[Taxacom] scientific predictions concerning Wallacean marsupials and primates
John Grehan
calabar.john at gmail.com
Thu Jun 21 10:12:26 CDT 2018
I have found Ken's assertions about chance dispersal to be quite
informative as it shows that the manner of reasoning about chance dispersal
has not changed from the time of Darwin who imagined the process and felt
that it could not be denied. Some of the inherent problems with
imagination, I would argue, is that they lack a coherent analytical
foundation and instead tend to ignore or explain away anything to the
contrary.
For example, with the statement:
"The examples given in the last paragraph of your post are indeed
fascinating and indicative of vicariance, And although they could also
involve some short distance island hopping as well, no long distance
dispersals would be required."
If they could involve island hopping then the patterns cannot be indicative
of vicariance. What is left out of the picture here is that there is a
cohesive set of patterns correlated with tectonics which does provide
evidence of a vicariance history. Here there is a direct link in reasoning
between evidence and conclusion.
"They are therefore somewhat similar to the marsupial expansion westward
into Wallacea. "
Of course this begs the question of what is the empirical evidence that
marsupials ever 'expanded' into the region.
"This is exactly the opposite of tarsiers that came from the other
direction and have their easternmost occurrences there."
Again, no empirical evidence has every been presented for such a notion.
"None of your examples are of taxa going from west to east, so it is sort
of like comparing apples and oranges."
Do you mean taxa that have an eastern boundary in SEA?
"In any case, all those examples just make me even more impressed that
cuscuses (which also expanded westward)"
No evidence.
"were actually able to become established even further westward into
Sulawesi. Of course, it could be that they just entered Wallacea a lot
earlier than the examples you gave. Or perhaps they are very competitive
and very good at dispersal."
Or perhaps not. No evidence. I have no problem with propositions of chance
dispersal, but in a science of biogeography one needs more than just an
assertion of a possiblity.
"I would predict that carefully designed experiments..."
Problem with these experiments is that they do not actually provide a
exclusionary text - the outcomes either way cannot exclude the alternative.
Thus one might find they do not coexist which might be dismissed as an
artifact of the current conditions, or they do coexist which might also be
explained away by the same argument. Early on Darwin 'tested' snail
dispersal by showing that pond snails could adhere to duck's feet or that
seeds could survive immersion in salt water. But these experiments actually
did not demonstrate anything in particular about whether or not such
ecological processes were involved in the origins of allopatric
distribution. Rather than ecology being informative about the mechanisms of
biogeographic origin, it is biogeographic analysis that is informative
about the role of ecological processes in biogeographic differentiation
(evolution).
"And thirdly, any evidence showing that cuscuses might have entered
Sulawesi earlier than monkeys could also be scientifically valuable. My
hypothesis would predict that cuscuses probably got there before monkeys."
Herein lies the crux of the matter. Is the occurrence of cuscuses in
Sulawesi linked to the tectonic history of the region or the result of
chance dispersal. The biogeography would indicate that tectonics has been
the main player in the structure of allopatric distributions in SEA. While
some on this list have been critical of the emphasis on vicariance, this
emphasis is analysis driven, not simply imagined and asserted.
While at least one person has characterized this discussion as a waste of
time etc., I do find Ken's perspective of interest to discuss and critique.
It is actually quite rare in the literature to directly view any direct
interactions between supporters of chance dispersal and supporters of
vicariance evidence. So in that respect the discussion, even if it may not
resolve anything in any absolute sense, may be helpful to those who are in
the process of developing their perspectives (and I am sure that there are
some on this list in that situation).
John Grehan
On Thu, Jun 21, 2018 at 7:36 AM, Kenneth Kinman <kinman at hotmail.com> wrote:
> Hi Michael,
>
> The examples given in the last paragraph of your post are indeed
> fascinating and indicative of vicariance, And although they could also
> involve some short distance island hopping as well, no long distance
> dispersals would be required.
> However, all these examples are taxa whose westernmost occurrences
> are on islands such as Peleng and Sangihe. They are therefore somewhat
> similar to the marsupial expansion westward into Wallacea. This is exactly
> the opposite of tarsiers that came from the other direction and have their
> easternmost occurrences there. None of your examples are of taxa going
> from west to east, so it is sort of like comparing apples and oranges.
