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The Freshie: Australian crocodile, seemingly from the north


(crocodiles part V)
By Darren Naish | July 31, 2012 |
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Freshw ater crocodile, photographed in captivity by Richard Fisher. Licensed under Creative Commons
Attribution 2.0 Generic license.

Its time to carry on once more with the Tet Zoo crocodile series for previous parts,
see the list of links below. In the previous article we looked at the New Guinea
crocodile Crocodylus novaeguineae and Philippine crocodile C. mindorensis, and
before that the Saltwater, Indopacific or Estuarine crocodile C. porosus.
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This last species famously occurs in


(but is not unique to) Australia where
its definitely the largest, most
formidable and most feared crocodile.
For its not the only one there:
Australia is also home to the
Freshwater or Johnstons crocodile C.
johnstoni or C. johnsoni, also called the

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Dorsal osteoderm compliment of Freshw ater crocodile,


from Ross & Mayer (1983). Note that the cervical and
dorsal shields are nearly contiguous and that there are
typically tw o row s of ovate scutes betw een the cervical
shield and the back of the skull. Greg Mayer kindly
points out that I've been citing the w rong Ross brother
w hen referencing Ross & Mayer (1983) - apologies!

Freshie [photo of captive Freshie


above by Richard Fisher]. Most people
interested in crocodilians know that G.
Krefft mis-spelt the specific name
when naming this species (in 1873)
after Robert A. Johnston; the incorrect
spelling johnsoni has been used widely
ever since. According to the ICZN,
incorrectly spelt names have to be
retained for reasons of stability so
technically C. johnsoni is right, even
though its definitely wrong. Despite
this, many authors (including most
Australian herpetologists) have used
the correct, but wrong C. johnstoni,
mostly since Cogger et al. (1983)
argued that it deserves precedence.

Its difficult today to work out which name appears more frequently in the literature,
though it has been said that C. johnstoni is most commonly applied in the
scientic and general literature (Webb & Manolis 2010, p. 66). If you work on the
basis that language exists to serve us, not the other way round, it seems most sensible
to use the wrong spelling C. johnstoni, and Ill follow many others in doing that here.
Oh, its not directly relevant to the content of this article but if youre wondering
heres how that montage of fossil crocodyliforms is coming along (for the previous
version go here). Still numerous taxa left to add

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Some of crocodyliform diversity, by Darren Naish. We'll be seeing lots more of this image later on.

The Freshie is a particularly long-snouted member of Crocodylus, with a snout thats


about three times longer than it is wide at its base. In contrast to the Saltwater croc,
the Freshie lacks unarmoured bands between its dorsal and cervical osteoderm
shields. At maximum it perhaps reaches 3 m in length. Freshies are (so far as we
know, read on) uniquely Australian, being found only in northern Western Australia,
Northern Territory and Queensland. The species is of course mostly associated with
rivers, billabongs and (during the wet seasons) flooded grasslands and forests; it
generally avoids areas frequently by the Saltwater crocodile, but the two are known to
be sympatric in places. When Saltwater crocs declined due to hunting, Freshwater

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crocs moved in, but when the Saltwater crocs recovered, Freshwater crocs retreated.
In other words, competitive exclusion seems to be in operation between the two
species. The Freshie appears to be at healthy population levels overall, though its
sensitivity to Cane toad poison is a cause for concern and the topic of ongoing research.
[Photo below by Guillaume Blanchard].

Another captive Freshie; photo by Guillaume-Blanchard, licensed under Creative Commons Attribution-Share
Alike 1.0 Generic license.

In terms of ecology, its a fish predator that uses rapid lateral strikes of the open jaws
to grab passing prey; frogs, mammals (bats and murid rodents), snakes, turtles, birds
and other vertebrates are on record as prey items as well (Tucker et al. 1996). An
ontogenetic shift from invertebrates to vertebrates occurs (Tucker et al. 1996).
The historical view

Hypothesis of crocodile phylogeny, show n in


simplified form but mostly based on Oaks
(2011). The Freshw ater crocodile is a
member of the 'reduced' Indopacific
assemblage. Photos (top to bottom) by Mo
Hassan, Davric, Herbert Ponting, Dave Hone,
Naish, W ilfried Berns. Image of New Guinea
crocodile (icon for 'reduced' Indopacific
assemblage) licensed under Creative
Commons Attribution-Share Alike 2.0
Germany license. Click to enlarge.

