Log In or Register

Subscribe & get

both print AND
tablet editions
Subscribe
Buy This Issue
Subscribe to Digital
Give as a Gift

Search ScientificAmerican.com

Subscribe

News & Features

Blogs

Multimedia

Blogs

Education

Citizen Science

Topics

About the SA Blog Network

Magazines

Choose a blog....

Tetrapod Zoology

About

Search All Blogs

Blog Network Highlights

Amphibians, reptiles, birds and mammals - living and extinct

Tetrapod Zoology Home

Products

Guest Blog

Context and Variation

Getting a Grip on the Goose

Barnacle

SciAm Beginnings: When a

Beginning is Not a Beginni

Contact

Earth: Crocodile Empire homeworld (crocodiles part I)

By Darren Naish | June 12, 2012 |
Share

Email

47

Print

ADVERTISEMENT

Prior to the epic

destruction
wrought upon their
kind by the noble
and beautiful Homo sapiens,
crocodiles had it pretty good.
One thing that Ive read several
times and found especially
annoying is the idea that
crocodiles, alligators and
gharials are the last remnants
of a once mighty empire, or
something along those lines. No,
crocodiles and other living
Highly simplified cladogram show ing one of the several
crocodilians are not
phylogenetic hypotheses proposed for crocodiles (this topology
based on Brochu 2000a, b and McAliley et al. 2006). Photos (top
anachronisms, relics or sorry
to bottom) by Mo Hassan, Davric, Naish, Thesupermat (see
below for licensing on last image).
vestiges of a former glory. Its
certainly true that living
crocodilians do have a superficial similarity to the sorts of crocodilians known from
fossils to have been around for more than 100 million years. But modern crocodilians
dont, contra all claims to the contrary, have especially long evolutionary histories and
theyre no more anachronistic or archaic than numerous other living things that we
take for granted and never consider of the past.

Most Read Posts

Most Commented Posts

Observations

Smart Headlights Let Drivers See Through

Rain and Snow
Observations

Deny This: Contested Himalayan Glaciers

Really Are Melting, and Doing So at a Rapid
Pace--Kind of Like Climate Change
Observations

In fact, crocodiles alone (that is,

totally ignoring alligators,
caimans and gharials) include

Abandoning Algebra Is Not the Answer

Observations

Is Pop Music Evolving, or Is It Just Getting

caimans and gharials) include

about 13 species that,
historically, occurred across
much of Africa, the American
tropics, southern Asia and the
Indopacific region. This
diversity and this global range
parallels or exceeds that of
many living animals often
regarded as the epitome of
modernity and success, so last
remnants of a once mighty
empire my arse.
While those of us who care
about such things will always
remain frustrated by continuing
Close to the smile of a Salw ater, Estuarine or Indopacific croc (C.
descriptions of crocodiles and
porosus). Verily, a thing of such great beauty.
kin as living dinosaurs, or as
living fossils, it is at least
becoming better appreciated that crocodilians are actually complex creatures with
interesting social lives, uncanny sensory abilities and a surprising, weird and
sophisticated anatomy.
The crocodile consensus

Louder?
Observations

Did Modern HumansNot Environmental

CatastropheExtinguish the Neandertals?

Image
of Shrimp
the Week
The Ghost
Video of the Week

A tree-ring record player

Latest Posts by Scientific American Editors
The Network Central

#SciAmBlogs Monday - rare diseases, Marley

parasite, math poetry, beginnings, self-citing
bloggers, and more.
The Network Central

Open Laboratory 2013 - submissions so far

On to a bit of phylogeny. One thing thats been universally agreed upon for a while is
that dwarf crocodiles (Osteolaemus) form the sister-group to Crocodylus [though,
read on]. And I talk of dwarf crocodiles in the plural sense since, as you might
remember from the previous crocodile article, its increasingly agreed that
Osteolaemus contains more than one species. Incidentally, please assume in this
article and those that follow that Im using the word crocodile in the specific (correct)
sense.

