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Journal of Biogeography (J. Biogeogr.

) (2005) 32, 739–740


Goodbye Gondwana

The southern end of the world is where the
two central ideas of historical biogeography
– vicariance and dispersal – have had their
most intense encounter. The geography
itself promotes controversy. The splinters
of the megacontinent of Gondwana are now
separated by vast stretches of ocean, studded
with isolated chains of volcanic archipelagos. Tracks and oceanic baselines have
proliferated in the vast information void of
the ocean basins, which are haunted by
rumours of vanished landmasses, such as
Pacifica, fragmented into any number of
conveniently potent pieces. All explanations
seem both possible and improbable at the
same time in the south. It is no accident,
therefore, that some of the most extreme
views in this debate have involved the
2000 km of ocean that have separated
Australia and New Zealand for the last
80 Myr.
Vicariance biogeography has never
explained the trans-Tasman relationship
well. Many features of current New Zealand
biogeography map perfectly onto those of
true oceanic island archipelagos such as
Hawaii, strongly suggesting a history of
dispersal, as does the apparently recent
derivation of many Australasian groups
from the Northern Hemisphere. The Australasian fossil record is clearly in conflict
with a pure vicariance interpretation for
many taxa. Therefore, if non-dispersal
explanations were the only ones permitted,
something stronger than cladistic biogeography was required.
This stronger medicine came in the form
of panbiogeography, modified and extended
by New Zealand practitioners from the late
1970s onwards. The question they addressed
was how can widespread, post-Gondwana
clades on isolated continental fragments be
explained without recourse to that devil
child, dispersal? The panbiogeographic
answer came in two forms: denial and
something close to special creation. Allegedly young post-Gondwana clades were in
fact old, dating back to pre-Cretaceous
times in some instances. More ingeniously,
it was proposed that many apparently

identical taxa in widely separated locations
‘crystallized’ out of a widespread ancestor
complex (for a full discussion see Mayben,
1991). No credible evidence has been produced to support these suggestions and the
panbiogeography programme has been well
and truly deconstructed in recent times
(Cox, 1998; McDowall, 2004). Perhaps it
can now be decently laid to rest.
While vicariance remains a respectable
and likely explanation for many disjunct
biotic distributions around the globe, recent
developments have undermined the preeminence it has enjoyed since plate tectonics
and cladistics elevated it in the 1960s and
1970s. We are now in the middle of a
dispersalist counter-revolution (see De Queiroz, 2005), fuelled by an outpouring of
molecular phylogenies. To the credit of
those who stuck by it, dispersal biogeography continued to advance through the latter
half of the last century, despite the jibes that
it was mainly a producer of ‘just-so-stories’
and despite the exhausting and unproductive task of countering the constricted
worldview of cladistic biogeography and
the alternative universe of panbiogeography.
The recent flood of evidence supporting
transoceanic dispersal (Givnish & Renner,
2004) has put it in the driving seat in the
South. Moreover, increasing recognition of
the damaging effects that invasive species
have wrought around the globe has sharpened interest in the science of dispersal and
Half a century of relative neglect means
that there are a number of underexploited topics. There has been much
recent exciting work that fully encompasses the dispersal paradigm. To take a
few recent examples: Clark et al. (2003)
tackle the thorny issues of how to model
long distance dispersal (LDD) and show
that parameterization might well be beyond our grasp; Ackerly (2004) looks in
detail at how plant trait selection occurs
during the formation of a biota by longdistance dispersal; Jordan (2001) demonstrates that recent plant dispersal events
can be interrogated to get an idea of how

ª 2005 Blackwell Publishing Ltd

plant ecology and morphology relate to
LDD probabilities; Sanmartin & Ronquist
(2004) use a meta-analysis of plant and
animal phylogenies to show that different
patterns among groups reveal those for
which dispersal has to be invoked to
explain biogeographical patterns. And, in
this issue, Cook & Crisp (2005) show how
the sea change with regard to dispersal is
leading to a rethink of the methodologies
of historical biogeography, in particular
the use of phylogenies.
Cladistic biogeography is greatly affected
by its origin in a methodology designed to
yield an unambiguous, objective classification. As dispersal was thought to overwrite
in a stochastic fashion the initial parsimonic biotic pattern established by vicariance,
it was assumed it interfered with the
derivation of informative area relationships.
Two separate approaches were used to deal
to this inconvenience. Dispersal, it was
argued, is not capable of falsification. As
dispersal could potentially explain any
pattern, it could conclusively explain no
pattern in particular. Hence, it was not
amenable to the usual procedures of
scientific discourse as codified by Popper,
and could be disregarded. As Cook and
Crisp show, this ‘baby and the bath-water’
approach is simply wrong: a rich array of
evidence and probabilistic methodology can
be brought to bear. However, even if
dispersal was regarded as unsuitable for
scientific examination, it was an unpalatable fact that it actually had occurred. How
to deal with this? The response was to
contract the scope of historical biogeography: dispersal was irrelevant noise and
could be disregarded in the search for
vicariance relationships. Why disjunctions
created by vicariance were regarded as
worthy of study, and those that are a
product of dispersal were not, has never
been adequately explained.
Cook and Crisp investigate what effect
taking the ‘noise’ seriously will have on our
interpretation of taxon–area phylogenies.
They argue that the current suite of methods
(Fitch optimization, Subtree analysis, DIVA,

