news and update ISSN 1948-6596

book review
If the conceptual straitjacket fits, chances are, you’re already
wearing it
Molecular Panbiogeography of the Tropics. Michael J. Heads, 2012, University of California
Press. 565 pp. $75.00, £52.00 (harback) ISBN: 9780520271968; http://www.cambridge.org

This is part of a series of volumes aimed at, in the spective promised in the Preface.
words of its editors, examining “the role of de- Perhaps a brief refresher on the nuts and
scriptive taxonomy, its fusion with cyber- bolts of the field is warranted, since panbio-
infrastructure, its future within biodiversity stud- geographic reasoning can be perplexing for a
ies and its importance as an empirical science.” number of reasons, not the least of which is that
The author’s stated intention is to attempt to its proponents have created their own terminol-
demonstrate “patterns of community immobilism ogy. The field was proposed by Léon Croizat in
leading to allopatric differentiation, as well as 1958, and furthered by M.J. Heads as well as R.C.
other patterns of mobilism, range expansion, and Craw and J. Grehan. Panbiogeography was defined
overlap of taxa.” It was with these themes in mind by Craw et al. (1999) as a method whereby known
that I delved into Heads’ latest work. species distributions are first mapped and then
But first, in the spirit of full disclosure, I extended by drawing connecting lines on maps.
have encountered the author of this book, as well Lines are termed ‘tracks’ and are drawn such that
as several of his fellow panbiogeographers, from they ‘connect’ collection localities or disjunct dis-
the New Zealand as well as the New York schools tributions of a particular taxon. Multiple overlap-
of the subdiscipline, which together comprise a ping tracks for unrelated taxa form ‘generalized
vocal minority in biogeography. My interactions tracks’. The figures presented in the new book
with the panbiogeographers have come via a consist of dozens of black-and-white maps with
number of academic exchanges, each of which has lines and shapes superimposed to indicate distri-
unofficially ended in an understood ‘agree to dis- bution patterns.
agree’, intellectual truce. Receipt of this review Croizat was correct and ahead of his time in
request generated feelings of both hesitancy and realizing that speciation can occur after emer-
curiosity. I was hesitant to spend time on this gence of a barrier within the existing boundaries
topic because each prior exchange with Heads and of a distribution, a phenomenon that ultimately
his associates has boiled down to the willingness became known as vicariance. But this somehow
on my part (and that of our colleagues in main- led him, and the panbiogeographic movement
stream phylogeography), to approach bio- which followed, to vehemently deny, a priori, any
geographic data from a perspective that is open to role of dispersal in natural history. Croizat (1964)
both dispersal- and vicariance-based interpreta- stated “I intend to destroy these notions
tion. By contrast, panbiogeograpers strongly re- (pertaining to ‘means of dispersal’) because they
ject the significance of dispersal a priori, in any positively interfere with the advance of knowl-
form. Based on past interactions and published edge on far-reaching points of evolution over
exchanges with this group (e.g., Holland and space, in time, by form” (p. vi). This vague notion
Cowie 2006, Nelson 2006), a clear way forward in of evolution in space and time by form is repeated
this dialogue has yet to materialize. My curiosity frequently in panbiogeographic publications. Ac-
came from the chance to evaluate this recent of- cording to Cox and Moore (2005) “Even after the
fering, being due in part to my admitted lack of theory of plate tectonics had been well docu-
appreciation of the usefulness of panbiogeogra- mented and widely accepted, Croizat refused to
phy, and the possibility that my views might accept it, and never integrated it into his method-
change given the “new” panbiogeographic per- ology” (p. 31). Heads also expresses a refusal to

