim that all the evidence for intergradation between species in the past was actually based on cases of infraspecific variation, and, in all honesty, it must be admitted that this claim is largely justified” (Mayr, 1942: 114).

The ‘bridgeless gaps’ were Goldschmidt’s conclusion (Goldschmidt 1940, p. 310), the ‘separate act of creation’, Kleinschmidt’s (Kleinschmidt 1930) – and earlier, that of Agassiz.

More on Stresemann Stresemann was also professor to Wilhelm Meise (1901-2002), who “was primarily a systematist studying problems of geographical variation, hybridisation, and speciation in birds” (Haffer, 2003, p. 117). Meise not only worked and published with Mayr (Meise & Mayr 1930) but at the same time was mentoring the 19 year old Willi Hennig (1913-1976), publishing two studies on Rassenkreis of the reptile genus Draco, the only 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 character polarity from geographical proximity, noting that “The use of the chorological method in zoology has become known particularly through the books of Rensch” (Hennig 1966a, p. 133, Brundin 1966, see also Hennig 1950, pp. 192-199 and Kiriakoff 1954), a view that led to Hennig’s ‘progression rule’ (Hennig 1960, 1966b), the view that the direction of dispersal was hypothesised as moving away from a particular centre of origin, those places inhabited by the most basal or plesiomorphic species. Hennig’s ‘progression rule’ was soon rejected (Nelson 1974, see Parenti 2006 [2007]). Bernhard Rensch was also one of Stresemann’s students (Junker 2003).

Thus, one might easily understand Stresemann’s descendants to include some of the leading proponents of the ‘modern synthesis’ (Mayr, Rensch) as well as, albeit indirectly through Wilhelm Meise, Willi Hennig – even including the following generation of phylogeographers (Avise 2000), providing an intellectual lineage from Formenkreis to phylogeography via the progression rule – or from Otto Kleinschmidt to Hennig, according to Croizat, the former “one of the first true biogeographers” (Croizat 1964), the latter irrelevant (Croizat 1978).

Discussion Both Kleinschmidt and Hennig, and perhaps all the others discussed above, understood species in the geographical dimension by investigating their ‘origin’,

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

The idea of a single origin for organisms can be traced to the Bible, a view given substance by Linnaeus (“If we trace back the multiplication of all plants and animals…we must stop at one original pair of each species”). Darwin found the idea of a single centre of creation captivating: “Undoubtedly there are many cases of extreme difficulty in understanding how the same species could possibly have migrated from some one point to the several distant and isolated points, where now found.

Nevertheless the simplicity of the view that each species was first produced within a single region captivates the mind. He who rejects it, rejects the vera causa of ordinary generation with subsequent migration, and calls in the agency of a miracle. It is universally admitted, that in most cases the area inhabited by a species is continuous; and when a plant or animal inhabits two points so distant from each other, or with an interval of such a nature, that the space could not be easily passed over by migration, the fact is given as something remarkable and exceptional” (Darwin, 1859, p. 352).

Mayr once wrote of the Origin that “…Darwin’s argument was directed against authors like Louis Agassiz, who explained discontinuous distribution of species by multiple independent creations” (Mayr 1982, p. 619). Agassiz wrote on centres of origin and independent creations: “The greatest obstacles in the way of investigating the laws of the distribution of organized beings over the surface of our globe, are to be traced to the views generally entertained about their origin. There is a prevailing opinion, which ascribes to all living beings upon earth one common centre of origin, from which it is supposed they, in the course of time, spread over wider and wider areas, till they finally came into their present state of distribution. And what gives this view a higher recommendation in the opinion of most men is the circumstance, that such a method of distribution is considered as revealed in our sacred writings” (Agassiz 1850, p. 181).

Agassiz saw his ideas as in conflict with the “sacred writings”, rather than embracing them. Agassiz saw the ‘origin’ of the same species occurring many times at many places, in the cases of reported disjunctions – much as Kleinschmidt did (Klienschmidt 1897).

Others, in the post-Darwinian, evolutionary literature, have seen something of value in Agassiz’ approach to biogeography, if not his explanation. Léon Croizat (1894-1982, see Nelson 1973, Craw 1984a, b, Zunino

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

Heads (1985) and Grehan & Ainsworth (1985) offered a view that the problem of species origin resides in the “nature of ancestors” (Heads 1985), such that ancestors are ‘polytopic’ rather than ‘uniform’, in the sense of their characters, a view that relates to Croizat’s polytopism (= “multiple origins”, Heads 1985, p. 209, 2005, p. 105, Grehan 2006, see Croizat 1971 and Aubréville 1969, 1974, 1975a, 1975b). This view can be contrasted with the monotopic ancestor, a position said to have been adopted by both Mayr and Hennig (Heads 1985, but compare with Rensch 1934, p. 102, figure 22, reproduced herein as figure 1). Thus, for Heads (1985) and Grehan & Ainsworth (1985), disjunct species may have arisen by some internal orthogenetic process, giving the illusion that the ‘same’ species occurs in different places, thereby offering an explanation for disjunction and (possibly, we will never know) satisfying Agassiz’ problems of their origin (compare figure 1a with Croizat 1964, p. 783, fig. 84 and Haffer’s interpretation of Kleinschmidt’s Formenkreis in Haffer 1997, p. 797, fig. 47].

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

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

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

When Ernst Mayr died in 2005 there was a veritable flood of obituaries, the good, the bad and the ugly. At times, referred to as the greatest evolutionary biologist since Darwin, some of Mayr’s philosophical pronouncements have become somewhat doubtful. Polly Winsor, for example, has begun to unravel the ‘typology-essentialism-morphology’ (the ‘populational’ viewpoint) story, her account bringing doubt to the accuracy of certain contributions of Arthur Cain (1958, 1959) and David Hull (1965). Mayr’s history of essentialism “in its broad sweep across the history of systematics, this story is not merely inaccurate in particulars, it is wrong and harmful in its basic message” (Winsor 2006, p. 3) the whole forming a “miserable history of depreciatory comment without a particle of truth” (Nelson 2004b). And so it may be with the ever changing species concepts, and their new and more exclusive categories.

One might follow the argument through the generations, contrasting Agassiz with Darwin, Kleinschmidt with Mayr, Croizat with Hennig, the phylogeographers with…well, no opponent has yet emerged. On the other hand, it might be that there really is no possibility of resolution to the problem, other than proposing new ways of discovering the ‘true’ nature of ancestors, determining its characters and proposing a mechanism to explain how those characters change (alter, transform) into their descendants.

In 1820 Candolle wrote a significant essay on biogeography suggesting that “All of the theory of geographical botany rests on the particular idea one holds about the origin of living things and the permanence of species” (Candolle 1820, p. 417, translation in Nelson 1978, p. 285). Perhaps it is now timely to set aside the ebb and flow of ‘the origin of living things’, putting the populational approach to one side, regardless of the contrasting ideas concerning polytopism (= “multiple origins”) or monotopism (= single origin), and examine the classificatory approach to taxa and the areas they occupy, a problem that has at the very least possibilities of solution (Nelson & Platnick 1981).

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Llorente J, Morrone JJ eds. pp. 67-123. Universidad Nacional Autónoma de México, México, D.F. Heads MJ. 2005b. Dating nodes on molecular phylogenies: A cri-

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David M. Williams Department of Botany, The Natural History Museum, Cromwell Road, London SW7 5BD, United Kingdom. dmw@nhm.ac.uk

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Journey to Coro

BY JOHN R. GREHAN My entry into biogeography came about in a rather unpredictable way. Biogeography was the last thing on my mind during the early 1980’s. But I then learned about the biogeographic interests of Robin Craw and Michael Heads and the rest is history.

By early 1988 New Zealand was in the grip of the 1987 stock market crash and there was a corresponding crash in government support for science research.

The entire New Zealand scientific establishment was in turmoil as positions were slashed and opportunities lost. This was no time to begin one’s scientific career. Having finished my PhD (on the evolutionary biology of the ghost moth Aenetus virescens) I decided to make a seven day visit to Coro (Venezuela), where Croizat lived and worked for the last years of his life before passing away in 1982. Michael Heads had earlier visited Coro to study Léon Croizat’s life’s work as a guest of his wife Catalina Croizat.

In 1976 Léon and Catalina Croizat moved to Coro from Caracas and became the first directors of the Jardín Botánico Xerófito, now named after Croizat. Catalina (see figure 1) kindly agreed to my visit and I found myself traveling to Venezuela in July 1988. At that time I had never traveled beyond the South Pacific (Australia, New Caledonia, and Fiji). Since there was to be an Entomology Congress at the University of British Columbia that included a symposium on primitive moths, it seemed like a good opportunity to combine this with my visit to Coro. I flew to Canada via the US and spent the next five days in Vancouver. From there I was joined

by Diane, a friend from New Zealand, and the two of us flew down to Caracas via Chicago and Miami. Neither of us spoke Spanish so it was by a bit of luck and perseverance that we managed to transfer ourselves to the domestic terminal and locate our flight to Coro.

The next leg of the journey was an hour’s flight along the Caribbean coast where we saw a dry, desert-like region of low scattered vegetation, farmland, and sand dunes. We arrived at a small airport not much unlike that of a rural city in New Zealand, with its open landscape and small terminal.

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

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were waiting for our luggage we noticed a small, elderly woman smiling at us and we wondered if she might be Catalina. After a tentative enquiry we found our assumption to be correct. I think we stood out from the usual crowd. I had sent Catalina our travel schedule but we did not know if it was received (mail was then sometimes unpredictable). As she drove us to her home we noticed that she drove quite slowly and with seeming disregard of other vehicles; however, we were later told that everyone in the town knew and liked her and made allowances for her driving.

The Croizats’ house was enclosed by a high wall tipped with broken glass. This was a common defense against forced entry in Coro. I was to learn that security was a major challenge due to the great disparity in economic prosperity in the region. After Catalina endured the experience of having a burglar enter and threaten her, the University arranged for a night security guard. Her house was surrounded by a wealth of euphorbias and other strange (for a home grown New Zealander) tropical plants. At the back of the house was a guest house where we stayed. The center of the house comprised a large, open lounge with a high ceiling and large windows at the top. The walls were lined with paintings, including some of Croizat (Fig. 2). Each morning Catalina, with help from a maid, prepared a breakfast of what were to us exotic fruits and drinks and other food. It seemed that we never had the same thing twice. It also seemed as if we could never eat enough to satisfy what Catalina thought we should have!