doi:10.1111/j.1365-2699.2005.01278.x 739

with each taxon having its unique history and the biota as a whole having a complex network of relationships reaching across the entire globe. McLachland.. (2004) Southern Hemisphere biogeography inferred by event-based models: plant versus animal patterns. For many. 216–243. G. Jordan. and usually regard biogeographical processes as independent of each other when they are competing explanations for nodes. unchanging Gondwanic heritage is an important cultural icon. 503–519. F.L. 4 : 1 and 6 : 1. International Journal of Plant Sciences. S1–S6. De Queiroz. Cook. the methodologies employed in constructing phylogenies have predetermined the outcomes to a surprising degree. Gondwana has been taken to heart by scientists and the public alike. population genetics will give convincing evidence as to the direction of dispersal. (2004) Tropical intercontinental disjunctions: Gondwana Journal of Biogeography 32. Popperians can rest easily: falsification is possible. (2004) What biogeography is: a place for process. PO Box 69. M. C. J. Sanmartin. 84. Mayben. Australian Journal of Botany. T. All in all. immigration from the boreotropics. an asymmetry in dispersal frequency of 2 : 1. cannot be as appealing. D. 53. 741–754. (2005) Directional asymmetry of long-distance dispersal and colonization could mislead reconstructions of biogeography. Cook and Crisp ask how radically will our biogeographical reading of a phylogenetic tree be affected if we accept that repeated dispersal may have happened and assume that there is not an equal probability of dispersal in every direction? The answer is a lot. Systematic Biology. L. if not particularly easy. is that we run a serious risk of being misled by phylogenetic trees if we do not take this wider view. if we can alter the frequency and direction of dispersal at will. Whereas in the Northern Hemisphere one can go for a long time without hearing about Laurasia. ‘The Land that Time Forgot’. 25. But there is no reason why a dispersalist universe. McGlone REFERENCES Ackerly. A.M.S.). 32.G. 31.. niche conservatism. most do not consider all nodes in a cladogram. R.S. McDowall. ACKNOWLEDGEMENTS I thank Rob Smissen for helpful suggestions on this commentary. For recent dispersal events. 813–828. or ‘The Ancient Dinosaur Forests of New Zealand’ unchanged for 80 & HilleRisLambers. & Renner. (2001) An investigation of longdistance dispersal based on species native to both Tasmania and New Zealand. Clark. the hebe clade. Repeated dispersal from an anagenetically evolving population in Y can lead to its extant representatives being nested within a series of persistent. anything is possible? Cook and Crisp show that there is a wide range of evidence that can be brought to bear to prevent biogeographical chaos breaking out.B. where a conventional interpretation of phylogenetic trees suggests New Zealand to be the ancestral area. 40. however. for New Zealanders in particular. Geological and fossil evidence will assist in some circumstances. Cox.J. 165(Suppl. I. (2005) The resurrection of oceanic dispersal in historical biogeography. respectively. time and form.D. 163. 68–73. dispersed lineages in X. What is clear. abandoning ‘Time Capsule of the South Seas’ for ‘Fly-paper of the Pacific’ will be a wrench.Commentary Maximum likelihood and Bayesian inference) have a range of problems: some favour vicariance rather than dispersal explanations. and convergence: comparative studies of leaf evolution in the California chaparral. R. (2003) Estimating population spread: what can we forecast and how well? Ecology. New Zealand E-mail: mcglonem@landcareresearch. with a much diminished role for Gondwana. in favour of Australian sources (a reasonable assumption given the strong westward wind flow in this region) is sufficient to make Australia the ancestral area. But it has to happen: goodbye Gondwana. the nature of the evolutionary change in the organisms undergoing allopatric speciation gives yet further clues. analysis of vector strength and independent estimates of 740 LDD frequency can put credible limits on the degree of dispersal asymmetry. ª 2005 Blackwell Publishing Ltd . (1998) From generalised tracks to ocean basins – how useful is Panbiogeography? Journal of Biogeography. Cook and Crisp demonstrate that if there is strong asymmetry in the direction of dispersal the interpretation of the pattern can be reversed. 654–671. Trends in Ecology and Evolution. & Ronquist. Landcare Research. 345–351. Ranunculus and Myosotis. Nevertheless. 49. Journal of Biogeography. Therefore. Meanwhile. J. But is this what happened? Surely. in Australia and New Zealand. 739–740. Lincoln. and transoceanic dispersal. & Crisp. (1991) The wilderness of panbiogeography: a synthesis of space. this supposed ancient. Givnish. (2004) Adaptation. 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