Taxacom: On describing new taxa without using phylogenetics: some suggestions

Thu Oct 9 06:40:50 CDT 2025

I have been following this discussion with some interest in particular with
respect to discussions of fossils and the overall "desirability" of
molecular datasets over morphological ones. I agree with the premise of
phylogenetics should be in any species description, I not Richard Pyles
point here I mean identifying and demonstrating a population is a species
and how it is related, not naming it thats another issue.

Throughout my career I have worked on both living and fossil species,
desribing and doing phylogenetic analyses on both, combined. Furthermore
working with turtles I am working with a group that has the unusual
situation of having 4 times more fossil taxa than living. Rounding off the
numbers about 370 living and about 1200 fossils. This disparity of numbers
has a major impact as when using molecular data, almost always on living
taxa though we do have a number of extinct taxa where DNA has been
preserved (a couple of dozen of the 1200 so dont get too excited and of
course all more recent). When you add 1200 species to a dataset of a little
under 400 species the tree explodes basically, completely detroying any
statistical confidence in the relationships.

The big issue in combining datasets is not philosophical it is practical.
Most people working with molecular data do not have the skills or funding
to do decent morphology, and vice versa. This requires complex
collaborations and difficult work. Note I am working on vertebrates, highly
endangered ones which generally have extremely difficult and expensive
access issues for morphological data.

Add to this is what people call morphology, in many largely molecular
papers on vertebrates it is mostly morphometrics, which cannot produce a
phylogenetic analysis, it is more akin to numerical taxonomy of the 1980s.
Not useless and can certainly help describe a taxa, but it does not infer
relationships. To develop the tree of a group and how they all evolved from
one another you must be infering relationships. If your using multiple
datasets (which I would encourage) then they must be comparable and hence
the morphology and the molecular data must be analised with phylogenetics.

Another problem in morphology, particularly in vertebrates, is that it is
dealing with significant legacy data, being recycled, that is based on 19th
century typological species concepts. So it needs to all be reworked into a
more modern framework and utilised in a way that can be phylogenetically
analysed under a more modern and aceted species concept. With turtles the
BSC is kind of hard to use because most turtles can and do hybridise, cross
species, cross genus, they do not care, they will breed with anything. In
the end the issue with some legacy data is the direct of the phylogenetic
signal. This is apparent in Chelid turtles, they would seem to have evolved
from a long necked ancestor not a short neck, despite there being multiple
short necks, a couple of medium necks and two long necked genera. The
morphology and the molecular data do not produce similar trees, but the
problem lies in the root of the tree. The closest living relatives of the
Family are from a parellel evolutionary path and are completely homoplassic
with respect to neck evolution with some characters, or evolved
indepenantly and differently on others.

As one who works significantly with fossils I am obviously a morphologist,
however I do work also with molecular data. Recently myself and others
weighed in on the recent Anaconda issue which was largely a nomenclatural
problem but it was created by the over simplification and over usage of
molecular data. Relying solely on mtDNA this proved highly problematic. In
our paper recommended against describing species solely on that dataset.
This is also particularly true of mtDNA genomic capture by species capable
of hybridisation (a number of species of turtles do not have their own
unique mtDNA genome, they have the genome of other species, sometimes
mixed).

In the end despite the limits of molecular data in covering the known
available species in turtles I consider molecular data essential. The
comparison of the trees of the two datasets can and often does identify
homoplasy. This forces the revisit and analysis of the morphological
characters, and as you look closer and in more detail at specific
characters, the homoplasic characters are often not so simple, nor are they
identical. Admitedly this is in animals with many complex internal
structures permitting more abundance of the characters available. I
recognise this would not be so easy in simple organisms, probably not so
relevant in groups with no (or little) fossil record.

Cheers Scott

On Thu, Oct 9, 2025 at 2:47 AM Lücking, Robert via Taxacom <
taxacom at lists.ku.edu> wrote:

> Dear Marko,
>
> for the placement of fossils, the technique described in this paper might
> be useful. It is also used for extant taxa and provides a more objective,
> quantitative and repeatable approach.
>
> "Accuracy of morphology-based phylogenetic fossil placement under Maximum
> Likelihood"
>
> Regards
> Robert
>
>
>
>
>
> -----Ursprüngliche Nachricht-----
> Von: Marko Mutanen <Marko.Mutanen at oulu.fi>
> Gesendet: Mittwoch, 8. Oktober 2025 21:02
> An: Lücking, Robert <R.Luecking at bo.berlin>; John Grehan <
> calabar.john at gmail.com>; Kuoi Zhang <zhang.guo-yi at outlook.com>
> Cc: taxacom at lists.ku.edu
> Betreff: RE: Taxacom: On describing new taxa without using phylogenetics:
> some suggestions
>
> Dear Robert, all,
>
> I agree with you. Morphological evolution is indeed fascinating, but it
> should be studied in the context of DNA-based phylogenies rather than by
> continuing to infer evolutionary relationships solely from morphology,
> which is highly prone to homoplasy. Of course, when dealing with fossils,
> we must still rely on morphological data, but this inevitably introduces
> substantial uncertainty into the taxonomy of extinct taxa - likely
> persisting indefinitely.
>
> Sincerely,
>
> Marko M.
>
> -----Original Message-----
> From: Taxacom <taxacom-bounces at lists.ku.edu> On Behalf Of Lücking, Robert
> via Taxacom
> Sent: keskiviikko 8. lokakuuta 2025 18.51
> To: John Grehan <calabar.john at gmail.com>; Kuoi Zhang <
> zhang.guo-yi at outlook.com>
> Cc: taxacom at lists.ku.edu
> Subject: Re: Taxacom: On describing new taxa without using phylogenetics:
> some suggestions
>
> Dear John,
>
> homoplasy in phenotype is much more frequent than homoplasy in DNA
> sequence data. Also, the number of characters in sequence data is larger by
> up to several orders in magnitude. So yes, I would say that there are
> substantial differences between these types of data and molecular data
> cannot be considered equal to phenotype data. That said, DNA sequence data
> are not the answer to everything and often provide conflicting results
> between different markers or for example multimarker vs. phylogenomic data.
> One always has to remain critical but unless DNA sequence data can be
> proved erroneous, I would go with the hypothesis that in case of conflict
> between molecular and morphological data, the molecular data are
> objectively more likely to show the correct relationships. E.g. crocodiles
> with birds and humans with chimps. If you get such constellations
> repeatedly with different kinds of molecular data using different
> analytical tools, it becomes really difficult to doubt or reject them.
>
> Regards
> Robert
>
>
>
> Von: John Grehan <calabar.john at gmail.com>
> Gesendet: Mittwoch, 8. Oktober 2025 17:38
> An: Kuoi Zhang <zhang.guo-yi at outlook.com>
> Cc: Lücking, Robert <R.Luecking at bo.berlin>; taxacom at lists.ku.edu
> Betreff: Re: Taxacom: On describing new taxa without using phylogenetics:
> some suggestions
>
> " rue, that's the point. But many of the new taxa are not proposed by
> monophyly and characters."
>
> Interested in an example, but even so, such entities may be falsifiable in
> the sense that subsequent analysis could show them to not be monophyletic,
> or that some such characters are insufficient to meet those standards.
>
> "I think molecule-based or morphology-based is not fundamentally
> different; they are all characters."
>
> In that sense one might say that cladistics and phenetics (overall
> similarity) are not all that different either. Might as well say that
> cladistically crocodile/alligator sister group with birds is not all that
> different from phenetically being sister group of lizards or some other
> extant reptile group.
>
> "P.S. Note, in my comments above I have avoided any reference to
> "molecular" vs. "morphological" characters as the basis for taxonomic
> analysis.  My comments above are all independent of this (somewhat dubious)
> distinction."
>
> Interested to know the context of the distinction being 'dubious' Agree
> that in general sense DNA molecules and developmental genetics (morphology)
> are both 'morphology, but they can generate very different conflicting
> results - e.g. the human-chimp clade (supported by molecules) vs the
> human-orangutan clade (supported by morphology). At present there is no
> objective way to chose between them.
>
> Cheers, John
>
> On Tue, Oct 7, 2025 at 10:24 PM Kuoi Zhang <zhang.guo-yi at outlook.com
> <mailto:zhang.guo-yi at outlook.com>> wrote:
> "disagree with the statement....all new taxa that are proposed are
> hypotheses that can be refuted and falsified" If a taxon is proposed based
> on evidence of monophyly then it would seem that the monophyly claim is
> potentially open to refutation and falsification.
>