Observations

Abandoning Algebra Is Not the Answer

A Blog Around The Clock

The Scienceblogging Weekly (July 27th, 2012)

The Network Central

#SciAmBlogs Friday - Gene duplication, moss

sex, global food, origin of scientific journals,
genetically engineered athletes, and more.

YES! Send me a free issue of Scientific American w ith

no obligation to continue the subscription. If I like it, I
w ill be billed for the one-year subscription.
Email Address
Name

Continue

Slender-snouted crocodile (Mecistops cataphractus), photographed in captivity by Thesupermat. I originally

thought that this photo actually show ed a False gharial (Tomistome schlegelii) but a clearer dorsal view of the
same animal reveals the scute pattern of Mecistops; it is definitely not a Tomistoma (SEE BELOW ). Image
licensed under Creative Commons Attribution-Share Alike 3.0 Unported, 2.5 Generic, 2.0 Generic and 1.0
Generic license.

Latest Headlines on ScientificAmerican.com

An interesting surprise is that one species often imagined to be part of Crocodylus

the highly distinct Slender-snouted crocodile, conventionally Crocodylus
cataphractus is no such thing (that is, it isnt a species of Crocodylus), but is in fact
probably closer to Osteolaemus. The data supporting this is from nuclear and
mitochondrial genes, and it clearly shows the Slender-snouted crocodile to be as
distant genetically from Crocodylus as is Osteolaemus (Schmitz et al. 2003, McAliley
et al. 2006). This means that cataphractus cant be included in Crocodylus anymore;
at least, not if we want to keep Osteolaemus separate from Crocodylus, as we do. The
sister-group to Crocodylus thus consists entirely of African taxa; an observation that

High Trans-Fat Diet Predicts Aggression

Unnatural Selection: Muscles, Genes and Genetic
Cheats
Why Siri Is Still the Future
Running Lights: Did Expelled Stars Reionize the
Ancient Universe?
Abandoning Algebra Is Not the Answer

ADVERTISEMENT

might indicate an African origin for Crocodylus itself (Brochu 2000a) (though see
later articles in this series).
The idea that cataphractus might not
be especially close to Crocodylus is not
actually novel. John E. Gray gave this
species its own genus Mecistops
back in 1844. More recently, Aoki
(1976, 1992) argued that Mecistops
should be treated as a distinct genus,
and as a close relative of gharials and
false gharials. Indeed, the common
Slender-snouted crocodile, from Wermuth (1953). Click
to enlarge.
name African gharial has sometimes
been used for this species. That last
idea hasnt been supported by other work (both molecular and morphological);
anyway, the name Mecistops is now back in business for the Slender-snouted
crocodile.
As for the relationships within the Crocodylus species, its fair to say both that
contrasting phylogenetic hypotheses have been published, and that certain species
have been recovered in radically different positions. Despite this, its probably most
useful to imagine Crocodylus to consist of three main lineages: an Indopacific one, a
New World one, and the Nile crocodile C. niloticus (e.g., Brochu 2000a, b, McAliley et
al. 2006), as shown in the simplified cladogram above. To reiterate, and as well see,
the monophyly and content of these lineages has been challenged by various recent
discoveries.

The scute pattern present on the anterior part of the dorsal shield, and in the cervical shield, show s that the
crocodile show n above is indeed a Slender-snouted crocodile (Mecistops cataphractus), not a False gharial
(Tomistoma schlegelii). Photo by Thesupermat, licensed under Creative Commons Attribution-Share Alike 3.0
Unported, 2.5 Generic, 2.0 Generic and 1.0 Generic license. Diagrams of scute patterns from Ross & Mayer
(1983).