> In any case, all those examples just make me even more impressed
> that cuscuses (which also expanded westward) were actually able to become
> established even further westward into Sulawesi. Of course, it could be
> that they just entered Wallacea a lot earlier than the examples you gave.
> Or perhaps they are very competitive and very good at dispersal.
> ANYWAY, if you want predictions that are scientifically testable,
> here are some. I would predict that carefully designed experiments placing
> male cuscuses with monkeys (male or female) of roughly equal weight
> (including cuscuses vs. macaques) might well find cuscuses able to hold
> their ground. If so, my hypothesis of well-established cuscuses on islands
> east of Sulawesi being able to competitively exclude dispersing pregnant
> females (in particular) could provide scientific evidence in its favor.
> And secondly, given the size of Sulawesi and its wide range of habitats,
> detailed maps (of "fine geographic scale") of cuscus and macaque
> distributions might also be scientifically valuable. And thirdly, any
> evidence showing that cuscuses might have entered Sulawesi earlier than
> monkeys could also be scientifically valuable. My hypothesis would predict
> that cuscuses probably got there before monkeys.
> -------------------Ken
>
> ________________________________
> From: Michael Heads <m.j.heads at gmail.com>
> Sent: Wednesday, June 20, 2018 5:12 PM
> To: Kenneth Kinman
> Cc: Taxacom
> Subject: Re: [Taxacom] Fwd: competition on Peleng (actually versus two
> specie of cuscus)
>
> Hi Ken,
>
> Consider a group present in area A and B, while its sister is in B and C.
>
> Dispersal theory (your model) proposes a centre of origin of the whole
> group in B, followed by chance dispersal to A in one group, and to C in the
> other.Vicariance theory (my model) proposes an ancestor present in A, B and
> C, followed by a vicariant break somewhere in or around B (mediated by
> geology/climate), followed by overlap (dispersal) in B by one or both
> groups (caused by geology/climate).
>
> Monkeys, apes etc. (simians or anthropoids) range through America, Africa
> and Asia to Sulawesi (not Peleng, Siau or Sangihe). Their sister goup,
> tarsiers, range from SE Asia to Peleng, Siau and Sangihe (PSS). In
> dispersal theory, the tarsiers on PSS migrated to these islands (not their
> precursors) from the Sulawesi mainland using means that are essentially
> mysterious (Shekelle). In vicariance, the records on PSS are not the result
> of random processes, but reflect the ancestral distribution of the simian +
> tarsier common ancestor, and the break between them, caused by geological
> change. PSS and eastern Sulawesi are the result of terranes and subduction
> zones that have migrated west from New Guinea and the Pacific, 'piling up'
> at Sulawesi. The PSS tarser belt is probably a small fraction, a relic, of
> the original insular distribution, while the large ocean basins they were
> distributed across have been subducted.
>
> The key question is: why is the 'tarsier belt' on these particular small
> islands (PSS) and not others in the region? How can we distinguish between
> the two models? Dispersal theory makes no special predictions, as chance
> dispersal is not related to any other factor, biological or physical.
> Vicariance theory makes the prediction that these geographically
> insignificant islands will also be significant biogeographically for many
> other groups, as they have been exposed to the same geology. Here are some
> examples:
>
> The bird genus Eutrichomyias is endemic on Sangihe (= Sanghir) and has its
> closest relative in Fiji (see 'Relicts of the lost arc..', Mol Phylogen
> Evol 120: 28. 2018). The closest relative of the bird Colluricincla
> sanghirensis, endemic on Sangihe, is on islands off the eastern tip of New
> Guinea. The Pacific clade of the skink Lamprolepis has western outliers on
> Peleng and Sangihe (Fig. 10.6 in my Australasia book). (Note that Peleng
> and Sangihe are 600 km apart, while Peleng is only 25 km off mainland
> Sulawesi). The parrot genus Eos ranges from New Guinea to Sangihe. Another
> parrot Loriculus amabilis is only on Peleng (also nearby Sula), Siau, and
> Sangihe. The diverse frog Platymantis s.l. ranges from Fiji to Peleng and
> Sangihe (and the Philippines), but not Sulawesi (Fig. 10.7 in my
> Australasia book).