My purpose in coming to Coro was to see the botanical gardens, see where Croizat worked, and see some of his works I had not previously accessed. Croizat’s office was left much as it was when he died, and one could almost feel a presence in the room’s quiet solitude. Unfortunately, I never took a photo of this space, but I remember the walls being lined with an array of fascinating books and publications. He even had a well worn, and annotated, copy of Brundin’s Transantarctic Relationships. Looking back I realize that there was much I overlooked – probably a combination of inexperience, culture shock, and jet lag; however, I felt I was in my element 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 supportive people, including Dr. Rodolfo Bastidas, an entomologist at the Coro campus of the Universidad Francisco de Miranda and chair of the Friends of the Jardín whose members worked to maintain funding and other support for the botanical garden operations. Bastidas kindly took Diane and me on a day-long trip to the coastal ranges behind Coro to see the local region and people. We drove through a region of low, dry vegetation with small scattered towns and houses. As we drove higher into the hills we entered an area of cloud forest that was maintained largely by atmospheric moisture. In some ways the vegetation was not much unlike parts of Australia with widely spaced trees, lack of undergrowth, and dry leaf litter. I looked for signs of ghost moth damage (ghost moths are my primary entomological interest) but without success. As we were driving back down towards Coro the road zigzagged back and forth with extremely tight turns decorated by many crosses and small chapels. Catalina later muttered that people too often drove down the hill while drunk and failed to make the turns.

Visiting the Jardín was one of the highlights of our journey. I had no idea what to expect as I had not seen any pictures or detailed descriptions. We drove out a little way on the main highway between Coro and Caracas. Located on the strip of land between the highway and the Caribbean coast, the Jardín entrance was from the highway through a high-walled façade with the name Jardín Botánico Xerófito “Dr. Léon Croizat” on the front and offices behind (Fig. 3). As we entered we found ourselves at a central pathway and low flowing shrubs. Beyond, we could see sand dunes obscuring the beach beyond. Other paths led between different focal areas with mesh-in enclosures and stepped terraces within. Apart from the euphorbias (Fig. 4) and cacti there were some surprises such as two large specimens of Welwitschia mirabilis (Fig. 5) that are native to the desserts of southern Africa. I had heard about these plants and their intriguing evolutionary position so I found their presence in the Jardín to be very exciting. I understand that these two plants were among the largest specimens in the New World. There was also the strange looking baobab tree (a group native to Africa, Madagascar, and northern Australia), a species I had never before other than in books.

Daily operations were supported by a small staff that tended the plants. As I walked around the Jardín I could see signs of the challenges faced by Catalina and supporters of the gardens. Although much of the garden was intact I could see signs of deterioration.

The enclosures were in a state of disrepair and the plants needed attention. Many other plants were still awaiting transfer from the nursery into the garden.

Catalina said the company contracted to work the Jardín was often not paid, and the barrier along the

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center of the highway prevented regular use by visitors such as school groups and university students. Potted plants were available for sale, but the money did not go to support the Jardín. She said there were only about 10 people working the Jardín when 40 were needed.

Dr. Bastida also took us on a tour of the scientific institutions in Coro, including the University campus, and the Centro de Investigaciones de Ecología y Zonas Áridas de la UNEFM. Through the support of Center director Dr. Miriam Díaz I was invited to give an evening presentation on Croizat’s theories to a small audience.

Although I was dealing with a subject that I was still working to understand, I had a few slides with me and I was able to give a brief outline of the principles of biogeography, symmetry, and evolution. Even though I could only communicate in English, the audience was forgiving and the slides at least made it possible to convey the main points. Afterwards Catalina said the presentation agreed very well with her understanding of Croizat’s work. This was most reassuring as I was somewhat nervous about speaking on this topic with 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 herself. I asked to interview her when I arrived, but she had put it off. She now said that rather than an interview she would tell me her story in her own words.

She said she was born in Hungary. Her father was a lawyer while her mother an intellectual and philosopher closely attuned to the importance of general knowledge. She attended a public school and then convent school where she did very well. She attended the Université de Paris Sorbonne, first with financial support from the convent and then through her academic performance. She learned several languages and then taught French and German in Hungary. She married a man from the family of her stepfather. As the Second World War started to unfold, she and her husband decided that they needed to leave Europe and Venezuela seemed to be a very good place to go. Her husband traveled to Venezuela first and she was then to follow. She had just boarded a ship sailing from Genova (Genoa) to Venezuela when Italy entered the war. The ship was attacked and sank. She and other survivors were rescued by another ship and returned to Genova where she came down with pleurisy and became very ill. She was still very sick when she boarded another ship that took her to Venezuela.

In Venezuela Catalina first worked as a cook and then started a career in child psychology, having studied psychology in Italy. In Caracas she established the first institute for this discipline. While returning from a visit to France, Catalina met Croizat. She described him as a person with a broad knowledge and interest - quite unlike other botanists who she generally found to be dull and uninteresting. She later amicably di-

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

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

Fig. 5. Welwitschia mirabilis in the Jardín (1988).

vorced from her husband as they had been separated for some time. She married Croizat and suggested that he leave his university position because there was a problem created by a demand that his articles be authored by another person. Catalina proposed that she support them through the development of a landscape gardening business. She made several trips to California to learn the practice. Having worked with various families in Caracas through her psychology institute she now had contacts for establishing her landscaping business. Catalina said she was initially reluctant to move to Coro, but she moved because of Croizat. But now she liked Coro even though it is a small town with limited commodities.

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

dates I was visiting, which shows that I was a bit clueless about such things at that time). We stayed with a Hungarian friend of Catalina for a couple of nights which gave us the opportunity to see some of Caracas (including a very nice restoration of a house belonging to Simón Bolívar or one of his colleagues, and a very nice bookstore where we were able to buy some natural history books) before catching a plane back to the US and then to New Zealand. Since that time I was unable to keep up regular contact with Catalina.

She passed away on July 29, 1997 and is buried along with Croizat. Last year it was reported in the newspaper El Nacional that the Jardín finally fell into such neglect that numerous plant collections from all over the world were lost, including the Welwitschia specimens. A recovery has been proposed. In February 2002 there was an exhibition of Croizat’s works, and a public event where presentations were made by Dr. José

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

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

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

The journey to Coro gave me a unique insight and understanding of the intellectual and practical challenges faced by Croizat in the development of his radical ideas on biogeography and evolution. Looking back at what he accomplished for panbiogeography, and the prominence that later came to his work, I am still amazed. For the most part, his leading contemporaries in evolutionary theory were established within the institutions of science and had resources that Croizat could only dream about. But through his extensive correspondence, Croizat was in surprisingly close contact with many of the influential biologists of his time.

That Catalina was also pivotal in changing the course of evolutionary biology through her total commitment to Croizat’s work is something I found beyond question. For me, the impact of their commitment extends well beyond their lifetimes in both space and time – as so it should.

Acknowledgements I am most appreciative to Jürg De Marmels, Ángel Viloria, and Mario Zunino for feedback on the draft article, and also to Ángel Viloria for illustrations of the dedication.

John Grehan Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, NY 14211-1193, USA. jgrehan@sciencebuff.org

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Una perspectiva latinoamericana de la biogeografía

BY R. WILLS FLOWERS

Una perspectiva latinoamericana de la biogeografía (1a. edición), 2003, Morrone JJ, Llorente Bousquets J, eds.

Universidad Nacional Autónoma de México, México, 307 pp.

ISBN 970-32-0498-8.

MXN $168 (USD 15.30).

Since the beginning of the 20th century, biogeography has been shaped by various “schools”. The dispersalist “First New York”, the vicariance “Second New York”, and the panbiogeographic “New Zealand” schools have each had their periods of prominence, and continue to generate both light and considerable heat among historical biogeographers. With the dawn of the 21st century, a new school has appeared that is giving a fresh perspective and direction to biogeographic research. Members of this school are from Latin America, and the focus of their research is the Neotropics and Mexico. Rather than promoting particular methodologies, the “Argentine-Mexican School” (named for the two countries where biogeographic workers have been the most active) is pragmatic, diverse, and focused on areas and biotas of the Neotropics that have been rather neglected by First World biogeographers.

In fact, South America has had an active cadre of native biogeographers for almost as long as the United States. For those wanting to learn more about the “Argentine-Mexican” biogeography school, and to read about recent developments in Neotropical biogeography, Una perspectiva latinoamericana de la biogeografía is an excellent introduction (with three articles in English, one in Portuguese, and 30 in Spanish, some reading knowledge of the latter is required). This volume is one of a series of books focusing on Latin American biogeography published since 2001 by the Uni-

versidad Nacional Autónoma de México. All were written primarily for a Latin American audience, with the goal of both publishing contemporary works and making some of the classic English biogeographical studies accessible to a Spanish-speaking audience.

Una perspectiva latinoamericana de la biogeografía is divided into three sections: history; theories, methods and concepts; and case studies. The historical papers include two essays on the works of Charles Lyell and Alfred Russel Wallace; both are intended to make these classic authors accessible to Spanish speakers but may be of limited interest to English speakers, who can readily find both the original works and innumerable English commentaries. Of greater interest are essays on Oswaldo Reig and Raúl Ringuelet, two Argentine biogeographers. Reig worked with South American mammals and Ringuelet pioneered the study of biogeographic subregions in the Southern Cone based on his studies of freshwater crustaceans and leeches.

Another essay evaluates the impact of Léon Croizat on Latin American biogeographers; this essay includes as appendices Spanish translations of two of Croizat’s later, hard-to-find, and relatively short publications.

The second section on theories, methods, and concepts covers topics from tracks to parsimony analysis of endemism to phylogeography. Most of the articles are intended to bring Spanish speakers up to speed on recent developments in the United States and Europe; thus, readers outside Latin America may find this section of limited value since much of the methodology has already been expounded and extensively discussed in English. I found one short article in English

by Nelson and Ladiges on geographic paralogy worthwhile because it actually includes a definition of geographic paralogy, something that is unaccountably lacking in many English articles on the subject (including the otherwise seminal Historical biogeography [Crisci et al. 2003]). In their article on biogeographical transitions, Ruggiero and Ezcurra provide a revealing graph of past climactic events, which shows that glaciations in the Southern Hemisphere began in the Eocene and, with a slight letup in the Miocene, have continued to the present. (Yet First World biogeographers continue to obsess over whether the much more

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recent Pleistocene glaciations were what shaped the biota of the Amazon.) In their article on phylogeography, Lanteri and Confalonieri cite two interesting cases in the Curculionidae (Coleoptera). One study showed that the boll weevil (Anthonomus grandis) in Argentina comprises two distinct populations: one identical to pest populations in the southern United States and one ancestral population related to Mexican boll weevils. In the other study of the genus Galapaganus, mitochondrial DNA shows that the age of the species on the Galapagos Islands predates the oldest extant island.