144 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.4, 2012
 news and update

acknowledge accepted tectonic models and in- Pacific margin of the Americas, and that further
stead focuses on his ideas of the ‘immobilism’ of support has been given by the hypothesis that
lineages. According to this idea, originally articu- some of the terranes might originally have been
lated by Croizat, continents are viewed as part of the sunken 'Pacifica' continent. According
‘composite terranes’ and come with their resident to this scenario, oceanic islands were formerly
biota intact. Panbiogeographers argue that organ- mountain peaks (e.g., Heads’ so-called “Mid-Pacific
isms have always occupied the areas where they Mountains”, Fig. 7-1, p. 315). This will sound famil-
are found today, and that these areas had been iar to many readers, since this extraordinary idea
colonized via slow continuous spread over land. was proposed in various iterations in the 18th and
Regarding the continental drift model, Craw 19th centuries. Given that this was an era where
(1985) wrote: “one cannot pull them back into a advanced technology consisted of wooden sailing
single supercontinent as Wegener did, and mod- ships, sextants and sounding lines, what sounds
ern plate tectonic proponents do.” Croizat (1958) peculiar today was perfectly reasonable, even bril-
stated that the concept of Pangea was “wrong” liant reasoning at the time. The original “mid-
because it conflicted with “evidence he had ob- Pacific plate” hypothesis preceded by several cen-
tained from his purely biogeographic studies”, turies the advent of modern scientific techniques
which I assume refers to the drawing of lines on (K–Ar radiocarbon dating; high-resolution, side-
maps with a pencil and a ruler. The panbio- scan, multibeam sonar; GPS tracking) and a de-
geographic position on geologic history and plate tailed understanding of the ages and dynamics of
tectonics has been reviewed at length by Cox tectonic plates. So the panbiogeographers inten-
(1998). tionally not only ignore or misinterpret many im-
Heads, like the other panbiogeographers, portant advances in natural science (molecular
fervently rejects dispersal followed by speciation clock theory, plate tectonics, continental drift, ra-
as a possible mechanism influencing an organism’s dio-carbon dating, oceanic island formation, ero-
distribution under any circumstance. This radical, sion and subsidence) but they seem to imply that
recurrent theme is seen throughout the book they would prefer to turn back the hands of time
(multiple times per chapter) as Heads forces ob- to a period of technological simplicity and scientific
served data to fit the preconception that there is ignorance, in order to fulfill their agenda.
no dispersal, aside from what he and other panbio- The first two chapters of Molecular Panbio-
geographers term ‘normal ecological dispersal’. geography of the Tropics revisit the father of pan-
The ideas presented in this book are anything but biogeography’s semi-metaphysical “evolution in
“new”. The underpinnings of panbiogeography space and time by form” ideas, and present
were summarized by Craw (1984) as Croizat’s Heads’ own take on his effort to go against the
“bold and novel attempts to empirically refute” grain of “biogeographic interpretation over the
the work of Charles Darwin, Alfred Russel Wallace last 2,000 years”. Chapters 3–5 present scenarios
and Alfred Wegener (p. 8). Implicit in many of the and offer explanations of origins and relationships
arguments of panbiogeography is the idea that between New and Old World primates, each un-
fossils of common ancestors represent minimum surprisingly chosen to favor the vicariant agenda.
(rather than the standard interpretation as maxi- The next three chapters concern what seems to
mum) ages of extant lineages, and there were be one of Heads’, and my own, favorite topics: the
emergent Pacific land masses, some of which con- evolution of life on insular Pacific islands.
nected the far-flung island archipelagos and It is here (Chapter 7) that Heads’ recurrent
continents surrounding the Pacific basin. The New thesis, that long-distance dispersal is not a factor
Zealand panbiogeographers—including Heads, in evolutionary history, reaches its pinnacle. This
Craw and Grehan—claim that Croizat's concept of chapter deals with what he refers to as “the near-
'horstian dispersal' has been validated by the rec- universal consensus” (p. 314) that Hawaiian Island
ognition of ‘accreted terranes’ embedded in the biota largely derives from long-distance dispersal,