In this and the following few articles, were going to go through all the worlds living 13
or so crocodile species. The article continues the theme initiated with Mays

Dissecting a crocodile article. Rather than discuss the same sort of information youve
heard before, my aim is to highlight recent ideas about phylogeny, history and
diversity the sort of material that rarely makes it into the popular and semi-popular
literature. We start with the Mugger and other species that appear to be close to (but
not necessarily part of) the Indopacific assemblage.
The Mugger or Marsh crocodile: from Iran to Burma
The Mugger C. palustris, also
called the Marsh or Swamp
crocodile (very occasionally, the
Iranian crocodile), is uniformly
dark with a relatively short,
broad snout (proportionally the
broadest of any crocodile,
actually, sometimes being only
1.3 times as long as it is wide at
its base). You can sometimes
understand why a naive person
might misidentify a Mugger as
an alligator. Adult Muggers
Basking Muggers in captivity. Photo by Kmanoj, licensed under
Creative Commons Attribution 3.0 Unported license.
generally reach 4 m in total
length, with 4.5 m probably
being a maximum. Lengths of up to 8 m (Wermuth & Fuchs 1978) are reported in
older literature. As with other crocodile species, extraordinary lengths of this sort
should be considered highly unlikely: they are either mistakes or the result of
exaggeration. The Mugger is mostly associated with freshwater habitats but its
presence in coastal marshes is on record too. [Adjacent image by Kmanoj.]
Though usually thought of as an Indian animal, the Mugger also occurs in Pakistan,
Nepal, Bangladesh, Bhutan and Burma. Or, at least, it did occur in these countries,
since it now seems extinct in Bangladesh, Bhutan and Burma. The last records from
Burma come from the 1860s, the last Bhutanese ones were seen in the 1960s, and the
Bangladesh population might have lingered on until the 1990s. Occasional, rumoured
sightings and the undetermined origins of captive specimens in small collections
suggest that a few crocodiles might persist in the wild in these countries; either way,
its clear that the animal has been eradicated from part of its historical range.

Mugger photographed in India by Herbert Ponting in 1907: the photo w as originally labelled 'An Indian
alligator'. Image in public domain.

It seems little known that the Muggers distribution extends west into southern
Afghanistan and Iran. A 2007 survey estimated a population of 200-300 Muggers in
Iran while the number present in Afghanistan seems unknown. Ive been unable to
learn anything about the alleged presence of the species in Afghanistan and am keen
to know more my only source for the presence of crocodiles in that country comes
from the minutes of a 2006 Crocodile Specialist Group Steering Committee Meeting.

I think I can just about identify a

Mugger from the proportions of its
head, and overall colour and body
shape. But the best way to know that
youre dealing with a Mugger is to look
at its dorsal osteoderm compliment.
Muggers sometimes have a rather low
number of scutes in each transverse
dorsal row (four), but they can also
have six or eight. The most anterior
transverse band in the dorsal shield is
often very short (that is, consisting of
just two or four scutes) and may even
be absent altogether. A band of
granular skin then separates the dorsal
shield from the cervical one; the latter
typically consists of six reasonably
large scutes (Ross & Mayer 1983). I
should use this an opportunity to note
that dorsal scute compliment really is
the most reliable way of distinguishing
Dorsal scute compliment of a typical Mugger, from Ross
& Mayer (1983).
crocodile species other features that
you might rely on, including size, colour
and the general appearance of the jaws and teeth, are often unreliable.
The Mugger population on Sri Lanka supposedly differs from the typical form just
described in having especially broad throat scutes and six longitudinal rows of dorsal
scutes (as opposed to four). Accordingly, there have been various suggestions that it
warrants taxonomic distinction. The names C. p. brevirostris and C. p. kimbula have
both been used for this population.
The Mugger is recovered as the sister-taxon to the Indopacific assemblage in some
studies (Brochu 2000b, McAliley et al. 2006), though positions especially close to C.
porosus, C. mindorensis and C. siamensis have all been proposed on occasion. In fact,
molecular data sets often find C. palustris, C. porosus and C. siamensis to group
together (Li et al. 2007, Gatesy & Amato 2008, Feng et al. 2010, Man et al. 2011),
and the Mugger and Siamese crocodile C. siamensis form a clade in some of these
studies (Li et al. 2007, Man et al. 2011).
More soon. For previous Tet Zoo ver 3 articles on crocodilians, see
Dissecting a crocodile
As youll know if you follow me on twitter or facebook, Tet Zoo ver 2 has now been
deleted in entirety. So, goodbye lengthy articles on sebecosuchians, giant fossil
caimans, mekosuchines and a ton of other stuff.
Refs Aoki, R. 1976. On the generic status of Mecistops (Crocodylidae), and the origin of
Tomistoma and Gavialis. Bulletin of the Atagawa Institute 6/7, 23-30.
- . 1992. Fossil crocodilians from the late tertiary strata in the Sinda Basin, eastern
Zaire. African Study Monographs 17, 67-85.
Brochu, C. A. 2000a. Congruence between physiology, phylogenetics and the fossil