The third section on case studies was for me the most interesting part of this volume. While vertebrates are not ignored, most of the articles featured invertebrate or plant groups as subjects for analysis. There is a definite emphasis on the Mexican Transition Zone and on biogeographic subregions of Argentina and surrounding countries. Some of these studies (e.g., Morrone’s article on trichodactyline decapods) are updates or continuations of earlier studies published in English (e.g., Morrone & Lopretto 1994). Many authors follow the recommendations of Crisci et al. (2003) in their selections of methods of analysis. A number of papers begin with track analysis followed by area cladograms and parsimony analysis of endemicity.

Some notable contributions include Kohlmann and Wilkinson’s delineation, using distributions of dung beetles, of a biogeographic barrier in Costa Rica along the Río Grande de Tárcoles at the foot of the Talamanca mountain range. Interestingly, Croizat (1976) repeatedly cites nearby Punta Mala as a major biogeographic node. As an entomologist, I found the articles on tropical Andean Satyridae (Lepidoptera) by Viloria, a worldwide review of scorpion biogeography by Lourenço, a review of Neotropical Muscidae (Diptera) and a biogeographic study of the arthropods of the precordillera of Argentina by Roig-Juñent, Flores, and Mattoni of particular interest. All except the last are in English.

My most serious complaint about this book is its small type size and sans serif font, which makes reading difficult except in very well-lit conditions. Some of the distribution maps would be more legible in a larger size, but in some cases this is a result of reproduction of small originals. However, for anyone with an intermediate reading knowledge of Spanish (and a good reading glass), this volume offers a wealth of biogeographic information from a mega-diverse part of the world.

References Crisci JV, Katinas L, Posadas P. 2003. Historical biogeography: An introduction. Cambridge: Harvard University Press.

Croizat L. 1976. Biogeografía analítica y sintética (‘panbiogeografía’) de las Américas. Caracas: Biblioteca de la Academia de Cien-

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cias Físicas, Matemáticas y Naturales.

Morrone JJ, Lopretto EC. 1994. Distributional patterns of freshwater Decapoda (Crustacea: Malacostraca) in southern South America: A panbiogeographical approach. Journal of Biogeography 21: 97-109.

R. Wills Flowers Center for Biological Control, Florida A&M University, Tallahassee, FL 32307, USA. rflowers7@earthlink.net

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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 comprises the Amazon basin and the lowlands of southern Colombia, southern Venezuela, eastern Ecuador, northern and eastern Peru, eastern Bolivia, northern Brazil, and the Guiana. Different approaches (Cabrera & Willink 1973, Müller 1973) have been used to identify its limits. However, the concern for its biogeographic history is recent (Amorim 2001). Just a few years ago the first quantitative approaches using phylogenetic hypotheses were generated. Cracraft & Prum’s (1988) study gave the first steps to resolve the question about Amazonia’s origin, but Amorim’s (2001) work was the first to challenge the traditional view of Amazonia as a single unit (Cabrera & Willink 1973, Hooghiemstra 1997).

The earliest attempts to reconstruct the Amazonian biogeographic history were non-quantitative approaches based on narrative descriptions of those events that contributed to today’s diversity of species.

These are known as the Amazonian diversification hypotheses (Wallace 1852, Haffer 1969, for a revision see Haffer 1997 and Nores 2000). Although these approximations were very useful at that time because they opened the question about Amazonian diversity, since the development of quantitative biogeography approaches they have been severely criticized for two main reasons. First, because they are based on explicit or implicit a priori assumptions (Amorim 2001), and secondly because their statements could not be empirically tested (Cracraft & Prum 1988, Patton & da Silva 1998, Alexio 2002). Under an explicit biogeographic framework the first cladistic and quantitative analyzes began with Prum (1988) and Cracraft & Prum (1988).

Although Cracraft & Prum (1988) did not explicitly describe the methodology used to obtain their area cladogram, Brooks Parsimony Analysis (BPA) was the most frequent method used. After them, da Silva & Oren (1996), Bates et al. (1998), Ron (2000), and Racheli & Racheli (2003, 2004) used a modification of PAE (Cracraft, 1991) to generate their hypotheses of area relationships. Amorim (2001) and Hall & Harvey (2002) have proposed the latest and most recent area relationship hypotheses using phylogenetic and dis-

tributional data, using “cladistic biogeography” and BPA, respectively.

In spite of all these efforts, the origin and the biogeographic history of the Amazonian diversity is still in discussion. This is evident from the amazing number of papers dealing with the subject (Peres & Terborgh 1995, Haffer 1997, Hooghiemstra 1997, Marroig & Cerqueira 1997, Colinvaux et al. 2000, Bates & Demos 2001, Rose & Grainger 2003). From the biogeographic framework, however, these area relationship hypotheses have failed to document explicit historical events, like those events resulting from cladogenetic events, since seven of the 16 area cladograms were developed without phylogenetic information of the Amazonian taxa (Table 1). For example, Racheli (2004) has criticized the use of the Matrix Parsimony Representation technique used by Hall & Harvey (2002) to produce their combined area cladogram, questioning the use of the competing topologies as source data. Additionally most of these hypotheses have been formulated from a single taxonomic group.

It is obvious that the history of the area had to have affected all the Amazonian taxa that coexisted over the region, producing their speciation, extinction, or dispersion pattern(s).

The diversification hypotheses: The non-quantitative approaches There have been proposed more than 16 diversification hypotheses (Nores 2000), these hypotheses formulated a causation perspective for the Amazonian diversity - lacking any approach to quantitative analysis and therefore these hypotheses have had a strong narrative tradition. Nores (2000) presented an extensive review of these hypotheses. One of the first to highlight the diversity present in the Amazonian forests was Alfred Wallace (1852), after his voyage along the Amazonian Basin. His observations lead him to propose the first diversification hypothesis to the Amazonia, known now as “The Riverine Hypothesis”.

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Table 1. Main hypotheses of area relationship proposed for Amazonia. For each pattern the author’s name, taxa used, method and explanation suggested are shown. Be, Belem; BPA, Brooks parsimony analysis; CAT, Caatinga; CB, “cladistic biogeography”, no method specified; 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

Prum (1988) Prum (1988) Cracraft & Prum (1988) da Silva & Oren (1996) Bates et al. (1998) Patton et al. (2000) Ron (2000) Ron (2000) Amorin (2001) Hall & Harvey (2002) Hall & Harvey (2002) Marks (2002) Marks (2002) Racheli & Racheli (2003) Racheli & Racheli (2004) Racheli & Racheli (2004)

Birds Birds Birds Primates Birds Rodents and marsupials Squamata Anura Insects and primates Butterflies Vertebrates and butterflies Birds Birds Birds Butterflies Butterflies

Method BPA BPA BPA PAE PAE BPA PAE PAE CB PH MRP PH PH PAE PAE PAE

Area cladogram ((GU,(PA, BE)),(RO, IM, IN, NP)) (GU,((RO,(BE,PA)),(IM,(IN,NP)))) (SD,((CH,CA),(GU,((SD,BE_PA),(IM,(IN,NP)))))) (GU, (PA, BE)), ((RO, IN1), (IM, IN2, NP)) (GU, (IM, (IN, NP)), (BE, (PA1, (PA2, (RO)))) (GU, (IM, (IN, NP))), (RO, PA, BE) ((PA,(RO,(BE,GU))),(IN,NP)) (BE,(RO,(GU,(IN,NP)))) (((IM,(GU,BE)),(NP,IN)),(PA,(SD,CAT))) (GU,((RO,(BE,PA)),(IM,(IN,NP)))) ((RO,(BE,PA)),(GU,(IM,(IN,NP)))) ((GU,(IN,(NP,(CH,(CA,IM))))),(RO,(PA,SD))) (RO,((GU,(IN,(NP,(CH,(CA,IM))))),(PA,SD))) ((IM,(NP,IN)),(GU,((BE,PA2),(PA1,RO)))) ((GU,(BE,(PA2,(RO,PA1)))),(IM,(IN,NP))) (GU,((BE,(PA2,(RO,PA1))),(IM,(IN,NP))))

Process invoked IV* IV* Vicariance Vicariance IV* Vicariance Vicariance Vicariance Vicariance Vicariance Vicariance none explicit** none explicit** IV* IV* 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 there is no explicit reference about the process involved.

The principal argument is that the current biodiversity is the result of the vicariant events depicted from the formation of the Amazon River and its tributaries. All of the diversification hypotheses have presented a vicariance perspective of the Amazonian history; clearly the speciation is given by allopatric events caused by a variety of barriers to gene flow: rivers, lagoons, ecological gradients, or the sea level. Marroig & Cerqueira (1997) offered some of the phylogenetic and biological implications of the statements proposed by the most important diversification hypotheses: the Pleistocene refuge hypothesis (Haffer 1969), the riverine hypothesis (Wallace 1852, Sick 1967), and the gradient hypothesis (Endler 1977).

The quantitative approaches Quantitative analyses gained ground during the last few years since the impact and acceptance of the phylogeography research program (Lougheed et al.

1999, Ditchfield 2000, Patton et al. 2000, Marks et al.

2002, Costa 2003). Population genetics analyses, undoubtedly, have contributed to the “Amazonian cause”, for example, testing some predictions of the riverine diversification hypothesis (Patton & da Silva 1998, Lougheed et al. 1999, Alexio 2002). Nevertheless, most

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of these analyses have been developed to answer the speciation history of one or two particular taxonomic groups and not to explore the relationship between the areas.