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news and update

followed by adaptation and radiation. The nature exaggerate controversy and overplay scientific
of his argument is that we should use caution uncertainty where convenient, in support of the
prior to deducing that species ever colonize oce- agenda. For example “maps of the seafloor are far
anic islands and evolve phyletically in place. He from complete”, and rates of molecular evolution
suggests that arcs of once-emergent ancient sea- are deemed universally “elastic” and therefore
mounts between, say, the Hawaiian and Marque- unreliable. In Heads’ world, even great white
san islands played a role, by harboring the line- sharks are not permitted to have attained their
ages we see today (which would surely necessi- present distribution and genetic structure via dis-
tate dispersal via island hopping?), over tens of persal in the Pacific basin, but “instead, the pat-
millions of years. But again this would require nu- tern could be the result of vicariance in a wide-
merous departures from what is understood and spread ancestor…the westward annual migrations
accepted by geophysicists. He presents maps of the populations may have been extended a few
showing filled-in outlines of implied land masses centimeters every year in step with the geological
in places where there is no hydrographic or geo- movement” (p. 332).
physical evidence that such landscapes ever ex- Early in Chapter 8, regarding the islands of
isted (Fig. 7-1 p. 315 and 7-2 p. 316). Even though Maui and Hawaii, and against all geophysical evi-
these features are described in the figure captions dence, Heads states that “there may have been
as “2,000”, “4,000” and “5,000 m isobaths”, the land between the two islands” (p. 357). Later he
figures appear to be a disingenuous and mislead- goes on to explain the extraordinary endemism of
ing depiction aimed at advancing the vicariant the Hawaiian Islands using wild speculation: “For
agenda. To me, this seems a high interpretive example, if all of North America were flooded ex-
price to pay in order to avoid founder speciation. cept for one central area the size of the Hawaiian
Heads argues further, attempting to cast Islands, and this was preserved and isolated from
doubt on established geological principles, that other continents, after 70 million years it would
the Pacific hot spot position is not well character- probably have a very high level of endemism, as-
ized, that mantle plumes may not exist, that the suming there was little or no colonization of the
Hawaiian Islands may have once been joined with community from outside areas” (p. 388). He then
other land masses, and “In fact the whole subject attempts to critique the widely held, useful bio-
of intraplate volcanism is currently being debated geographic concept of disharmonic distributions
among geologists” (p. 317). I wonder which geolo- of island biota, by stating that were comparisons
gists? Each case study ultimately devolves into an made among continental species, since they are
endeavor to cast suspicion on conventional rea- not identical, they could be termed “disharmonic”
soning, and hence promote one or more of the as well (p. 388). Conveniently overlooked in
recurring themes, discounting: the role of long- Heads’ interpretation here are ideas of bio-
distance dispersal, founder speciation, center of geographic dispersal barriers, filters, missing line-
origin and the use of fossil ages and/or molecular ages, sweepstake routes (e.g., Cox & Moore
divergence values to estimate lineage ages. The 2005), and the idea that geographically isolated
author invokes vicariant forces in each and every islands tend to lack entire categories of low-
instance. Examples include endemic oceanic island vagility flora and fauna and have disproportion-
lineages, whether strong-flying insects and birds, ately high biodiversity of dispersing lineages
marine taxa such as cetaceans or invertebrates (Cowie & Holland 2008). Heads’ approach renders
and fish with a planktonic larval phase. All of this scientific evaluation of the panbiogeographic
adds up to a strategy reminiscent of other efforts ideas presented difficult. This book inadequately
aimed at undermining confidence in empirical sci- addresses the stated goals of the series, especially
ence, namely the creationist/intelligent design in terms of examining the role of descriptive tax-
and climate change denial movements. Similari- onomy in biodiversity and empirical science. There
ties in approach include attempts to cast doubt, is nothing empirical about determining an out-