record on crocodylian historical biogeography. In Grigg, G. C., Seebacher, F. &

Franklin, C. E. (eds) Crocodilian Biology and Evolution. Surry Beatty & Sons
(Chipping Norton, Aus.), pp. 9-28.
- . 2000b. Phylogenetic relationships and divergence timing of Crocodylus based on
morphology and the fossil record. Copeia 2000, 657-673.
Feng, G., Wu, X. B., Yan, P., & Li, X. Q. 2010. Two complete mitochondrial genomes of
Crocodylus and implications for crocodilians phylogeny. Amphibia-Reptilia 31, 299309.
Gatesy, J., & Amato, G. (2008). The rapid accumulation of consistent molecular
support for intergeneric crocodilian relationships Molecular Phylogenetics and
Evolution, 48, 1232-1237
Li, Y., Wu, X., Ji, X., Yan, P. & Amato, G., 2007. The complete mitochondrial genome
of salt-water crocodile (Crocodylus porosus) and phylogeny of crocodilians. Journal of
Genetics & Genomics 34, 119-128.
Man, Z., Yishu, W., Peng, Y. & Wu, X. 2011. Crocodilian phylogeny inferred from
twelve mitochondrial protein-coding genes, with new complete mitochondrial genomic
sequences for Crocodylus acutus and Crocodylus novaeguineae. Molecular
Phylogenetic and Evolution 60, 62-67.
McAliley, L. R., Willis, R. E., Ray, D. A., White, P. S., Brochu, C. A. & Densmore, L. D.
2006. Are crocodiles really monophyletic? Evidence for subdivisions from sequence
and morphological data. Molecular Phylogenetics and Evolution 39, 16-32.
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.
Schmitz, A., Mansfeld, P., Hekkala, E., Shine, T., Nickel, H., Amato, G. & Bhme, W.
2003. Molecular evidence for species level divergence in African Nile crocodiles
Crocodylus niloticus (Laurenti, 1786). C. R. Palevol 2, 703-712.
Wermuth, H. 1953. Systematik der Rezenten Krokodile. Mitteilungen aus dem
Zoologischen Museum im Berlin 29, 375-514.
- . & Fuchs, K. 1978. Bestmmen von krokodilen und ihrer Hute. New York, Gustav
Fischer.
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.

More

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

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

Previous: Terrifying sex

organs of male turtles

More
Tetrapod Zoology

Post a comment | Read Comments (47)

Like

47 Comments

106

Next: The once far and

wide Siamese crocodile

Add Comment

1. himi-cat
8:19 am 06/12/2012

Is there any record of the contents of Tet Zoo v2? It would be

a tragedy to lose it forever (not least because I was in the
process of reading through the archives systematically).

A little more on-topic, your description of the dorsal scute

patterns on the Mugger crocodile suggest that theres a lot of
variability if thats the case, what rules/patterns are used to
distinguish between crocodile species? Is it simply that the
various species range of variation doesnt overlap?

himi
Link to this

2. vdinets
9:38 am 06/12/2012

Well, crocs are living fossils in the sense that their presentday diversity is much lower than it was, say, 70 mya. And their
lifestyle probably hasnt changed much, while birds, mammals
and squamates have diversified into all kinds of new niches.
Link to this

3. naishd
10:41 am 06/12/2012

Thanks for comments.

Response to himi on the fate of Tet Zoo ver 2: apart from

whatever is (temporarily) cached online, there seems to be no
record of any of it anywhere, not even behind the scenes at
wordpress. However, while the comments are essentially lost
forever, the articles themselves (and their accompanying
illustrations) are not lost I of course have everything saved
in my own files. Im not yet sure how to deal with this vast
amount of material I mostly want to post new stuff here at
Sci Am, not flood it with old content. We will see.