>
>
>
> On Thu, Jun 21, 2018 at 1:14 AM, Kenneth Kinman <kinman at hotmail.com
> <mailto:kinman at hotmail.com>> wrote:
> Hi Michael,
>
> Although my hypothesis of Wallacea being a battleground between
> primates and cuscuses ("marsupial monkeys" as they are sometimes called)
> may seem ad hoc, but I think it is worth pursuing. I finding it
> particularly interesting that the bear cuscus is unusual in being diurnal,
> so we have both diurnal and nocturnal cuscuses and primates in this
> battleground.
>
> Sulawesi being a much larger island seems able to accomodate a mix
> of species that would not be possible on the much smaller islands to the
> east. That tarsiers were able to spread to some of those much smaller
> islands could be due to their small size and secretive nature. Or perhaps
> they just got there before cuscuses.
>
> All of these small islands in Wallacea seem to be an evolutionary
> filter in both directions, and as a mammalogist, I am finding the mix of
> primates and marsupials particularly interesting. And cuscuses ("marsupial
> monkeys") are not as well studied, and it is thought that there are still
> undiscovered species in this region.
>
> But there are admittedly other factors that make this battleground
> region more difficult to understand. The changing ocean currents and
> changes in sea level that Scott mentioned could be a major factor for some
> organisms and not for others. It is no wonder why biogeographers find
> Wallacea so interesting (and frustrating).
>
> -----------Ken
>
>
>
> ________________________________
> From: Taxacom <taxacom-bounces at mailman.nhm.ku.edu<mailto:taxacom-bounces@
> mailman.nhm.ku.edu>> on behalf of Michael Heads <m.j.heads at gmail.com
> <mailto:m.j.heads at gmail.com>>
> Sent: Wednesday, June 20, 2018 6:33 AM
> To: Taxacom
> Subject: [Taxacom] Fwd: competition on Peleng (actually versus two species
> of cuscus)
>
> On Wed, Jun 20, 2018 at 11:06 AM, Michael Heads <m.j.heads at gmail.com
> <mailto:m.j.heads at gmail.com>> wrote:
>
> > Hi Ken,
> >
> > Monkeys range east only to Sulawesi, while the only primates east of
> there
> > are tarsiers on the interesting chain of islands: Peleng, Siau and
> > Sangihe. The cuscuses Strigocuscus and Ailurops are both on Sulawesi
> > and Peleng, Strigo. is also on Siau and Sangihe. Why would the cuscuses
> > prevent monkeys establishing on Peleng, Siau and Sangihe by competition
> > when they all live together on Sulawesi?
> >
> > A closely related question is why and how tarsiers are on Peleng, Siau
> and
> > Sangihe. The expert on this group, Myron Shekelle, admitted that 'How
> > tarsiers ever came to these islands is a mystery'. Also, why are
> > tarsiers on *these *islands, but not further east?
> >
> > The islands in this 'tarsier belt' - Peleng, Siau and Sangihe - are
> > geographically insignificant, but form one of the most interesting areas
> in
> > the world for biogeographers and geologists (distributions and geology in
> > the region are illustrated in Figs. 5-15 and 5-16 of my Tropics book).
> E.g.
> > apart from the tarsiers it has one of the highest densities of
> > endemic birds in the world. Why? Many of the birds are related to groups
> > further east (including eastern New Guinea groups, 1000s of km away), not
> > to groups on the much closer Sulawesi. The geology is also very unusual
> - the
> > Sangihe and Halmahera arcs are undergoing the only active arc collision
> on
> > Earth. Most biogeographers understand that at a subduction zone, such as
> > the Sangihe SZ, the plates move towards each other. What many
> > biogeographers do not understand is that active subduction zones
> (producing
> > islands) themselves often move long distances, at rates of ~10cm/yr.
> >
> >
> >
> > On Tue, Jun 19, 2018 at 2:30 PM, Kenneth Kinman <kinman at hotmail.com
> <mailto:kinman at hotmail.com>>
> > wrote:
> >
> >> Dear all,
> >>
> >> I am still catching up on the changing taxonomy of the cuscuses.
> >> Back when I was working on our 1st Edition of the reference book "Mammal
> >> Species of the World" (1st Edition, by Honacki, Kinman, and Koeppl,
> 1982),
> >> Strigocuscus was included within genus Phalanger, but genetic data since
> >> then has apparently shown them to be in separate subfamilies.