> True, that's the point. But many of the new taxa are not proposed by
> monophyly and characters.
>
> "cladistics” is now often understood as referring specifically to
> parsimony-based approaches" I believe this is a misunderstanding.
> Cladistics, in its original form, was about grouping taxa according to
> uniquely shared features. Parisony was just one of many potential
> algorithms for making a judgement about how to balance out conflicting
> relationships indicated by different uniquely shared features. I know of
> some cladists who are not enamoured with standard parsimony methods (e.g.
> PAUP). I am not sure that molecular methods are 'cladistic' in the original
> sense, especially if uniquely shared molecular characters are only defined
> after the tree is selected.
>
> I remember there was a discussion on a blog, but anyway, it's a language
> philosophy. I followed the common usage, especially after the "parsimony
> gate", which is a choice by the "previous" Cladistics journal, Willi Hennig
> Society, and Ernst Mayr's comments to people who supported the opinion of
> Willi Hennig. I think molecule-based or morphology-based is not
> fundamentally different; they are all characters.
>
> "as fossil taxa can indeed be placed within a phylogenetic framework using
> morphological characters." Problem occurs when clades are supported by
> molecular features that are not known for fossils. For example, molecular
> folk argue for a human-chimp sister group, in which case there is no
> evidence for fossils such as Australopiths being placed within a
> human-chimp clade. Ironically there is evidence to place them in a
> human-orangutan clade. In another irony, there was a prominent
> paleoanthropologist, Pilbeam, who dismissed morphology in favor of
> molecules because he made a mistake with one of his fossil assignments, but
> then later went on to present himself as an authority on what fossils were
> hominids (i.e. within the human lineage). He had his cake and ate it too.
>
> Thanks for sharing these dramas. Paleoanthropology always has quite a lot
> of dramas. You may miss the point. The point is that someone thinks fossils
> can't be placed in phylogenetics because most of them have no molecules
> available. It's not about conflicts between morphology and molecule.
>
> As for having a phylogeny, whether morphological or molecular, not always
> possible.  For example, in my group of study (Hepialidae), there are two
> genera in S Africa, Eudalaca and Gorgopis, for which neither have any
> identifiable autapomorphy. Probably they are a mix, and maybe in the future
> prove to be a single taxon, or two or more taxa under a different
> arrangement. It is possible to find new species and see that they fall
> within one or other genus as currently circumscribed by currently named
> species, and in one of them, to compare with the type species as a
> reference point. Under this circumstance I will make the best argument
> possible for assigning a new species to one or other genus and leave it to
> future research to conform or falsify my current choice. To me this is
> better than not describing and naming these entities at all just because
> there is no resolved phylogeny.
>
>  I may disagree with you on describing species. We wrote the last
> paragraph on describing new taxa and open nomenclature, including the
> morphological code widely used in Australia.
>
> Best regards
> Guoyi
>
> Guoyi Zhang (she/her), MRes, PhD candidate Personal Web<
> https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Fmalacology.net%2F&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368353811%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=dE1HOLfGl4I37QgCeCmnNBEvfYxx1hO%2FIIIoKHmN8pM%3D&reserved=0> | OCRID<https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Forcid.org%2F0000-0002-3426-9273&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368374329%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=M3ofp%2F5xoOoaUC28lSwfKorrO3Y9E6Ddnrg7iurFiG4%3D&reserved=0>
> School of Biological, Earth and Environmental Sciences University of New
> South Wales Australian Museum Research Institute Australian Museum
>
> ________________________________
> From: John Grehan <calabar.john at gmail.com<mailto:calabar.john at gmail.com>>
> Sent: Wednesday, 8 October 2025 12:49
> To: Lücking, Robert <R.Luecking at bo.berlin>
> Cc: Kuoi Zhang <zhang.guo-yi at outlook.com<mailto:zhang.guo-yi at outlook.com>>;
> taxacom at lists.ku.edu<mailto:taxacom at lists.ku.edu> <taxacom at lists.ku.edu
> <mailto:taxacom at lists.ku.edu>>
> Subject: Re: Taxacom: On describing new taxa without using phylogenetics:
> some suggestions
>
> Hi Guyoi,
>
> Just a couple of off the cuff remarks as I have not read the paper in the
> link and am not deep into the metaphysics of taxonomy or systematics (I