Darren
Link to this

4. naishd
10:50 am 06/12/2012

On intraspecific scute variation (see himis comment # 1): yes,

there is a reasonable amount of variation within species. In
cases, this might be because those species actually include
extremely distinct taxa that deserve species-level distinction.

However, the bottom line is that you need to take note of

numerous details of scute compliment into consideration
when trying to identify an individual. Muggers, for example,
may be variable with respect to number of dorsal scutes in
each transverse row and in the number of scutes present in the
post-occipital row, but, as a whole, this animal is still
distinctive in other details (e.g., presence of typically six,
relatively large scutes in close contact in the cervical shield, a
distinct break between the cervical and dorsal shields, and so
on). In these general respects, species tend not to overlap.
And we of course combine this knowledge of scute
compliment with details of skull form, overall size,
proportions, pigmentation and so on. As well see in later
articles, there are certain populations whose specific identities
and affinities remain the areas of debate.

Darren
Link to this

5. Heteromeles
10:59 am 06/12/2012

Darren, you could start TetZombie 2 on WordPress or

similar to republish the material from TetZoo 2. Or just call it
TetZoo 2.1.

A semantic question: which came first etymologically, the

mugger crocodile, or the human mugger who grabs purses?
Link to this

6. naishd
11:11 am 06/12/2012

Vlad (comment 2): I didnt know that a certain level of

diversity was anything to do with living fossil status to be
honest, I dont think its relevant.

As for the idea that modern crocodilians are conservative in

lifestyle compared to birds, mammals and so on.. this is
complicated. Its true that there are, say, fossil alligators from
the Oligocene that really dont look all that different from
modern kinds, but the same is true of (for example) waterfowl
and swifts. And whats especially interesting is that while
modern crocodilians do look superficially similar to, say,
Jurassic and Cretaceous goniopholidids it seems that any
such similarities are likely superficial: that is, the lineage
leading to all living species went through a small-bodied phase,
apparently containing taxa (e.g., Isisfordia, Susisuchus) not
necessarily similar to the classic, large, crocodile-like species
of the Mesozoic.

Darren
Link to this

7. dinosauricon
11:13 am 06/12/2012

Well there was a fair bit of croc diversity in the Late

Pleistocene, into the Holocene even actually. Land-based

Mekosuchids in New Caledonia and a host of other islands in

the region. And I think there was also a lineage of marine
gavialids in the South Pacific in the Late Pleistocene, if im not
wrong. Ill to check it out though. Not to mention the
terrestrial predatory Quinkana in Australias Pleistocene as
well.
Link to this

8. naishd
11:16 am 06/12/2012

On that fair bit of croc diversity in the Late Pleistocene

(comment 7), Tet Zoo ver 2 famously included several articles
on mekosuchines, Indopacific gharials, Aldabrachampsus
from Aldabra and Voay from Madagascar. Hmm, something
else to prioritise, perhaps.

Darren
Link to this

9. dinosauricon
11:23 am 06/12/2012

Darren, just wondering, how much is known about Quinkana

anyway? Ive been trying but I cant find much on it.

Did it play an important role as a predator in the Pleistocene

ecosystem? How big did it get anyway? Ive seen sites quoting
anywhere between 3 to 7m.Intraguild predation with
Megalania perhaps?

Would be exciting to watch a death match like that.

Link to this

10. Heteromeles
12:02 pm 06/12/2012

Id point out that bird body design has been pretty freaking
conservative since the Mesozoic. The reason is that flying
imposes strong constraints on both body shape and size.
Despite this, nobody calls birds living fossils.

Similarly, crocodilian lifestyles impose a set of substantial

constraints on morphology, so we shouldnt be surprised that
they all look similar and have for a long time. The same
constraint issues apply to say, designing an attack submarine.
No one calls subs obsolete warcraft, even though the ones now
look similar to the ones in WWI or even in the American Civil
War (unlike, say, tanks or aircraft carriers). No matter whats
inside, theyre going to all look similar, just because of what
they have to do.
Link to this

11. BrianL
12:16 pm 06/12/2012

Tet Zoo 2 completely deleted? Thats pretty dreadful news for

us and anyone seriously interested in tetrapods in general!