How does C. johnstoni fit into the crocodile


radiation? Meganathan et al. (2010) found the
Freshwater crocodile to be the sister-taxon to a
C. mindorensis + C. novaeguineae clade in
several of their trees, and Oaks (2011)
supported this position as well. Oaks (2011) put
the divergence between C. johnstoni and the C.
mindorensis + C. novaeguineae lineage in the
Serravalian or Tortonian part of the Miocene
(somewhere round about 13-11 million years
ago or so), so the Freshie might not be an
especially young species but, then again, it
might, since we dont know much, if anything,
about the extinct populations that form the
stem of its lineage. We might regard them as
separate species if only we had better fossils of
them all we have are a few fragments.

Willis & Archer (1990) described an isolated,


incomplete dentary from Pleistocene deposits at
Riversleigh that they referred to C. johnstoni. Intriguingly, its exceptionally large for
this species (preserved total length 183 mm; when complete, it would have been
several cm longer this is dentary length, not whole jaw length). This might show that
modern Freshies are unnaturally small due to human hunting pressures, or it could
show that Pleistocene Freshies were larger on average than Holocene ones. The
specimen, however, also differs from modern C. johnstoni individuals in being
relatively narrower, and in having a noticeable gap between the 5 th and 6th alveoli
(Willis & Archer 1990). Maybe these differences are inconsequential, but maybe they
hint at the presence of a distinct Pleistocene member of the C. johnstoni lineage.
If youve read the previous
articles in this series (or if you
know a lot about crocodiles

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already), youll know that the


clade that contains C. johnstoni,
C. mindorensis and C.
novaeguineae can be regarded
as a pared-down version of the
Indopacific assemblage
hypothesised in previous studies
(e.g., Brochu 2000). With the
Saltwater croc removed (as
discussed previously, it seems to
be part of the same clade as the
Siamese crocodile C. siamensis
and Mugger C. palustris), this
group can be imagined as a
reduced Indopacific
assemblage, as shown in the
simplified cladogram above.
Given that the Freshwater
crocodile is exclusively
Australian, its difficult to know
what these results mean in terms of biogeographical history. It obviously seems
sensible to conclude that the reduced Indopacific assemblage clade originated in the
Indopacific region (as, maybe, did crown-group Crocodylus crocodiles as a whole), but
did it diversify in Australasia before one lineage dispersed to the Philippines, or did
the two Australasian taxa (the New Guinea and Freshwater crocodile) reach
Australasia independently after originating further north? If the latter possibility is
true, we should expect fossil Freshwater crocodiles (or stem members of the
Freshwater crocodile lineage) to be discovered north of Australia.
From W illis & Archer (1990). The big Pleistocene Freshie jaw is
visible as the data point at extreme right.

Theres a lot left to find out.


Non-standard taxonomic proposals, and the Hoser Problem
In recent decades, the Freshie
has been universally included
within the genus Crocodylus, and
molecular and morphological data
shows it to be deeply nested
within a clade that has long been
associated with this name. For
the sake of completeness,
Galloping Freshw ater croc, photo by Adam Britton. This is a
however, its only right to note
McKinlay River Freshie - the sort suggested to represent the
new species Philas w ebbi by Wells & Wellington (1985).
that Wells & Wellington (1983,
1985) sought to reinstate the
generic name Philas, coined by John Gray in 1874, for both this crocodile and the New
Guinea crocodile. Furthermore, Wells & Wellington (1985) also suggested (as they did
with the Saltwater crocodile) that the Freshie might represent a species complex
rather than a single species. According to these authors, a Northern Territory
population (the holotype is from the McKinlay River) that reaches a smaller
maximum size than Freshies elsewhere warrants recognition as the distinct species
Philas webbi. As with the other new Australian crocodile named by Wells &
Wellington (1985) (C. pethericki), the idea that P. webbi might deserve recognition
has not been accepted by other workers. [Adjacent photo of a galloping McKinkay
River Freshie by Adam Britton; from McKinlay River Freshwater Crocodile Project
page at the invaluable Crocodilians: Natural History & Conservation.]