Under an explicit biogeography framework, the area relationship for the Amazonia has been reconstructed eleven times (Table 1), but since some authors presented more than one, there are 16 proposed area cladograms. All these biogeographic studies stated that the history of Amazonia was driven mainly by vicariance events that have produced a hierarchical pattern (Moritz et al. 2000, Ron 2000, Amorim 2001, Bates 2001, Hall & Harvey 2002). Ron (2000) and Hall & Harvey (2002), based on the congruence between the history of the areas for taxonomically distant groups (butterflies, primates, birds, anurans, and lizards), presented a strong common history of vicariant isolation events proposing as the first event the disjunction between Guiana and the Atlantic Brazilian Forest. These studies also have shown some persistent relationships, for example a consistent Inambari-Napo group. The Guiana position, however, continues to be uncertain, its relationship with the so-called upper (Imeri and Napo) and lower (Inambari, Rondonia, Para, and Belem) Amazonia is inconsistent among the hypotheses and sometimes appears to be an independent unit. Additionally, it is still unclear what barriers were

responsible for the proposed vicariant patterns (Ron 2000, Hall & Harvey 2002) and the timing of the diversification. The controversy continues at this point, and several hypotheses have been proposed to explain that vicariant pattern (the diversification hypothesis) and although the Pleistocene Refuges model (Haffer 1969) has generated discussion, this is still accepted for some groups of animals and within the neontologists, to explain the diversification patterns (Brower 1994, Hall & Harvey 2002). One major problem that involves all of the above mentioned hypotheses of area relationship is that they were formulated to account for events of the late Pleistocene period, leaving out earlier periods (Cretaceous or late Tertiary) which molecular phylogenies of frogs, lizards, birds, and mammals suggest to be a critical time for the main events of speciation (Moritz et al. 2000, Ron 2000).

The most important contributions about the origin of the Amazonian biota probably have come from recent phylogeographic studies (Patton et al. 2000, Marks et al. 2002, Costa 2003). Marks et al. (2002), using the diversification pattern of Glyphorynchus spirurus, showed that some Amazonian haplotypes (Imeri population) are more closely related to Central America and Chocó populations than they are to other Amazonian taxa (Napo populations), a pattern that they documented to be common within the Amazonian birds (Marks et al. 2002, p. 164). Costa’s (2003) analysis of small mammals in Amazonia and the Brazilian Atlantic forest showed that these two regions are not exclusive in their fauna composition, and that the genetic similarity exhibited between the mammals in the two areas is often higher than the similarity within each of the areas. Although others works had already achieved a similar conclusion (Cracraft & Prum 1988, Amorim 2001), it is interesting to see how this result repeats when new taxa are studied, and how wrong was the idea of two different units (provinces or domains) that now appear to be a single biogeographic unit. Another conclusion produced by Costa’s (2003) work, which must be tested with strictly historical biogeographic approaches, predicts that a general area cladogram for the history of the Neotropical area is not likely and that a single vicariance model will not explain the speciation events. Although these works are focused on birds and mammals, they have produced new insights about the origin of the Amazonian biota and have placed new challenges to the biogeography research program. Given this scenario if we want to understand the biotic diversity of the Amazonia, and the reasons to explain why it supports that diversity we have to search for a reconstruction of the Amazonian history using all the distributional and phylogenetic information available, using an explicit cladistic method to account all the possible events that have taken place in Amazonia.

Acknowledgments We are deeply thankful to Isabel Sanmartín, Leonora Costa, Juan J. Morrone, Jim Patton, Paula Posadas, Carla Penz, André Freitas, Keith Brown Jr., Salvador Arias, Mario Quijano, and Claude Gascon for their constructive comments. The first author is much indebted to Isabel Sanmartín for her constant incentive and encouragement. All the members of the Laboratorio de Sistemática & Biogeografía are acknowledged for their individual efforts. Malte Ebach helped us as Editor. The financial support of the Vicerectoría de Investigaciones y extensión, UIS, is kindly acknowledged.

References Aleixo A. 2002. Molecular systematics and the role of the “Varzea”“Terra-Firme” ecotone in the diversification of Xiphorhynchus woodcreepers Aves: Dendrocolaptidae. The Auk 119: 621-640.

Amorim DS. 2001. Dos Amazonias. Introducción a la biogeografía en 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 for historical complexity and a vicariance model from a basic pattern of 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 the neotropical lowlands: An hypothesis based on raw distributions of Passerine birds. Journal of Biogeography 25: 783–793.

Bates JM, Demos T. 2001. Do we need to devalue Amazonia and other large tropical forest? Diversity and Distributions 7: 249-255.

Brower AVZ. 1994. Rapid morphological radiation and convergence among races of the butterfly Heliconius erato inferred from patterns of mitochondrial DNA evolution. Proceedings of the National Academy 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 and Neotropical plant communities on glacial time-scales: The failure of the aridity and refuge hypotheses. Quaternary Science Reviews 19: 141-169.

Costa, LP. 2003. The historical bridge between the Amazon and the Atlantic forest of Brazil: A study of molecular phylogeography with small mammals. Journal of Biogeography 30: 71-86.

Cracraft J. 1991. Patterns of diversification within continental biotas: Hierarchical congruence among the areas of endemism of Australian vertebrates. Australian Systematic Botany 4: 211-227.

Cracraft J, Prum RO. 1988. Patterns and processes of diversification: Speciation and historical congruence in some Neotropical birds. Evolution 42: 603-620.

Ditchfield AD. 2000. The comparative phylogeography of Neotropical mammals: Patterns of intraspecific mitochondrial DNA varia-

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tion among bats contrasted to nonvolant small mammals. Molecular 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 in Amazonia: An overview. Biodiversity and Conservation 6: 451-476.

Hall JPG, Harvey DJ. 2002. The phylogeography of Amazonia revisited: New evidence form Riodinid butterflies. Evolution. 56: 14891497.

Hooghiemstra H. 1997. Tropical rain forest versus savanna: Two sides of a precious medal? A comment. The Hague, pp. 31-43. The Netherlands Organization for Scientific Research.

Lougheed SC, Gascon C, Jones DA, Bogart JP, Boag PT. 1999.

Ridges and rivers: A test of competing hypotheses of Amazonian diversification using a dart-poison frog Epidobates femoralis. Proceedings of the Royal Society London 266: 829-1835.

Marks BD, Hackett SJ, Caparella AP. 2002. Historical relationship among Neotropical lowland forest areas of endemism as determined by mitochondrial DNA sequence variation within the Wedgebilled Woodcreeper Aves: Dendrocolaptidae: Glyphorynchus spirurus. Molecular Phylogenetics and Evolution 24: 153-167.

Marroig G, Cerqueira R. 1997. Plio-pleistocene south American history and the Amazon lagoon hypothesis: A piece in the puzzle of Amazonian diversification. Journal of Comparative Biology 2: 103-119.

Moritz C, Patton JL, Schneider CJ, Smith TB. 2000 Diversification of rainforest faunas: An integrated molecular approach. Annual Review of Ecology and Systematics 31: 533-563.

Müller P. 1973. The dispersal centers of vertebrates in the Neotropical realm. Biogeographica 2: 10-198.

Nores M. 2000. Species richness in the Amazonian bird fauna from an evolutionary perspective. Royal Australian Ornithologists Union 100: 419-430.

Patton JL, da Silva MNF. 1998. Rivers, refuges and ridges: The geography 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 Rio Juruá and the evolutionary and ecological diversification of Amazonia. Bulletin of the American Museum of Natural History 244: 1-306.

Peres CA, Terborgh JW. 1995. Amazonian nature reserves: An analysis of the defensibility status of existing conservation units and design criteria for the future. Conservation Biology 9: 34-46.

Prum RO. 1988. Historical relationships among avian forest areas of endemism in the Neotropics. Acta XIX Congressus Internationalis Ornithologici. Pp. 2563–2572. Ouellet H de. XIX Internat. Ottawa.

Racheli L. 2004 The nigthmare of the combination: Comments on matrix representation with parsimony and its first application in biogeography. Cladistics 20: 208-211.

Racheli L, Racheli T. 2003. Historical relationship of Amazonian areas of endemism based on raw distributions of parrots Psittacidae.

Tropical Zoology 16: 33-46.

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ships based on raw distributions of papilionid butterflies Lepidoptera: Papilionidae. Biological Journal of the Linnean Society 82: 345-357.

Ron S. 2000. Biogeographic area relationships of lowland Neotropical 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 variation in biodiversity in Peruvian Amazonia. Diversity and Distributions 9: 237-249.

Sick H. 1967. Rios e enchentes na Amazonica como obstaculo para a 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 analysis of endemicity in Amazonian biogeography: an example with primates. Biological Journal of the Linnean Society 39: 427–437.

Wallace AR. 1852. On the monkeys of the Amazon. Proceedings of the Zoological Society of London 20: 107-110.

Ivonne J. Garzón-Orduña Department of Biological Sciences, University of New Orleans, 2000 Lakeshore drive, New Orleans, LA, 70148, USA Daniel Rafael Miranda-Esquivel Laboratorio de Sistemática & Biogeografía, Escuela de Biología, Universidad Industrial de Santander, A. A. 678 Bucaramanga, Colombia.

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Ghosts of Gondwana

BY JOHN R. GREHAN

Ghosts of Gondwana, 2006, Gibbs GW. Craig Potton Publishing, Nelson, New Zealand.

232 pp.

ISBN 13: 978-1877333-48-4.

NZD 49.99 (USD 34.50).

George Gibbs faced no small challenge in writing Ghosts of Gondwana. New Zealand has long been a major center of attention for global biogeographic studies, and its origins and evolution were subject to intense debate over the last quarter century. Ghosts of Gondwana attempts to portray the origins and evolution of organisms in the context of New Zealand’s linear temporal sequence, so that different groups are discussed in each chapter or section according to their origin before or at various times following New Zealand’s geological isolation according to fossil, molecular, or dispersal criteria. In Ghosts of Gondwana Gibbs attempts to grapple with new biogeographic research present a balanced view on the historical arguments over the origins and evolution of the fauna and flora.

As noted in the introduction, this is not just another book describing the plants and animals of New Zealand, but a selection of examples that best illustrate the unique features of New Zealand life by drawing on biological and geological information to trace their history. In many respects Ghosts of Gondwana is very successful in its exploration of a broad range of issues and examples in a coherent and sequential picture of traditional theories about New Zealand’s biogeographic origins and evolution and some of the central issues of contention. The issues are arranged in five sections. Section I emphasizes New Zealand’s endemism, section II looks at the historical issues for biogeography, section III looks at various explanations for New Zealand’s life, section IV examines specific

groups or ecologies within New Zealand, and section V provides some final thoughts on the problem of distinguishing vicariance and dispersal. Each of the major issues and prominent examples are complemented by extensive illustrations, beginning with a geological timescale that includes an overview of dominant lifeforms and some major historical geological events, and followed by various focal inserts summarizing the pertinent geographic, systematic and biological features or individual organisms or various groups.