> >>
> >> In any case, although the species of Strigocuscus on Sulawesi and
> >> Peleng Island is a "pygmy" species, the other species on Peleng is a
> more
> >> normal size. I had overlooked is that the bear cuscus is also on Peleng
> >> Island. So any monkey trying to become established on Peleng Island
> would
> >> have faced two species of well-established species of cuscuses.
> >>
> >> So it is even less surprising that monkeys did not expand east of
> >> Sulawesi. The battle ground between monkeys and marsupial cuscuses
> seems
> >> most intense between Sulawesi and nearby Peleng Island. This
> battleground
> >> in Wallacea seems to have favored cuscus over monkeys, even though they
> >> managed to coexist on the westernmost Wallacean island of Sulawesi.
> Just a
> >> matter of Sulawesi being a larger island with more competition allowing
> a
> >> stalemate of sorts. Both monkeys and cuscuses survived there, but not
> >> further east, and cuscuses did not survive further west. Something that
> >> neither Wallace's Line nor Weber's Line seems to have anticipated.
> >>
> >> -----Ken Kinman
> >>
> >> P.S. When cuscuses were first discovered they were thought to be a
> >> species monkey. So similar to monkeys in their form, habitat, and food
> >> sources, but only the discovery that cuscuses were pouched marsupials
> did
> >> their true relationship become known. I'm not sure if you could say
> that
> >> they are more like monkeys or lemurs. In any case, monkeys would have
> met
> >> their match in both Madagascar (versus lemurs) and east of Sulawesi
> (versus
> >> the two species of cuscuses).
> >>
> >>
> >> ------------------------------
> >> *From:* Taxacom <taxacom-bounces at mailman.nhm.ku.edu<mailto:
> taxacom-bounces at mailman.nhm.ku.edu>> on behalf of
> >> Kenneth Kinman <kinman at hotmail.com<mailto:kinman at hotmail.com>>
> >> *Sent:* Monday, June 18, 2018 8:54 AM
> >> *To:* Michael Heads; taxacom at mailman.nhm.ku.edu<mailto:
> taxacom at mailman.nhm.ku.edu>
> >> *Subject:* Re: [Taxacom] competition on Peleng (was: Jurassic
> >> primates???)
> >>
> >> Hi Michael,
> >>
> >> Well, I was talking about monkeys in general, but even with
> >> macaques, I would think a male cuscus could take on a female macaque.
> When
> >> I talk about transoceanic dispersal of primates, I am usually thinking
> >> about of a group of pregnant females. No adult males required, as long
> as
> >> one or more of the females are pregnant with male offspring.
> >>
> >> Such a group of pregnant female monkeys landing on a small island
> >> like Peleng that has a well-established cuscus population could have a
> very
> >> rough time trying to get established. If not killed by male cuscuses,
> the
> >> cuscuses could drive the pregnant monkeys away from the best food
> sources
> >> (and they just die from malnutrition). Likewise, cuscuses haven't
> spread
> >> further east because monkeys were already well established there.
> >>
> >> Monkeys introduced by humans in New Guinea is a different story.
> >> They get established around human habitations and then spread out from
> >> there. And there is a lot more territory than on a small island like
> >> Peleng.
> >>
> >> --------------Ken
> >>
> >>
> >> ________________________________
> >> From: Michael Heads <m.j.heads at gmail.com<mailto:m.j.heads at gmail.com>>
> >> Sent: Monday, June 18, 2018 2:09 AM
> >> To: Kenneth Kinman
> >> Subject: Re: [Taxacom] competition on Peleng (was: Jurassic primates???)
> >>
> >> Hi Ken,
> >>
> >> You say: 'Cuscuses are a very monkey-like marsupial', and suggest that
> >> they prevented monkeys from establishing east of Sulawesi.
> >>
> >> I know cuscuses and monkeys quite well (I lived for a long time in
> Africa
> >> and in New Guinea) and I wouldn'y describe them as very similar at all
> >> really - for a start, the monkeys concerned (macaques) are at least
> twice
> >> the size of a cuscus, much smarter and much more aggressive. But in any
> >> case, New Guinea is very rich in cuscuses and, as I mentioned,
> introduced
> >> monkeys have established there with no problems.