> leave that to those who are).
>
> "disagree with the statement....all new taxa that are proposed are
> hypotheses that can be refuted and falsified" If a taxon is proposed based
> on evidence of monophyly then it would seem that the monophyly claim is
> potentially open to refutation and falsification.
>
> "cladistics” is now often understood as referring specifically to
> parsimony-based approaches" I believe this is a misunderstanding.
> Cladistics, in its original form, was about grouping taxa according to
> uniquely shared features. Parisony was just one of many potential
> algorithms for making a judgement about how to balance out conflicting
> relationships indicated by different uniquely shared features. I know of
> some cladists who are not enamoured with standard parsimony methods (e.g.
> PAUP). I am not sure that molecular methods are 'cladistic' in the original
> sense, especially if uniquely shared molecular characters are only defined
> after the tree is selected.
>
> "as fossil taxa can indeed be placed within a phylogenetic framework using
> morphological characters." Problem occurs when clades are supported by
> molecular features that are not known for fossils. For example, molecular
> folk argue for a human-chimp sister group, in which case there is no
> evidence for fossils such as Australopiths being placed within a
> human-chimp clade. Ironically there is evidence to place them in a
> human-orangutan clade. In another irony, there was a prominent
> paleoanthropologist, Pilbeam, who dismissed morphology in favor of
> molecules because he made a mistake with one of his fossil assignments, but
> then later went on to present himself as an authority on what fossils were
> hominids (i.e. within the human lineage). He had his cake and ate it too.
>
> As for having a phylogeny, whether morphological or molecular, not always
> possible.  For example, in my group of study (Hepialidae), there are two
> genera in S Africa, Eudalaca and Gorgopis, for which neither have any
> identifiable autapomorphy. Probably they are a mix, and maybe in the future
> prove to be a single taxon, or two or more taxa under a different
> arrangement. It is possible to find new species and see that they fall
> within one or other genus as currently circumscribed by currently named
> species, and in one of them, to compare with the type species as a
> reference point. Under this circumstance I will make the best argument
> possible for assigning a new species to one or other genus and leave it to
> future research to conform or falsify my current choice. To me this is
> better than not describing and naming these entities at all just because
> there is no resolved phylogeny.
>
> Just my two cents worth, and mostly out of ignorance no doubt.
>
> Cheers, John
>
> On Tue, Oct 7, 2025 at 1:53 AM Lücking, Robert via Taxacom <
> taxacom at lists.ku.edu<mailto:taxacom at lists.ku.edu>> wrote:
> Dear Guoyi,
>
> your point is well founded. The confusion may be in the term
> "phylogenetics". This term relates to the establishment of genealogies,
> regardless of the methods used. What you refer to is the use of
> quantitative methods, even when only morphological data are available. I
> believe the term is then "cladistics", which came long after the use of
> "phylogenetics". There are also other quantitative methods, including an
> array of multivariate analyses and phenotype-based phylogenetic binning
> (which was precisely designed for fossils but can be used for any kind of
> phenotype data).
>
> Regards
> Robert
>
>
> -----Ursprüngliche Nachricht-----
> Von: Taxacom <taxacom-bounces at lists.ku.edu<mailto:
> taxacom-bounces at lists.ku.edu>> Im Auftrag von Kuoi Zhang via Taxacom
> Gesendet: Dienstag, 7. Oktober 2025 03:52
> An: Taxacom (taxacom at lists.ku.edu<mailto:taxacom at lists.ku.edu>) <
> taxacom at lists.ku.edu<mailto:taxacom at lists.ku.edu>>
> Betreff: Taxacom: On describing new taxa without using phylogenetics: some
> suggestions
>
> Zhang G, Feng Q. 2025. Why we should not describe new taxa without using
> phylogenetics. Comment on Chen et al. (2025). Journal of Natural History
> 59(37-40): 2355-2359.
> https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Fdoi.org%2F10.1080%2F00222933.2025.2564347&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368384631%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=HbFtqIyqDAOC8pxvr7wZ%2Fnc3HamyU8zwgtkrUVOGhu4%3D&reserved=0
>
> pdf available at
> https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F396239492_Why_we_should_not_describe_new_taxa_without_using_phylogenetics_Comment_on_Chen_et_al_2025&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368394573%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=%2BdMheunzQbNgNrjP19PTES4SxSpu2e8csPg5%2BdHPRc4%3D&reserved=0
> <