Ghosts of Gondwana represents a significant signpost in the history of biogeographic and evolutionary research in New Zealand. During the mid 20th century an almost universally accepted history of the New Zealand biota was established by Sir Charles Fleming, who used geological stratigraphy to construct a center of origin and dispersal model that divided organisms into ancient endemics (later to become Gondwana elements) and later colonists from various centers of origin or derived from New Zealand itself as a center of origin within the region. Only with rare exceptions, such as the pioneering New Zealand palynologist Lucy Cranwell (1962, 1964), was there any early willingness to consider geological influences in the evolution of New Zealand plants and animals. The scientific exploration of New Zealand’s evolution is characterized in Ghosts of Gondwana as a search for criteria to distinguish New Zealand’s original biota before separation from the hypothetical super-continent of Gondwana, and those that subsequently dispersed to or from New Zealand after its geological and geographic isolation about 80 million yeas ago.

The division of New Zealand’s biota into ancient endemics and later dispersal was established as the foundation for New Zealand’s evolutionary history by Sir Charles Fleming. With popular acceptance of continental drift and plate tectonics the ancient endemics became the icons or New Zealand’s original Gondwana inheritance. But all the elements of the biota continued to be seen as the product of dispersal either before or after isolation and arriving from various centers of origin in the local region or other parts of the globe.

Life was seen as being continually on the move as various newly evolved groups moved along one or other

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dispersal pathway to sometimes finally even end even up at the most isolated locations such as New Zealand. It was not until the 1980’s that Fleming’s evolutionary model was challenged by local support for Léon Croizat’s panbiogeography. It would be an understatement to say that this was received with strong opposition from the scientific establishment. George Gibbs found himself in the middle of the debate through his role as the graduate advisor for Robin Craw, Ian Henderson, and John Grehan, who were finding panbiogeography to provide a much more scientifically defensible approach to New Zealand’s biological evolution. While taking a critically cautious path in the face of some very radical propositions about New Zealand history, George Gibbs also found the opportunity to apply panbiogeographic methods to his interest in the primitive micropterigid moths, and even explore the possibility their biogeographic origin was linked to a Pacific geo-history (Gibbs, 1983) and that their New Zealand distributions may reflect patterns dating back to the Cretaceous (Gibbs, 1990). Following a very successful symposium on panbiogeographic and non-panbiogeographic perspectives in 1989 (Matthews 1990), the panbiogeographic synthesis failed to receive the necessary support from New Zealand’s research and funding institutions to continue as an active research program and there has now been no endemic panbiogeographic research program for many years.

Through Ghosts of Gondwana, one is in a position to see how various animal and plant groups are interpreted through theoretical interpretations of fossil and molecular evidence. The principle deficiency of this approach lies in what is left out. The panbiogeographic analyses of New Zealand are presented as a largely historical event as Ghosts of Gondwana looks to molecular systematics as the fuel for the “latest revolution in biogeographic thinking” that can provide a direct measure of time in biogeography, whereas Croizat’s panbiogeography could only analyze space. The development of routine methods of sequencing the DNA molecule which “contains information on the evolutionary history of every living organism” is characterized as the most exciting advance in recent years, and molecular technology is presented as a biogeographical revolution generating “new” interpretations of how and where the present fauna and flora reached New Zealand. In one sense these characterizations are true, but constructing a phylogenetic tree requires appropriate concepts of character homology to separate primitive and derived character states involving only four base combinations that provide no trace of individual replacements at each site. Ghosts of Gondwana does not show any empirical evidence for molecular similarity being necessarily any more precise about phylogenetic relationships than morphology so it is not possible to discern whether the molecular

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emphasis and historical relegation of panbiogeography is scientifically justified.

Throughout Ghosts of Gondwana the molecular approach is used to generate divergence estimates for taxa where fossils are not available. Even though the first fossil appearances are acknowledged to be potentially be unreliable and that molecular biology only gives “approximate hypothetical answers” these concerns are overruled by “a cascade of studies that offer very persuasive arguments for efficacy for trans-oceanic dispersal”. Molecular claims for over-water dispersal when divergence estimates post-date earlier geological events are accepted at face value, even though molecular divergences are in reality minimal estimates calibrated by fossils representing minimal ages of fossilization. It is noted that a discrepancy of 10 million years either way may not matter much, but if the discrepancy between age of origin and age of fossilization may be many times greater, molecular divergence cannot be assumed to falsify earlier geological events (Heads 2005). A few examples will serve to illustrate the perspective presented in Ghosts of Gondwana and illustrate the methodological and conceptual boundaries of what is in many respects the most comprehensive popular book on New Zealand’s origin and evolution in thirty years.

Chapter II presents recent DNA research to suggest ancestors of the short-tailed bat, Mystacina tuberculata, were separated from ancestors of noctilionoid bats in South America about 54 Ma, after which they became widespread across Antarctica and Australia before they “probably reached New Zealand by flying across the Tasman sea”. The empirical evidence for this South American center of origin and the dispersal story involving Antarctica and Australia is not identified. The molecular center of origin story ignores the biogeographic pattern that connects New Zealand and Australia (through fossil representatives) directly with northern South America and Central America directly over the Pacific (Fig. 1). This is a pattern found in many other animal and plant groups that do not conform to the expectations of a Gondwanic origin, but predicts a Pacific geohistory responsible for this type of biogeographic pattern as well as the origin of allocthonous terranes around the Pacific margins (Craw et al. 1999).

The origin of southern beeches (Nothofagus) in New Zealand through geological isolation is described as a theory long supported by the strong belief that the trees are incapable of long-distance dispersal.

“Modern” molecular studies (Knapp et al. 2005) are said to now suggest that probably all of New Zealand’s modern Nothofagus species arrived from Australia, and replaced the previous species within the last 35 million years. Knapp et al. (2005) claim to have “unequivocal molecular clock evidence” that trans-Tasman Sea

Fig. 1. Biogeography of the New Zealand short-tailed bat and its fossil relative in Australia as red polygons, and their probably nearest relatives the South American Mormoopidae and Noctilionidae (blue) (Kennedy et al. 1999, Koopman 1984).

distributions can only be explained by long-distance dispersal and that the absence of Lophozonia and Fuscospora pollen types in the Cretaceous of New Zealand is evidence for Tertiary dispersals to New Zealand.

The dates were calibrated by fossil pollen divergence of for subgenera Lophozonia, Nothofagus, and Fuscospora at 75 million years ago along with a “minimum” divergence between N. cunninghammii and N. moorei based on “intermediate” fossils and others that “closely resembled” N. moorei. There is no indication of why the 75 million year fossil record should represent the upper limits of this group. A calibration at 65 million years for sister taxa between New Zealand and Australia produced “unrealistic age estimates for the basal nodes”, which might actually suggest that the molecular clock model is itself unrealistic. This problem aside, the dispersal speculations would have been enhanced by reference to the vicariant main massings of the four Nothofagus subgenera (Heads 2006). These patterns (Fig. 2) are concordant with the distributions of each subgenus having their origins in a process of vicariant differentiation well into the Mesozoic, and the patterns may well represent an independent ‘test’ of the molecular clock theories.

Dispersal directionality is also a common feature of molecular biogeography. For example, the basal position of the New Zealand kauri (Agathis astralis) is seen as evidence that its relatives dispersed out of New Zealand. Similarly, Metrosideros has been “shown” by Wright et al. (2000) to have dispersed out of New Zealand to reach a number of Pacific islands as far away as Hawaii. In neither case is this dispersal demonstrated. Rather, each represents an application of Hennig’s progression rule that assumes derived taxa represent more recent dispersal events so the distribution of primitive and derived groups tracks the path of dispersal. This purely theoretical Darwinian model of evolution ignores the possibility that the same pat-

tern may arise from sequential differentiation of a widespread ancestor. This is particularly pertinent for Metrosideros, which comprises three largely vicariant (exception of a node at Fiji) taxa involving New Zealand and the Pacific (Fig. 3). If these tracks are the result of three serendipitous chance dispersal events, one is confronted with explaining just how each manage not only to jump across the ocean, but to do so in such a way as to end up on only a few islands (and not other islands or mainland areas) without overlapping each other than one case in Fiji, and also track

Fig. 2. Biogeography of Nothofagus main massings.

Large text – extant species, small text – fossil pollenleaves-cupules (from Heads 2006).

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

the same path found in other organisms such as the Tahiti-Hawaii connection that is also found in some Pacific shorefishes (Heads 1983) or find their limits on Bonin Island – another classic biogeographic boundary in the Pacific. Gibbs’ suggestion of a very recent origin for one lineage scarcely varies by more than one nucleotide may be true or it shows that for this lineage there has not been much in the way of molecular differentiation (i.e. effectively this means the ‘clock’ slowed down or is in stasis).

The treatment of Hebe biogeography provides a global example of the sometimes too limited scope of comparison between different current biogeographic frameworks and their bearing on the origin of the New Zealand biota in Ghosts of Gondwana. A molecular phylogeny by Wagstaff et al. (2002) is said to show that the Hebe ancestor reached New Zealand about 10 million years ago. Their molecular phylogeny is problematic in that it does not always correspond to patterns of relationship derived from morphological analysis, yet they presume that the molecular results are somehow more correct. They note that the earliest fossil appearance is Pliocene for Hebe, mid-Miocene for Scrophulariaceae, and mid-Eocene for Lamiales, and admit that their divergence estimates are “crude” and that older fossils could be found in the future. But they then claim that it would be inconsistent with the current fossil record to assume a Gondwana or earlier divergence even though the earliest divergence of Hebe is not constrained by the current fossil record other than a pre-existing fiat on the part of the authors. Gibbs cites the study by Wagstaff et al. (2002) as evidence for dispersal away from a New Zealand center of ori-

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gin, but again this dispersal is read into the relationships based on Hennig’s progression rule – a purely theoretical construction based on one of the many conflicting theoretical criteria for defining the theoretical center of origin. Whatever position one might take on the molecular study, the Hebe example would have been greatly enhanced by referencing and illustrating the study by Heads (1994) showing the nodal position of New Zealand with respect the four subgenera (Fig. 4), the similarity of Alpine fault disconnections shared between Hebe, Leonohebe, Nothofagus, and Coprosma as a tectonic correlation potentially falsifying 10 million year molecular divergence as an underestimate.

Spatial analysis, acknowledged to be so important in Croizat’s panbiogeography, represents the major absent methodology in Ghosts of Gondwana, where spatial correlation is briefly illustrated for the alpine fault translocation. Fossils from the mid-Miocene Lake Manuherikia of inland Otago are cited as a evidence for reconstructing the forest communities that grew around its shores. This example could have been greatly enhanced by reference to the living ring species formations in Otago that are spatially correlated with the shores of this progressively declining lake and illustrating the integrated biogeography of fossil and living members of the New Zealand biota (Heads 1990).