I should also note that Raymond T. Hoser a notorious individual who has published
tens of new taxonomic names for Australian reptiles in his self-published, strangely
written, non-technical works has also just (within the last week or two) proposed
some revisions to the taxonomy of Freshwater crocodiles as well as for crocodiles in
general (Hoser 2012a).
For starters, he argues that
Crocodylus as currently
conceived should be split into
several genera: Crocodylus
Laurenti, 1768 is retained for C.
niloticus alone, Oxycrocodylus
Hoser, 2012 is coined for the
African species C. suchus, Motina
Gray, 1844 is resurrected for the
New World species, and Oopholis
Gray, 1844 is also resurrected for
Asian and Australasian species. If
youre wondering,
Tw o of the crocodile taxa affected by Hoser's
Oxycrocodylus is named after
recommendations: Saltw ater croc (conventionally Crocodylus
porosus) and New Guinea crocs (conventionally C.
Hosers dog, Oxyuranus itself
novaeguineae). Photos by Darren Naish and W ilfried Berns.
New Guinea croc photo licensed under Creative Commons
named after the Australian
Attribution-Share Alike 2.0 Germany license.
snake. A lengthy etymology
section explains why Hosers dog, nicknamed Oxy, has provided a sterling
contribution to Australian herpetology (Hoser 2012a, p. 13).
Within Oopholis, Hoser (2012a) regards Philas as a subgenus for the Australasian
species. If you read the previous article on the New Guinea crocodile C. novaeguineae,
you might recall the discussion of distinct northern and southern forms, neither of
which have been granted taxonomic separation. Hoser acts on this, naming the
southern one O. adelynhoserae. Its named after one of his daughters (Hoser 2012a,
p. 13). [New Guinea croc photo here by Wilfried Berns.]
Among Australian crocodiles, he also names the Liverpool River crocodiles of
Northern Territory as the new species O. jackyhoserae. The etymology isnt given,
but I assume the name honours his other daughter. These crocodiles Hoser (2012a)
says that males average under 1.5 m (versus 1.7 m for an average male Freshie)
have otherwise been identified as a population of C. johnstoni. Hoser (2012a) doesnt
provide any comprehensive measurement data, nor any skeletal or molecular data
their separate species status is based on scute and scale number, small overall size,
and proportionally longer limbs. If there really is data confirming the validity of these
supposedly diagnostic features, then frankly the case for species status doesnt sound
unreasonable. Alas, the data if it exists is not presented, so all we have is an
assertion or, at best, a very poorly presented hypothesis of separate taxonomic status.

Photo of Liverpool River freshw ater crocodile, by Grahame Webb.