I can see not wanting to flood Tet Zoo 3 with old articles, so I
second the proposal for a Tet Zombie site of some sort. Might
you not allow someone you trust to post those articles there,
so that you can focus on Tet Zoo 3 yourself?

As for starting another series, now on crocodiles: Awesome as

always, but to quote Sergeant Wilson in Dads Army: Do you
really think thats wise, sir? I could imagine you losing track
of all the series you want to finish. That being said, the vesper
bat series did get finished, despite being lengthy.
Link to this

12. naishd
12:24 pm 06/12/2012

On starting another series (BrianL, comment 11): youre right,

its not wise. But my philosophy on blogging is (as in that
Simpsons episode) Do what you feel. In other words, write
(and publish) on whatever it is that grabs your enthusiasm.
Having said that, Ive learnt my lesson from past yet-to-becompleted article series, so these days I actually finish the
WHOLE SERIES _BEFORE_ I start publishing. So
crocodiles of the world: all done. Petrels of the world: also, all
done.

While were here, which series didnt get completed on Tet Zoo
ver 2? (1) Temnospondyls, (2) Gekkotans, (3) Anurans, (4)
Sacs and pouches in the heads, necks and chests of mammals
Im sure there are others. Coincidentally, I just started a Tet
Zoo To-Do-List on twitter, #TZTDL.

Darren
Link to this

13. Andreas
Johansson
1:03 pm 06/12/2012

Speaking of Pleistocene diversity, I suppose one could, on

that count, call the last surviving Homo species a living fossil.

But enormous diversity doesnt stop many people from

thinking anurans or squamates somehow archaic, so I suspect
the real driver of the perception is a progressivist view of
evolution where amphibians are succeeded by reptiles who are
succeeded by mammals. Any offshoots from the trunk from
primitive vertebrates to familiar mammals that fail to go
extinct tend to be regarded as relicts (birds being the big
exception.)
Link to this

14. BrianL
2:01 pm 06/12/2012

If we assess living fossil-status by phylogenetic decline since

the Pleistocene, is there any living clade that is not composed
of living fossils? To hell with asking a genie in the lamp to
grant you the wish of perpetual wealth, a cure to cancer or
world peace: Ask him to bring back healthy populations of all
species that went exist because of human impacts, except ones

that cause diseases! Fascinating though the subject is, it tends

to be bad for my mood to think of the sheer number of species
lost since the late Pleistocene because of anthropogenic
causes.

Regarding crocodiles, is there any possibility that

*Crocodylus* might someday reconquer the Mediterranean
via the Suez Canal? I suppose its unlikely.

As for *Mecistops*, Diergaarde Blijdorp in Rotterdam has two

individuals whove been there since before World War 2. They
actually survived the German bombardments on the city (and
the zoo). Theyve laid eggs in the past, but have never bred
despite being over 80 years old by now. I used to see them at
least once a week and the two seemed to be still going strong.
Its been nearly a year since I last got to see them though.

Beautiful creatures and very distinctive, as crocodiles go.

While theyre not overly big (About 2.5 to 3 meters, I guess),
especially the larger individual has a pretty impressive width,
both absolutely and compared to the adjacent Nile Crocodiles.
Is *Mecistops* truly broadbodied compared to other
crocodiles or is this an illusion created by the narrow skull? Or
would this just be individual variation or physiology at work?
The one in the top picture seems to be nothing special in that
regard, but the lower one does seem to have a broad body.
Link to this

15. vdinets
3:27 pm 06/12/2012

Darren: there is no official definition of living fossil as far as

I know. My impression was that the taxa usually called living
fossils (i. e. Nautilus, Monoplacophora, Tuatara, Latimeria
etc.) are just that: descendants of formerly more diverse
groups, similar to their ancestors in overall appearance. They
dont have to look identical, just similar. And they dont have
to be barely alive: Nautilus, for example, has a few species and
seems to be doing generally well in its extensive range. But
birds, for example, are too diverse to qualify, and extant
Monotremes are too modified. Of course, all these criteria are
very arbitrary, and there is no consistency in use: Ive seen
rhinos called living fossils simply because they look ancient.
Link to this