> >>
> >> I'm sure you can think up an ad hoc reason for that particular
> >> phenomenon, but I prefer to look at the group, and all its biogeographic
> >> boundaries, overall. As Edgar Allan Poe said, 'the ingenious are always
> >> fanciful, and the truly imaginative never otherwise than analytic'.
> >>
> >>
> >> On Mon, Jun 18, 2018 at 2:49 PM, Kenneth Kinman <kinman at hotmail.com
> <mailto:kinman at hotmail.com>
> >> <mailto:kinman at hotmail.com<mailto:kinman at hotmail.com>>> wrote:
> >>
> >> Hi Michael,
> >>
> >> Sulawesi and Peleng (being in the western part of "Wallacea") are
> >> an interesting battleground between the Asian and Australian faunas. In
> >> answer to your question, I suspect that monkeys could have encountered
> >> competitive exclusion on Peleng by a combination of not only the
> tarsiers
> >> there, but even more so by the marsupial species Strigocuscus
> pelengensis.
> >> This marsupial is also in the Sula Islands, which might also explain why
> >> monkeys didn't get into the Molucca Islands either (although a greater
> >> distance gap might also explain that).
> >>
> >> The other species of Strigocuscus does live on Sulawesi, but being
> >> called a dwarf cuscus, I assume it is smaller (and therefore less
> >> competitive and aggressive) than the species on Peleng. So I would say
> >> that studying the possible battle between the cuscus marsupials and
> monkeys
> >> in western Wallacea (especially on Peleng) could be far more fruitful
> than
> >> just tarsiers vs. monkeys. It could be a combination of both battles in
> >> this transition zone.
> >>
> >> The males of cuscuses in particular can be quite aggressive. And
> >> the cuscus on Peleng (presumably being larger than those on Sulawesi and
> >> being on a much smaller island) might be the main reason any attempted
> >> monkey dispersals would fail. Probably more so than tarsiers. Cuscuses
> >> are a very monkey-like marsupial.
> >>
> >> -----------------Ken Kinman
> >>
> >>
> >> ________________________________
> >> From: Michael Heads <m.j.heads at gmail.com<mailto:m.j.heads at gmail.com
> ><mailto:m.j.heads at gmail.com<mailto:m.j.heads at gmail.com>>>
> >> Sent: Sunday, June 17, 2018 6:31 PM
> >> To: Kenneth Kinman
> >> Cc: Taxacom
> >> Subject: Re: [Taxacom] Jurassic primates???
> >>
> >> Hi Ken,
> >>
> >> Mesozoic mammal fossils are very different from modern groups, and their
> >> affinities are often very controversial or simply admitted as unknown.
> If
> >> modern morphological work identified the phylogenetic position of groups
> >> such as cetaceans incorrectly - when the vast wealth of morphological
> >> information was available from complete specimens - isn't it very likely
> >> that they have misidentified many, fragmentary Mesozoic fossil groups in
> >> which the key features are not preserved?
> >>
> >> Did you see the lizard paper I mentioned (pushing back dates by 75
> m.y.)?
> >> Do you agree with my earlier statement that Goswami & Upchurch's
> reasoning
> >> is illogical (they wrote that fossils provide minimum ages, but then
> >> treated the fossil age of eutherians as a maximum age for primates)?
> Also,
> >> you haven't said which examples of vicariance you accept.
> >>
> >> Some other 'extraordinary' evidence for primates:
> >> Monkeys are diverse on Philippines/Sulawesi, but they reach their limit
> >> there and are entirely absent from islands to the east, such as Peleng,
> 20
> >> km off eastern Sulawesi, the Moluccas etc. If monkeys dispersed across
> the
> >> Atlantic long after it opened, why would a 20 km strait block their
> >> dispersal?
> >>
> >> Peleng is a small island, so you might think that large monkeys could
> not
> >> exist there. But macaques are famously 'weedy' and Peleng is the same
> size
> >> as Zanzibar (30 km off the African mainland), which has endemic monkeys.
> >>
> >> You might hypothesize that the tarsiers on Peleng prevented the monkeys
> >> from establishing there by competition, but monkeys and tarsiers overlap
> >> through most of the tarsiers' range.
> >>
> >> You might suggest that islands east of Philippines/Sulawesi are
> >> ecologically unsuitable for some other reason, but monkeys introduced to
> >> Palau and to western New Guinea have thrived and are now pests.