Id rather this didnt become the time or place to discuss what has become known as
Hoser taxonomy, but it does need to be said that Hosers works are decidedly
unprofessional, the numerous taxonomic changes he proposes are typically highly
questionable and without adequate support, and few if any working experts agree
with his proposals. Of the allegedly new Liverpool River crocodile, Grahame Webb,
their discoverer, said that They are stunted because of a lack of food; they are a clinal
variation and not a new species. Of Mr Hoser, Webb added The guys a f*****g
idiot (Im not kidding, this quote was genuinely reported in an online news article
from the Northern Territory News).
Hoser is so prolific and has self-published so many new names (including species,
subgenera, genera and tribes the count was c. 60 new taxa as of May 2012) that
hes been accused of taxonomic vandalism. The ICZN seemingly refuses to impart
any kind of blanket ban on the names Hoser proposes, since it argues that the
maintenance of taxonomic freedom that is, the right of researchers to make
taxonomic changes as they see fit, so long as they operate within the rules is
important. However, this ignores the fact that Hosers works (which arent peerreviewed, nor written in the style or tone expected of technical contributions) contain
undoubted examples of unethical behaviour (e.g., knowingly scooping other authors
because their work was preceding too slowly (Hoser 2000)) and frequently include
personal rants directed at professional herpetologists, officials, politicians and judges.
In other words, he isnt operating within the normal rules. Hoser, incidentally, writes
at length about alleged corruption in the Victorian police force and legal profession and
he has been found guilty of (and fined for) scandalising the court. Wikipedias page
on him is pretty good.
Hosers main response to those who
criticise his herpetological articles is
that they are truth haters who
reject his work due to personal
vendettas or biases. It should be
clear to any outside party that
Hosers works are woefully
inadequate as pieces of honest
scholarship: if you dont believe me,
look at any of pp. 3-13 in Hoser
(2009a) or pp. 16-18 in Hoser
Raymond Hoser, one of many photos available online. His
(2009b). It should also be noted that
venomous snakes have their venom glands surgically
removed.
his proposed nomenclatural acts are
not trivial and irrelevant if youre
interested in conservation, ethics and welfare, since many of them concern venomous
snakes a group of animals for which a stable nomenclature, accessible via a clearly
authoritative literature, is something of a must (for reasons related to law, medicine
and communication).
Im hardly the first to bring attention to the Hoser issue and I definitely wont be the
last. As youll know if you follow Hosers Australasian Journal of Herpetology, a
group of qualified herpetologists recently put together a manuscript in which they
argue why Hosers work qualifies as taxonomic vandalism, and why his output
necessitates the creation of some sort of ICZN-approved vetting system for new
herpetological names. This manuscript was leaked to Hoser, and he has published it in
full, together with a lengthy response that includes screenshots of facebook pages and

chains of email correspondence (Hoser 2012b). I think its time that as many of us as
possible stand up and denounce Hosers work. Taxonomic freedom is all very well and
good, but due process, conservatism and appropriate levels of scholarship, rigour and
evidence are essential if a researchers output is to be taken seriously.
I didnt realise wed end up here, giving that this article was meant to be about
crocodiles. More soon.
For previous Tet Zoo articles on crocodiles, see
Dissecting a crocodile
Earth: Crocodile Empire homeworld (crocodiles part I)
The once far and wide Siamese crocodile
The Saltwater crocodile, and all that it implies (crocodiles part III)
Crocodiles of New Guinea, crocodiles of the Philippines (crocodiles part IV)
Refs Brochu, C. A. 2000. Phylogenetic relationships and divergence timing of Crocodylus
based on morphology and the fossil record. Copeia 2000, 657-673.
Cogger, H. G., Cameron, E. E. & Cogger, H. M. 1983. Amphibia and Reptilia In
Zoological Catalogue of Australia. Vol. 1. Australian Government Publishing Service,
Canberra.
Hoser, R. 2000. A new species of snake (Serpentes: Elapidae) from Irian Jaya.
Litteratura Serpentium 20, 178-186.
- . 2009a. Creationism and contrived science: a review of recent python systematics
papers and the resolution of issues of taxonomy and nomenclature. Australasian
Journal of Herpetology 2, 1-34.
- . 2009b. A reclassification of the rattlesnakes; species formerly exclusively referred
to the genera Crotalus and Sistrurus. Australasian Journal of Herpetology 6, 1-21.
- . 2012a. A review of the taxonomy of the living crocodiles including the description of
three new tribes, a new genus, and two new species. Australasian Journal of
Herpetology 14, 9-16.
- . 2012b. Robust taxonomy and nomenclature based on good science escapes harsh
factbased criticism, but remains unable to escape an attack of lies and deception.
Australasian Journal of Herpetology 14, 37-64.
Oaks, J. R. 2011. A time-calibrated species tree of Crocodylia reveals a recent
radiation of the true crocodiles. Evolution 65, 3285-3297.
Ross, F. D. & Mayer, G. C. 1983. On the dorsal armor of the Crocodilia. In Rhodin, A.
G. J. & Miyata, K. (eds) Advances in Herpetology and Evolutionary Biology. Museum
of Comparative Zoology (Cambridge, Mass.), pp. 306-331.
Tucker, A. D., Limpus, C. J., McCallum, H. I. & McDonald, K. R. 1996. Ontogenetic
dietary partitioning by Crocodylus johnstoni during the dry season. Copeia 1996,
978-988.
Webb, G. J. W. & Manolis, S. C. 2010. Australian Freshwater Crocodile Crocodylus
johnstoni. In Manolis, S. C. & Stevenson, C. (eds) Crocodiles. Status Survey and