Further geological correlations may have enhanced the book by illustrating the extensive biogeographic arcs and their correlation with zones of tectonic activity such as plate and terrane margins, fracture zones, and belts of granitic intrusion as well as the role of uplift transforming mid-Tertiary lowland-coastal communities (Craw 1989, Heads 1990).

It is somewhat surprising to see that biogeographic maps are noticeably absent from a book with biogeography as its central theme. The global context of New Zealand is represented by several geohistorical maps, three geographic maps, and one showing the distribution of tuataras and their fossil relatives. Within New Zealand there are two maps illustrating the al-

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 biogeographic maps, Ghosts of Gondwana looks to theories from historical geology, ecology, and phylogeny to obtain answers on the origins of New Zealand’s terrestrial plants and animals. Since New Zealand biogeography has been so intensively studied by both Darwinian and nonDarwinian research programs, the inclusion of biogeographic maps could help the reader evaluate competing theories about the origin of New Zealand’s plants and animals. For example, the map illustrating the location of fossil tuatara relatives (Sphenodontia), in all continents except Antarctica does not show that the closest relatives appear to be found only in the Jurassic and early Cretaceous of Mexico and the western United States (Fig. 5). This is a pattern again clearly incongruent with the Wegenerian model of Gondwana requiring a permanent Pacific Ocean.

According to Ghosts of Gondwana, the apparent inability of the giant carnivorous land-snails (Rhytidae) to travel over oceans along with their presence on parts of Gondwana (Africa, Madagascar, eastern Australia) suggest they could represent a Gondwana founder in New Zealand. This is contrasted with their presence also on “oceanic islands” such as the Seychelles, Indonesia and Melanesia and this is said to imply “a capacity for trans-oceanic dispersal” (Fig. 6). The alternative possibility, that such “oceanic” islands are either not oceanic in origin, or they inherited a non-oceanic component from former non-oceanic landscapes is not considered. These possibilities, demonstrated for the Galapagos and Fiji (Grehan 2000, Heads 2006), suggest the label ‘oceanic’ is not necessarily informative for biogeographic analysis. The real question, perhaps never really addressed in this book, is why should Gondwanic landsnails, distributed over former parts of Gondwana as have a completely different biogeographic orientation (Indian Ocean) from the Pacific oriented tuatara, frogs and southern beeches that are also distributed over former parts of Gondwana and supposedly Gondwanan in origin? This kind of question is extensively addressed in panbiogeographic approaches to New Zealand, so their omission from this book is problematic for achieving a balanced overview of its biogeography and evolution.

A biogeographic correlation between New Zealand’s biota and its geological history is addressed by Ghosts of Gondwana in general reference to Wegener’s model of Gondwana as a unified geological entity. The existence of accreted Pacific terranes is acknowledged along with the possibility of former Pacific land, but their biogeographic significance is discounted by the “view of tectonic geologists” that such land would have been “insignificant, perhaps in the form of island arcs” and larger (how much larger?) terrestrial models are ridiculed by “mainstream geologists”. These

opinions raise the very important question of whether such island arcs (or any other mobile units such as micro-continents), however small, were also necessarily biogeographically insignificant. Ghosts of Gondwana presents a very different view on the role of geology in determining biogeographic interpretations to that of Gibbs (1983) where it was the evidence of biological distributions represented by the Micropterigidae that were seen to suggest that both New Zealand and New Caledonia are composite areas comprising elements from the Pacific as well as from Australia (Gondwana). Gibbs (1983) noted that the New Zealand micropterigids cannot be interpreted on the assumption that the present land mass is nothing more than a fragment of eastern Australia, yet in Ghosts of Gondwana that is what New Zealand has largely become in terms of its biogeographic evolution. In view of the extensive and diverse Pacific based distributions (including that of the Micropterigidae), the historical role of the Pacific in the ‘Gondwana’ biota may be far less resolved than implied by Ghosts of Gondwana. Because Wegener’s model of Gondwana (that proposes a permanent Pacific ocean) is accepted as an empirical reality, any phylogenetic relationship linking New Zealand closer to Australia than South America is nec-

Fig. 5. Biogeography of tuatara and closest fossil relatives (from Reynoso and Clark 1998).

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

· Biogeografía 1 · January, 2007 · Page 25 ·

essarily classified as over water dispersal because the classical breakup sequence requires New Zealand to have separated earlier than Australia and South America. The alternative possibility, that trans-Tasman patterns are derived from a Pacific accretion pre-dating New Zealand’s geological isolation remains a critically important biogeographic question to be addressed in the popular literature.

According to Ghost of Gondwana, molecular studies “can tell us the location of ancestral groups, when the founders arrived, and how rapidly they diversified”.

But this is not quite the whole story. The location of ancestral groups is not derived from molecules, but a theoretical speculation, proposed by Darwin, that vicariant distributions evolve from geographically narrow centers from which they disperse according to their individual means of dispersal. The theorized center of origin is located according to one or more sometimes contradictory theoretical criteria. Molecular studies usually use the location of the basal lineage, the oldest fossil, or area of greatest taxonomic diversity to denote the theorized center of origin. None of these criteria empirically demonstrate anything other than records of past or present distribution. The timing of ‘founders’ is also problematic as it represents a theoretical extrapolation of the fossil record. Since fossils represent minimal age of fossilization, any clock-like model can also only predict minimal dates of divergence.

In practice molecular dispersalists treat the fossil record as more or less the actual history of a group and molecular divergences are therefore said to falsify earlier events if such events predate the divergence estimate.

Such claims are methodologically invalid; for all that they are extremely popular (Heads 2005).

Molecular dispersalism relies on theory (e.g. models of molecular evolution, molecular substitution, and fossil calibration), whereas panbiogeography relies on the facts of distribution that are accepted by everyone, including molecular biogeographers. While Ghosts of Gondwana treats the molecular model as an independent ‘test’ (of what, is not quite clear), so too is the spatial analysis that has generated the facts of New Zealand’s biogeographic structure (its composite biogeography, its correlation with accretion and plate tectonics, parallel arcs, etc.). Ghosts of Gondwana’s characterization of panbiogeography as a spatial analysis lacking the element of time is also incorrect since the method provides a temporal estimate through the spatial correlation of distributions and tectonic structure. Geological correlations sometimes predict ages older than the oldest fossil and so provide a potential falsifier of the minimal estimates predicted through fossils and the molecular-fossil clock.

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

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lar clock speculations, and double invasions, Ghosts of Gondwana seems somewhat reminiscent of Sir Charles Fleming’s (1979) Geological history of New Zealand and its life dressed up in molecular technology. Since there is such an overwhelming focus on molecular clock theory as both a conceptual and technical advance over panbiogeography and other biogeographic approaches, the book seems to inevitably falls short of the author’s goal of providing an overall balanced description of current biogeographic research on New Zealand’s evolution. But these concerns aside, the book does stand out as a unique and significant contribution to the popularization of New Zealand biogeography and evolution. Ghosts of Gondwana covers a tremendous range of examples and stands out in stark contrast to other major works supporting a traditionalist view on New Zealand evolution by at least acknowledging the existence of an alternative. Because so many organisms are discussed, and so many questions are raised, it will make an excellent student textbook (indeed it is already being used for the course New Zealand Flora & Fauna at Victoria University of Wellington) if students are encouraged to explore the biogeographic literature in general. Only in this way will the author’s goals, and real potential and value of Ghosts of Gondwana, be achieved.

References Cranwell LM. 1962. Endemism and isolation in the Three Kings Islands, New Zealand - with notes on pollen and spore types of the endemics. Records of the Auckland Institute and Museum 5: 215-232.

Cranwell LM. 1964. Ancient Pacific floras. University of Hawaii Press, Honolulu.

Craw RC. 1989. Continuing the synthesis between panbiogeography, phylogenetic systematics and geology as illustrated by empirical studies on the biogeography of New Zealand and Chatham Islands. Systematic Zoology 37: 291-310.

Craw RC, Grehan JR, Heads MJ. 1999. Panbiogeography: Tracking the history of life. Oxford University Press, New York.

Emberton KC. 1990. Acavid land snails of Madagascar : Subgeneric revision based on published data (Gastropoda: Pulmonata: Stylomatophora). Proceedings of the Academy of Natural Sciences of Philadelphia 142: 101-117.

Fleming CA. 1979. The geological history of New Zealand and its life. 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 primitive Lepidoptera 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. Biological Journal 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 complex (Scrophulariaceae): Leonohebe and Chionohebe. Candollea 49: 81-119.

Heads MJ. 2005. Dating nodes on molecular phylogenies: A critique of molecular biogeography. Cladistics 21: 62-78.

Heads MJ. 2006. Panbiogeography of Nothofagus (Nothofagaceae): Analysis of the main species massings. Journal of Biogeography 33: 1066-1075.

Heads MJ. 2006. Seed plants of Fiji: An ecological analysis. Biological 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 and the 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 dispersal of Nothofagus (southern beech). PLoS Biology 3: 38-43.

Koopman KF. 1984. Orders and families of recent mammals of the world. John Wiley & Sons, Inc. New York.

Matthews C. 1990. Panbiogeography special issue. New Zealand Journal of Zoology 16: 1-815.

Reynoso VH, Clark JM. 1998. A dwarf Sphenodontian from the Jurassic La Boca Formation of Mexico. Journal of Vertebrate Paleontology 18: 333-339.

Van Bruggen AC. 1998 Gondwanaland connections in the terrestrial molluscs of Africa and Australia. Journal of the Malacological Society of Australia 4: 215-222.

Wagstaff SJ, Bayly MJ, Garnock-Jones PJ, Albach DC. 2002. Classification, 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 evolution of Metrosideros subg. Metrosideros (Myrtaceae) inferred from nrDNA. Proceedings of the National Academy of Science, USA 97(8): 4118-4123.

John Grehan Buffalo Museum of Science, 1020 Humboldt Parkway, Buffalo, NY 14211-1193, USA. jgrehan@sciencebuff.org

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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 and biogeographic ideas of the German physician, writer, and natural philosopher Julius Minding (November 8, 1808 - September 7, 1850). His book entitled “Über die geographische Verteilung der Säugethiere” (Minding 1829) played a rather important role in the development of regional approach of zoogeography.