146 © 2012 the authors; journal compilation © 2012 The International Biogeography Society — frontiers of biogeography 4.4, 2012
 news and update

come in advance, then forcing all observed data to “…geophysical findings…promise to corroborate Croi-
fit the preconception. I will go one step further zat’s biogeographic work, in novel and exciting ways”.
and argue that this approach is antithetical to the Alas, this was not to be.
scientific process.
After spending time with this book I view it Acknowledgments
as less of a scientific publication than a 10-chapter I am grateful to Markus Eichhorn for the opportu-
philosophical argument, continuing the promo- nity to discuss this interesting, and in its own idio-
tional efforts of the panbiogeographic agenda in syncratic way, important book and subject. Thanks
all of its a priori glory. This is an attempt to sway to Richard Field for suggestions that improved this
the reader to consent to a peculiar conceptual manuscript, and to my students at the University
framework lacking in scientific rigor. The author of Hawaii, including Zak Williams and Melissa
has expended a commendable amount of effort to Wright, for lively discussions of panbiogeography,
review the relevant systematic literature, only to all of which were enjoyable, some of which led to
incessantly return to the central views delineated clarification of my own appraisal of these topics.
by preceding panbiogeographic efforts. Where
this book succeeds is in convincing me that the Brenden S. Holland
central incongruities between approaches of con- Pacific Biosciences Research Center, University of Ha-
ventional biogeographic science and those of pan- waii, Honoululu, USA; [email protected]
biogeography are fundamental, and emanate
References
from opposing interpretations of scientific data.
Cowie, R.H. & Holland, B.S. (2008) Molecular biogeogra-
In one interesting passage, Heads states “It seems phy and diversification of the endemic terrestrial
that the progression rule has become a concep- fauna of the Hawaiian Islands. Philosophical
tual straitjacket” (p. 360). On the contrary, the Transactions of the Royal Society of London B,
progression rule is used by conventional scientists 363, 3363–3376.
as a theoretical starting point, a model from which Cox, C.B. & Moore, P.D. (2005) Biogeography: an eco-
logical and evolutionary approach. Blackwell
we discover and acknowledge both departures
Publishing, Malden MA, USA.
and adherence equally, readily and frequently Cox, C.B. (1998) From generalized tracks to ocean basins
(Cowie and Holland 2008). Close evaluation of – how useful is Panbiogeography? Journal of
panbiogeography suggests that the colorful term Biogeography, 25, 813–828.
“conceptual straitjacket”, used by Heads himself, Craw, R.C. (1984) Leon Croizat’s biogeographic work: a
is a flawless fit for their own mode of reasoning. personal appreciation. Tuatara, 27, 8–13.
Craw, R.C., Grehan, J.R. & Heads, M. (1999) Panbio-
Heads claims, and I agree, that “a healthy skepti-
geography: tracking the history of life. Oxford
cism is one of the pillars of science” (p. 3). Perhaps Biogeography Series No. 11, Oxford University
where we begin to disagree is on precisely what is Press, USA.
a “healthy skepticism”. Panbiogeographic skepti- Croizat, L. (1958) Panbiogeography. Published by the
cism is in my view agenda-driven, obdurate and author, Caracas.
misdirected. This relates to my personal tolerance Croizat, L. (1964) Space, time, form; the biological syn-
thesis. Published by the author, Caracas.
for both how far afield alternative viewpoints in
Grehan, J. (1994) The beginning and end of dispersal:
science can go, as well as the validity of the justifi- the representation of panbiogeography. Journal
cation underpinning iconoclastic positions. of Biogeography, 21, 451–462.
I echo the statement in the review of panbio- Holland, B.S. & Cowie, R.H. (2006) Dispersal and vicari-
geography by Cox (1998) that it is time for panbio- ance in Hawaii: submarine slumping does not
create deep inter-island channels. Journal of Bio-
geographers to abandon their defensive position, geography, 33, 2155–2156.
stop objecting that conventional biologists do not Nelson, G. (2006) Hawaiian vicariance. Journal of
take their work seriously, and instead begin to ad- Biogeography, 33, 2154–2157.
dress the criticisms that have been leveled at their
work for decades. In the wishful words of Craw (1984) Edited by Markus Eichhorn

frontiers of biogeography 4.4, 2012 — © 2012 the authors; journal compilation © 2012 The International Biogeography Society 147