Conservation Action Plan. Third Edition. Crocodile Specialist Group, Darwin, pp. 6670.
Wells, R. W. & Wellington, C. R. 1983. A synopsis of the Class Reptilia in Australia.
Australian Journal of Herpetology 1, 73-129.
- . & Wellington, C. R. 1985. A classification of the Amphibia and Reptilia of Australia.
Australian Journal of Herpetology, Suppl. Ser. 1, 1-61.
Willis, P. M. A. & Archer, M. 1990. A Pleistocene longirostrine crocodilian from
Riversleigh: first fossil occurrence of Crocodylus johnstoni Krefft. Memoirs of the
Queensland Museum 28, 159-163.
About the Author: Darren Naish is a science writer, technical editor and palaeozoologist
(affiliated with the University of Southampton, UK). He mostly works on Cretaceous
dinosaurs and pterosaurs but has an avid interest in all things tetrapod. He has been
blogging at Tetrapod Zoology since 2006.

Follow on Twitter @TetZoo.


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The views expressed are those of the author and are not necessarily those of Scientific American.

Tags: archosaurs, crocodiles, crocodilians, crocodyliforms, herpetology, reptiles

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3 Comments

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1. Christopher Taylor
9:40 am 07/31/2012

Once again, I feel compelled to ward off the mention of


Raymond Hoser with the words of C. T. Simpson: Life is too
short and valuable to be wasted in any attempt at
deciphering such nonsense. The guy shows every sign of
being a complete arse. I dare anyone to look at his website and
not fall about laughing: its like something from Geocities ca
1997.

this quote was genuinely reported in an online news article


from the Northern Territory News

The Northern Territory News is known for being a little


quirky. And they take their crocodiles seriously, dammit.

Re Crocodylus johnsoni vs C. johnstoni: if Krefft stated that he


was naming it after Johnston, then that would seem a pretty
clear case where the change in spelling would be justified (Art.

32.5.1: If there is in the original publication itself, without


recourse to any external source of information, clear evidence
of an inadvertent error, such as a lapsus calami or a copyists
or printers error, it must be corrected. Incorrect
transliteration or latinization, or use of an inappropriate
connecting vowel, are not to be considered inadvertent
errors.) Anyone know what the original publication was?
Link to this

2. Christopher Taylor
9:49 am 07/31/2012

Scratch that: the original publication (see here) cites Mr.


Johnson. Any inference that Mr Johnston was the intended
would fail the without recourse to any external source of
information requirement, so by that measure johnstoni
would be an unjustified emendation. However, Art. 33.2.3.1
does allow for an unjustified emendation to be accepted as if it
were justified if it has entered prevailing usage.
Link to this

3. naishd
9:59 am 07/31/2012

Ah, you just beat me to it on Kreffts naming of the species.

The Hoser Problem is somewhat similar to some other miseducational efforts Ive been involved in, but its a far more
serious problem with real-world implications. Action is
desperately needed. Hoser seems to think people hate his stuff
because they just do; he cant seem to appreciate that it isnt
sufficient to look at published phylogenies and name taxa
willy-nilly whenever a genus is recovered as nonmonophyletic (there is where a lot of his new taxa come
from).

Darren
Link to this

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