> https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Fwww.researchgate.net%2Fpublication%2F396239492_Why_we_should_not_describe_new_taxa_without_using_phylogenetics_Comment_on_Chen_et_al_2025%3F_sg%255B0%255D%3DhqDEV92UeyWCOKjJ17ed1jbtmejHJ9MvHWGQaWxxhz91RNCiYocNZhGRj2BRWUdscZ9UF6XUnkfkZoKRIVT8RXi0FUkNPb4Rchgejiww.9eeTE4s8-qcLQo1Mb6y-bvvUmMI_J0sUIayNKpVofl3fGnDJw9mz4G9HP665eLzB2opS-IMV26GiwEDPg5TCgA%26_tp%3DeyJjb250ZXh0Ijp7ImZpcnN0UGFnZSI6InByb2ZpbGUiLCJwYWdlIjoicHJvZmlsZSIsInBvc2l0aW9uIjoicGFnZUNvbnRlbnQifX0&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368404271%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=HKfWRl2KO9MgzcC0N%2B8i33395qKA9BUAN40YyvfX174%3D&reserved=0
> >
>
> Under the pressure of an overwhelming number of problematic new taxa, we
> call for taxonomists to incorporate phylogenetic methods and reasoning into
> their taxonomic work. I understand that many may disagree, but please allow
> me to explain.
>
> This paper briefly summarizes why taxonomy is a science, how taxonomy can
> be a science, how the ideas and methods of Kant, Darwin, Hennig, and Popper
> have influenced modern taxonomy, and why contemporary taxonomic research
> should not be separated from phylogenetics, both from the perspectives of
> history and the philosophy of science.
>
> To reach readers who may be new to phylogenetics, we must emphasize that
> phylogenetics is not limited to molecular data. Although I belong to
> Generation Z, I am aware that phylogenetics was first widely applied to
> morphological characters long before molecular sequencing became common.
> While morphology-based phylogenetics may suffer from homoplasy, something
> is still better than nothing. Simply describing morphological patterns
> without an explicit phylogenetic framework is less informative and less
> scientific than conducting morphology-based phylogenetic analyses.
>
> I am aware of previous arguments presented in the paper "Should we
> describe genera without molecular phylogenies?", whose author strongly
> opposed our views and directly criticized our commentary during the review
> process. That paper lacked a clear understanding of phylogenetics based on
> morphological data, as also revealed during the review process of our own
> manuscript. The author wrote, "not to mention fossil taxa, which also need
> to be placed in the system obviously without molecular support." I believe
> most paleontologists would disagree with this statement, as fossil taxa can
> indeed be placed within a phylogenetic framework using morphological
> characters.
>
> In response to claims such as "we should invest more trust in the
> taxonomic evaluations of the decreasing number of taxonomists and allow
> more freedom for morphology-based grouping," I would say: "Taxonomy should
> not become theology, where people are expected to believe without
> evidence." Furthermore, I personally disagree with the statement "taxonomy
> is a science, and all new taxa that are proposed are hypotheses that can be
> refuted and falsified." A taxon described solely by a fixed morphological
> pattern, as the author often does, cannot truly be falsified, since any
> additional variation can easily be reinterpreted as representing a new
> taxon. Such direct assertions are not falsifiable, much like religious
> doctrines. When synonyms or new taxonomic acts are later proposed, it
> merely reflects a redefinition of original taxa using new assertions or
> occasionally falsifiable methods.
>
> Ultimately, this commentary only represents our perspective on taxonomic
> practice. We are also happy to receive any constructive feedback.
>
> Best regards
> Guoyi
>
> Guoyi Zhang (she/her), MRes, PhD candidate Personal Web<
> https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Fmalacology.net%2F&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368413753%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=gtlr9p4rmobgj8bllp7OWmzvE97zmVCxbFo83NrFcZ8%3D&reserved=0> | OCRID<https://nam10.safelinks.protection.outlook.com/?url=https%3A%2F%2Forcid.org%2F0000-0002-3426-9273&data=05%7C02%7Ctaxacom%40lists.ku.edu%7Ca37f8f9615d34d5f178708de0728bc92%7C3c176536afe643f5b96636feabbe3c1a%7C0%7C0%7C638956070368423448%7CUnknown%7CTWFpbGZsb3d8eyJFbXB0eU1hcGkiOnRydWUsIlYiOiIwLjAuMDAwMCIsIlAiOiJXaW4zMiIsIkFOIjoiTWFpbCIsIldUIjoyfQ%3D%3D%7C60000%7C%7C%7C&sdata=hJ89e1LPQoOFAHXaADn8Jv8AaTCivgXdfBRTZdUO6KM%3D&reserved=0>
> School of Biological, Earth and Environmental Sciences University of New
> South Wales Australian Museum Research Institute Australian Museum
>
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-- 
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Centro de Estudos dos Quelônios da Amazônia - CEQUA
Petrópolis, Manaus
State of Amazonas, 69055-010
Brasil

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