A small book by Julius Minding of only 104 pages entitled “On geographic subdivision of mammals” (Minding 1829) (Fig. 1) is deposited in the library of the Zoological Institute of the Russian Academy of Sciences in St. Petersburg. Strangely enough, this book is not listed in data bases of the largest libraries of the world (for instance, The Library of Congress, The British Library, etc.) though it result Engelmann (1846) and Agassiz (1852). Neither is it listed by Wood (1931).

Carus (1872) mentions Minding only in passing, when he cites another book (Minding 1832). Hofsten (1916) writes that Minding explains an origin of the disjunct faunas by the action of “creative force” [Schöpfungskraft] “…which aspires everywhere to formation of similar organisms which however there are only analogous and not absolutely similar essences under influence of local conditions in the removed districts” [“J. Minding… nahm seine Zuflucht zu einer “Schöpfungskraft”, die überall zur Bildung gleichartiger Organismen strebt, von den lokalen Verhältnissen aber derart beeinflußt wird, daß in entfernten Gegenden nur analoge 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 in question.

The major part of Minding’s book (Minding 1829, pp. 46-104) borrows extensive “Tables of geographical subdivision of mammals” [“Tabellen über geographische Vertheilung der Säugethiere”] (Fig. 2). In tables on the basis of added data (Illiger 1812) Minding presents a list of 1230 Mammalia species for the basic regions of the Earth (Europe, northern and southern Asia, Africa, North and South America, Australia).

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

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geographical views are presented. He considers that taking into account physical geography alone is not sufficient for “natural” delimitation of continents. Of equal importance is geographical distribution of organisms, which closely interact with environmental conditions. For instance, the Sahara and Libian deserts, rather than the Mediterranean Sea, should be considered as the border between Europe and Africa [“Andern Theils möchen auch die alten Annahmen der Scheide zwischen Europa und Afrika verschiedene geographische und geognostische Ansichten gegen sich haben, da Binnenmeere, wie das Mittelländische, wirklich mehr alsgrosse Flussbetten, deren Gebiet sich nach beiden Seiten ihres Laufes erstreckt, zu betrachten wären; eine Annahme, in deren Folge die eigentliche Grenze zwischen Europa und Afrika durch die saharische und libische Wüste gebildet würde”] (Minding 1829, p. 5).

On the basis of the subdivision of the Earth into the Eastern and Western hemispheres (rather than into the Northern and Southern, as generally accepted nowadays), Minding proposed the following original division of the Earth’s surface: the Northeastern land [Festland, the firm ground] (30-78° N, 8-190° E) including Europe and North Asia; the Northwestern land (to the north of 23° N and 190–360° E) including North America; 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, and Australia. The “Festlands” established by Minding, although Minding himself does nor use the term “region”, obviously correspond to faunistic regions.

Minding did not consider rivers, narrow straits, etc. as barriers to species distribution, but referred to them as conditions (like climate and soil) promoting expansion of species. [“Flüsse sind so wenig als schmale Meerengen für Thierscheiden anzusehen; im Gegentheil pflegen sich verschiedene Arten gern längs ihrem Laufe zu beiden Seiten aufzuhalten, was wohl in der Gleichmässigkeit des Bodens und Climas seinen Grund haben mag” (Minding 1829, p. 11)].

Minding is consistent in his belief in the permanency of continents and oceans, rejecting the concept

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

of multiple “land bridges”. “The assumption that land masses were connected in the past, although extensively argued for, especially by earlier authors, seems to be so highly conjectural and inadequate, that no further arguments are needed to disprove the statement that such bridges may have ever existed. – A more precise comparison suggests that such assumptions are needless; clear differences between distinct land masses are as ancient as the contemporary pattern of the Earth’s surface, which has been affected only by recent revolutionary changes”. [“Die Annahme verbindender, untergegangener Länder, zu der so Viele, insbesondere in frühener Zeiten ihre Zuflucht nehmen zu müssen glaubten, ist so hypothetisch und zugleich so unzureichend, dass es keiner Beweise wider die Möglichkeit solcher Existenzen bedarf. – Eine genauere Vergleichung führt auch zugleich auf das Unnöthige derselben hin, indem die strengen Verschiedenheiten getrennter Länder mehr auf eine Trennung hindeuten, die so alt ist, als die jetzige Gestalt der Erdoberfläche, denn auf eine, erst durch spätere Revolutionen aufgehobene Verbindung”] (Minding 1829, p. 11).

Minding suggests that there might have been two or more centers of Creation for each species. He considers 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”.

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

Unlike Buffon (1776), Minding suggested that faunas 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 way that depending on the environmental conditions either similar or different forms populated different continents at any given period (Minding 1829, p. 11).

Summarizing what has been said above, one clearly sees that Minding’s (1829) book is the second greatest, after C. Illiger’s works (1811, 1812), contribution to the regional approach in zoogeography and therefore is worth referring to as a classical treatise on biogeography. Surprisingly, personal information about J. Minding is hardly available. Some data about him can be found in the free Wikipedia encyclopaedia (http://de.wikipedia.org/wiki/Julius_Minding), which, without mentioning the book (Minding 1829) in question, says the following: “Julius Minding was born on November 8, 1808 in Breslau, Germany (Wroclaw, as it now called, Poland).

In 1833 he graduated from Berlin University with his doctor of medicine degree [Dr. Med.] after submitting his thesis ‘De vitæ functionum perturbationibus notiones generales. Dissertatio inauguralis pathologica, etc.’ At that time he entered private practice and simultaneously started his literary work (for instance, he translated ‘Frithjofs-Saga’ by Esaias Tegnér from Swedish into German). Later, he involved himself into a risky commercial venture. Being on the verge of bankruptcy he moved to New York in 1850. There he and his friend established a medical bureau. This venture also failed. On September 1, 1850, J. Minding committed suicide” (http://de.wikipedia.org/wiki/Julius_ Minding).

Since J. Minding is undoubtedly an influential figure in biogeography, further studies of his biography and development of his expertise are needed. Of particular interest is the origin of his biogeographic views.

Acknowledgements I am grateful to Drs. Igor Smirnov and Sophia Stepanjants (both at the Zoological Institute, Russian Academy of Sciences) for providing much literature.

This paper is partly supported by the grant of the Far

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Eastern Branch, Russian Academy of Sciences.

References Agassiz JLR. 1852. Bibliographia zoologiæ et geologiæ. A general catalogue of all books, tracts, and memoirs on zoology and geology. Vol. 3. London: Print. for the Ray Soc. Buffon GLL. Comte de. 1776. Histoire naturelle, générale et particulière, etc. Supplément. T. 3 : …servant de suite à l’histoire des animaux quadripèdes. Paris, De l’Impr. Royale.

Carus JV. 1872. Geschichte der Zoologie bis auf Joh. Müller und Charl. Darwin. München, R. Oldenbourg.

Engelmann W. 1846. Bibliotheca historico-naturalis. Verzeichniss der Bücher über Naturgeschichte, welche in Deutschland, Scandinavien, Holland, England, Frankreich, Italien und Spanien in den Jahren 1700-1846 erschienen sind. Bd 1. Bücherkunde.

Hülfsmittel. Allgemeine Schriften. Vergleichende Abatomie und Physiologie. Zoologie. Palaeontologie. Mit einem Namen- und Sachregister. Leipzig: W. Engelmann.

Illiger JKW. 1811. Prodromus systematis mammalium et avium; additis terminis zoographicis utriusque classis, eorumque versione germanica. Berolini: C. Salfeld.

Illiger JKW. 1812. Überblick der Säugethiere nach ihrer Vertheilung über die Welttheile. Abhandlungen der Königliche Akademie der Wissenschaften zu Berlin. Physische Klasse für 1804-1811: 39-160.

Menzbir MA. 1882. Ornithological geography of European Russia. Pt. 1. Moscow: Imperial Moscow University [in Russian].

Minding J. 1829. Über die geographische Verteilung der Säugethiere. Berlin: Anslinsche Buchhandlung.

Minding J. 1832. Lehrbuch der Naturgeschichte der Fische. Berlin: A. Rücker.

Wood CA. 1931. An introduction to the literature of vertebrate zoology: Based chiefly on the titles in the Blacker Library of Zoology and other libraries of McGill University, Montreal. Oxford: Oxford University Press.

Alexander I. Kafanov A.V. Zhirmunsky Institute of Marine Biology, Far Eastern Branch, Russian Academy of Sciences, Vladivostok, 690041, Russia kafanov@mail.primorye.ru

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Una herramienta automatizada para realizar análisis panbiogeográficos POR CAMILO ANDRÉS ROJAS PARRA En la actualidad, la geoinformática, el creciente desarrollo de algoritmos o secuencias automatizadas de instrucciones y la tendencia a incorporar tecnologías modernas, como los Sistemas de Información Geográfica (SIG), han motivado a que los biólogos se enfrenten con el problema de representar en un mapa los patrones espacio-temporales de la biodiversidad (Morrone 2000, Tikunov 2002). Uno de los métodos que involucra principios, técnicas y tecnologías a nivel geográfico e informático es el método de la panbiogeografía. De acuerdo con Grehan (2001), la práctica del método de la panbiogeografía en lugares como América Latina, el suroeste del Pacífico y Europa se ha extendido desde la década de los 1990’s, no solo porque algunos autores, como Morrone y Crisci, han adoptado el método panbiogeográfico para integrarlo con los análisis biogeográficos cladísticos, sino también porque el método panbiogeográfico involucra el aspecto geográfico dentro del análisis de la evolución, parámetro que en gran parte de los análisis cladísticos se reduce a las ramas terminales de los cladogramas, construidos a partir de análisis entre caracteres biológicos.

Por los motivos anteriores se planteó el reto de formalizar una aproximación al método de la panbiogeografía desde un punto de vista informático, desarrollando una herramienta para crear trazos individuales, en lenguaje de programación AVENUE® para ArcView GIS 3.2® (ESRI 1996).

Antecedentes El uso en la biología de algoritmos como modelos de predicción de patrones espaciales de la biodiversidad, ha adquirido gran importancia en los últimos años para probar hipótesis biogeográficas, mejorar atlas florísticos y faunísticos, y definir áreas prioritarias para la conservación (Margules y Austin 1994, Hausser 1995, Kienast et al. 1996, 1998, Mourell y Ezcurra 1996). La mayoría de estos modelos se basan en el análisis de la relación especies-ambiente, asunto central de la bio-

geografía y la ecología (McArthur 1972, Myers y Giller 1988, Espinosa et al. 1993, Arita y Rodríguez 2002, Llorente et al. 2002), que permiten predecir la organización u ordenamiento espacial de los individuos y ayuda a tener mayores parámetros para el conocimiento sobre la biodiversidad. Los ejemplos más importantes del uso de algoritmos en la biología están relacionados principalmente con los modelos de predicción de los patrones espaciales, que involucran la instrumentación de los Sistemas de Información Geográfica (SIG) y el uso y desarrollo de algoritmos genéticos como el COVER (Moeur 1985), BIOCLIM (Nix 1986), GARP (Genetic Algorithm for Rule-set Prediction) (Stockwell y Noble 1991, Stockwell 1999, Gouch y Rushton 2000, Murguía y Rojas 2001, Navarro et al. 2003) y GAP Analysis (Scout et al. 1993), entre otros.