> >>
> >> Of course, the 'extraordinary' break between Sulawesi and Peleng could
> >> just be due to the vagaries of chance dispersal, but wouldn't it be
> useful
> >> to investigate other possibilities?
> >>
> >>
> >>
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> >
> > --
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> Biogeography-and-Evolution-in-
> > New-Zealand/Heads/p/book/9781498751872
> >
> >
> > *Biogeography of Australasia: A molecular analysis*. Cambridge
> > University Press, Cambridge. 2014. www.cambridge.org/9781107041028<
> http://www.cambridge.org/9781107041028><http://www.cambridge.org/
> 9781107041028>
> >
> >
> > *Molecular panbiogeography of the tropics. *University of California
> > Press, Berkeley. 2012. www.ucpress.edu/book.php?isbn=9780520271968<
> http://www.ucpress.edu/book.php?isbn=9780520271968><http://www.
> ucpress.edu/book.php?isbn=9780520271968>
> >
> >
> > *Panbiogeography: Tracking the history of life*. Oxford University Press,
> > New York. 1999. (With R. Craw and J. Grehan). http://books.google.
> > co.nz/books?id=Bm0_QQ3Z6GUC<http://co.nz/books?id=Bm0_QQ3Z6GUC>
> > <http://books.google.co.nz/books?id=Bm0_QQ3Z6GUC&dq=
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> >
> >
> >
> >
> >
> >
> >
> >
> >
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> >
>
>
> --
> Dunedin, New Zealand.
>
> My books:
>
> *Biogeography and evolution in New Zealand. *Taylor and Francis/CRC, Boca
> Raton FL. 2017.
> https://www.routledge.com/Biogeography-and-Evolution-in-
> New-Zealand/Heads/p/book/9781498751872
>
>
> *Biogeography of Australasia: A molecular analysis*. Cambridge University
> Press, Cambridge. 2014. www.cambridge.org/9781107041028<http://www.
> cambridge.org/9781107041028><http://www.cambridge.org/9781107041028>
>
>
> *Molecular panbiogeography of the tropics. *University of California Press,
> Berkeley. 2012. www.ucpress.edu/book.php?isbn=9780520271968<http://www.
> ucpress.edu/book.php?isbn=9780520271968><http://www.
> ucpress.edu/book.php?isbn=9780520271968>
>
>
> *Panbiogeography: Tracking the history of life*. Oxford University Press,
> New York. 1999. (With R. Craw and J. Grehan).
> http://books.google.co.nz/books?id=Bm0_QQ3Z6GUC
> <http://books.google.co.nz/books?id=Bm0_QQ3Z6GUC&dq=
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> http://mailman.nhm.ku.edu/cgi-bin/mailman/listinfo/taxacom
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> taxacom-owner at mailman.nhm.ku.edu<mailto:taxacom-owner at mailman.nhm.ku.edu>
>
> Nurturing Nuance while Assaulting Ambiguity for 31 Some Years, 1987-2018.
>
>
>
> --
> Dunedin, New Zealand.
>
> My books:
>
>
> Biogeography and evolution in New Zealand. Taylor and Francis/CRC, Boca
> Raton FL. 2017. https://www.routledge.com/Biogeography-and-Evolution-in-
> New-Zealand/Heads/p/book/9781498751872
>
>
> Biogeography of Australasia: A molecular analysis. Cambridge University
> Press, Cambridge. 2014. www.cambridge.org/9781107041028<http://www.
> cambridge.org/9781107041028>
>
>
> Molecular panbiogeography of the tropics. University of California Press,
> Berkeley. 2012. www.ucpress.edu/book.php?isbn=9780520271968<http://www.
> ucpress.edu/book.php?isbn=9780520271968>
>
>
> Panbiogeography: Tracking the history of life. Oxford University Press,
> New York. 1999. (With R. Craw and J. Grehan). http://books.google.co.nz/
> books?id=Bm0_QQ3Z6GUC<http://books.google.co.nz/books?id=Bm0_QQ3Z6GUC&dq=
> panbiogeography&source=gbs_navlinks_s>
>
>
>
>
>
>
>
>
>
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>
> Nurturing Nuance while Assaulting Ambiguity for 31 Some Years, 1987-2018.
>
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