El problema del árbol de extensión mínima La base que fundamenta la instrumentación del método panbiogeográfico es el trazo individual, que extrapolado a un plano matemático es un conjunto mínimo de enlaces de N que conectan todos los nodos en A y por lo tanto al menos un árbol de expansión puede ser encontrado en un grafo G. El árbol mínimo de expansión denotado por T* es un árbol de expansión cuyo peso total de todo los enlaces es mínimo o “árbol de extensión mínima” (Campos 2002, Milková 2004). Es decir:

T * = min ∑ wij t

eij εE

Formalmente, un grafo G consiste en dos conjuntos finitos N y A. N es el conjunto de elementos del grafo, también denominado vértices o nodos. A es el conjunto de arcos, que son las conexiones que se encargan de relacionar los nodos para formar el grafo.

Los arcos también son llamados aristas o líneas. Los nodos suelen usarse para representar objetos y los arcos para representar la relación entre ellos. Cada enla-

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

Luego, para que un árbol de expansión sea mínimo 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 se resuelven acotados por un polinomio, se desarrollaron para solucionar el problema del árbol de extensión mínima entre los que se encuentran los algoritmos de Prim, Kruskal, Dijisktra y Solin (Milková 2004).

El algoritmo de Prim El algoritmo de Prim es una de las soluciones de tiempo polinomial desarrolladas para solucionar el problema del árbol de extensión mínima (Fig. 1). Las especificaciones para programar el algoritmo son de las menos complejas. Supongamos un grafo G = (N,A), con los arcos etiquetados con su costa. El algoritmo de Prim va construyendo un subconjunto de nodos U, aumentándolo hasta que contenga todos los nodos de N. Inicialmente, U contiene un único nodo de N, que puede ser cualquiera. En cada paso se localiza el arco más corto (u, n) tal que u ∈ U y n ∈ N, repitiendo el proceso hasta que U = N. Algoritmo Prim (G: Grafo; VAR T; Conjunto de Arcos); Variables U: Conjunto de arcos u, n: arcos Inicio T:={} U:={Cualquier n ∈ N} MIENTRAS U <> N HACER Sea (u,n) el arco de costo mínimo tal que u ∈ U y n ∈ N. T: = T {(u,n)} U:=U ∪ {(n)} FIN MIENTRAS Fin Prim Para encontrar fácilmente el arco de menor costo entre U y N-U, se mantienen dos matrices, MAS_CERCA NO[i], que devuelve el nodo en U más cercano a i en N-U y MENOR_COSTE [i] que devuelve el coste del arco (i, MAS_CERCANO[i]). En cada paso, se escoge k tal que k N-U es el nodo más cercano a U y se actualizan ambas matrices teniendo en cuenta que ahora k ∈ U. Para evitar que k vuelva a ser seleccionado, se asigna MENOR_COSTE[k]= .

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Fig. 1. EL algoritmo de PRIM. Representación de las soluciones de tiempo polinomial desarrolladas para solucionar el problema del árbol de extensión mínima.

Conclusiones La herramienta de trabajo Trazos 2004 ofrece una interfaz gráfica al usuario, dentro de la plataforma ArcView 3x, que permite llevar un control del proceso y desarrollo de las tres etapas básicas del método de la panbiogeografía, a través de temas espaciales gráficos referidos a un sistema de coordenadas, de manera sencilla. El hecho de catalogar como herramienta el desarrollo aquí presentado significa que su función es la de ser una extensión, un módulo adicional de nuestras capacidades, que facilita realizar una tarea específica dentro del método de la panbiogeografía, de manera automatizada pero bajo la supervisión y criterio del investigador.

La automatización computadorizada del método panbiogeográfico, a través de Trazos 2004, agiliza el delineado gráfico de los trazos, independientemente de la complejidad espacial que tenga la nube de puntos que representan las localidades del taxon o grupo de taxones que se estén investigando. A pesar de que no se trata de un programa computadorizado orientado a su comercialización, permite a diferentes grupos de investigación continuar con el desarrollo de otra serie de adiciones y aplicaciones informáticas que ayuden a mejorar la herramienta y el método de la panbiogeografía en sí mismo. Para el método de la panbiogeografía no se había desarrollado una herramienta automatizada como la que se presenta en esta investigación. La herramienta se encuentra en proceso de desarrollo y mejoramiento.

En la actualidad, se están desarrollando nuevos métodos biogeográficos para obtener modelos de predicción de los patrones espaciales de las especies, que involucran el uso de la panbiogeografía y otras áreas de estudio como los sistemas, la estadística y las matemáticas. Estas aproximaciones hacen que Trazos 2004 sea una herramienta sujeta a crecimiento sin presentar dificultades para la incorporación de nuevos métodos.

Referencias Arita H, Rodríguez P. 2002. Ecología geográfica y macroecología.

En: Llorente J, Morrone JJ (eds.). Introducción a la biogeografía en Latinoamérica: Teorías, conceptos, métodos y aplicaciones. Las Prensas de Ciencias, Facultad de Ciencias, UNAM, México, D.F. Espinosa DN, Llorente J. 1993. Fundamentos de biogeografías filogenéticas. Coordinación de Servicios Editoriales, Facultad de Ciencias, UNAM, México, D.F. Gough MC, Rushton SP. 2000. The application of GIS-modelling to 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 la biogeografía. en: Llorente J, Morrone JJ (eds.). Introducción a la biogeografía en Latinoamérica: Teorías, conceptos, métodos y aplicaciones. Las Prensas de Ciencias, Facultad de Ciencias, UNAM, México, D.F. Margules CR, Austin MP. 1994. Biological models for monitoring species decline: The construction and use of data bases. Philosophical Transactions of the Royal Society of London, B 344: 69-75.

McArthur RH. 1972. Geographical ecology: Patterns in the distribution of species. Harper and Row, New York.

Moeur M. 1985. COVER: A User’s Guide to the CANOPY and SHRUBS Extensions of the Stand Prognosis Model. General Technical Report INT-190.

Ogden UT. U.S. Department of Agriculture, Forest Service, Intermountain Research Station.

Morrone JJ. 2000. Atlas biogeográficos. PrIBES. http://entomologia. rediris.es/pribes.

Mourell C, Ezcurra E. 1996. Species richness of Argentine cacti: A test of biogeographic hypotheses. Journal of Vegetation Science, 7: 667-680.

Murguía M, Rojas F. 2001. Biogeografía cuantitativa. En: Llorente J, Morrone JJ (eds.). Introducción a la biogeografía en Latinoamérica: Teorías, conceptos, métodos y aplicaciones. Las Prensas de Ciencias, Facultad de Ciencias, UNAM, México, D.F. Myers AA, Giller PS (eds.). 1988. Analytical biogeography: An integrated 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 estudios de la biodiversidad. En: Morrone JJ, Llorente J (eds.). Una perspectiva latinoamericana de la biogeografía. Las Prensas de Ciencias, 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.), Australian Government Publishing Service, Cambridge.

Stockwell DRB. 1999. Genetic Algorithms II: Species Distribution Modelling. En: Fielding AH (ed.). Machine learning methods for ecological applications. Kluwer Academic Publishers. Cambridge, Massachussets, pp. 124-144.

Stockwell DRB, Noble IR. 1991. Induction of sets of rules from animal distribution data: A robust and informative method of data analysis. 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/tikunov 150.htm.

Camilo Andrés Rojas Parra 377 Placid Lake Drive, Sanford, 32773 Florida, USA. camian2000@yahoo.com

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U I D E L I N E S F O R

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U T H O R S

Biogeografía: Bulletin of the Systematic and Evolutionary Biogeographical Association is a forum for students, scholars and academics of biogeography to express their ideas and views about current or past issues in the discipline. We especially encourage students and young researchers to submit ideas for general articles and points of view on topics in biogeography, or about the projects with which they are currently involved. Contributed works should be written in an easily accessible and popular style. We look forward to your contributions.

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If you have been invited to contribute an article please submit your manuscript as an .odt, .rtf or .doc. We ask authors to only use Primary (bold) headings.

Citations All citations are to be made without using commas between author and year (Wallace 1855) and commas between multiple authors (e.g. Nelson & Platnick 1981, Brandon-Jones 1998). Quotes should be cited as (Willis 1922, p. 100). The ‘&’ symbol is used in all citations (but not in the references). Please refer to this issue for further usage of figures (see figure 1 or Fig. 1), tables and numerals.

References Please include the full titles of journals and books. Do not use abbreviations! Please keep your references in the styles listed below.

Brandon-Jones D. 1998. Pre-glacial Bornean primate impoverishment and Wallace’s line. In Hall R, Holloway JD eds. Biogeography and geological evolution of SE Asia. Leiden: Backhuys Publishers, pp. 393-404.

Heads M. 2006. Panbiogeography of Nothofagus (Nothofagaceae): Analysis of the main species massings. Journal of Biogeography 33: 1066-1075.

Merriam CH. 1898. Life zones and crop zones of the United States. U.S. Department of Agriculture Division Biological Survey Bulletin 10: 1-79.

Nelson G, Platnick NI. 1981. Systematics and biogeography: Cladistics and vicariance. New York: Columbia University Press.

Wallace AR. 1855. On the law which has regulated the introduction of new species. Annals and Magazine of Natural History, available at: http://www.victorianweb.org/science/science_texts/wallace_law.html Proofs Authors will be sent proofs of their articles. Authors should also make sure that they have the copyright for any photos or diagrams used in their articles.

Submissions Please submit your articles in electronic format to any of the three co-editors: Malte C. Ebach: mcebach@gmail.com John R. Grehan: jgrehan@sciencebuff.org Juan J. Morrone: jjm@hp.fciencias.unam.mx Note: This guide has been adopted from The Systematist: Newsletter of the Systematics Association (Ebach & Jenner 2006). Used with permission of the The Systematics Association.

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