16. vdinets
3:53 pm 06/12/2012

BrianL: AFAIK, there are no coastal croc populations in the

Red Sea (there are some in Kenya, but thats a bit far away). A
more likely route for recolonizing the Mediterranean would be
through the Nile (there are a few crocs above the Aswan Dam),
but for that to happen, Egypt would have to become
depopulated and the Dam would have to fail. And for that to
happen, Israel would have to elect a really, really hawkish
prime minister
Link to this

17. John Harshman

4:31 pm 06/12/2012

You make a slightly wrong statement near the beginning, when

you claim that Osteolaemus is the sister group of Crocodylus,
but then you fix it right after that. Since the idea that
Mecistops is sister to the former is so new, I dont know if it
would widely be considered settled (though I believe it), so it
isnt clear what single topology in that neighborhood you
could reasonably claim as the consensus, other than that
Osteolaemus and Mecistops, whatever their mutual
relationships, are the two closest relatives of Crocodylus.

I hope you intend to deal with (I mean, have dealt with) the
ages of taxa and the problems that causes for the inclusion of
various fossils in various extant families. Once you get away
from extant species, the phylogeny of
Crocodylia/Crocodylomorpha/whatever becomes
problematic.
Link to this

18. John Harshman

4:35 pm 06/12/2012

Could we consider a relic as a taxon that had a much higher

disparity in the past? There are, for example, currently no
terrestrial crocs. There have been times in the past in which
crocs occupied a much wider range of lifestyles than
currently. Especially if you consider them as the inheritors of
all the crododylian total group, which I hesitate to try to
assign a name to. (Does it really have to be Pseudosuchia?)
Link to this

19. naishd
5:06 pm 06/12/2012

Loving these comments cant respond to everything of

course. On the sister-group relationship between Osteolaemus
and Crocodylus (comment 17), what I meant to do there was
explain the status quo prior to the new discoveries about
Mecistops. Hopefully its clear now.

As for fossil crocodilians Im aiming to work through them in

time. But, then, I aim to do a lot of things.

Darren
Link to this

20. Bret Newton

6:25 pm 06/12/2012

In my view, the only animal you could call a living fossil is

something like Catagonus. An animal known from the fossil
record before it was know to be living by scientists.
Link to this

21. LeeB 1
6:58 pm 06/12/2012

Tet Zoo v 2 is not even stored on the internet wayback

machine because it disallowed robot crawlers from copying it.
Really clever.
And regarding fossil crocodiles, hopefully you plan to cover

the recently named large fossil Crocodylus species from East

Africa and their relationship to living African crocodylus
species.

LeeB.
Link to this

22. HowardRichards
7:37 pm 06/12/2012

I believe Pooh-Bah had the correct response to such

comments: Oh, my protoplasmal ancestor!
Link to this

23. Mythusmage
8:16 pm 06/12/2012

Have any crocodilian remains been found in Iraq? Because the

dragons of the Ishtar Gate had to come from somewhere.
Link to this

24. Christopher Taylor

10:12 pm 06/12/2012

Im not yet sure how to deal with this vast amount of material
I mostly want to post new stuff here at Sci Am, not flood it
with old content.

Darren, you could start TetZombie 2 on WordPress or

similar to republish the material from TetZoo 2. Or just call it
TetZoo 2.1.

Ahem darrennaish.blogspot.com is still sitting there

Link to this

25. Heteromeles
10:52 pm 06/12/2012

Personally, I tend to agree with one friend who said that the
only living fossils are exactly that: living and fossilized. I
think those putative living bacteria from amber-preserved
bees might count, but certainly no metazoan that Im aware of
qualifies.
Link to this

26. vdinets
11:12 pm 06/12/2012

Bret: interesting suggestion but the prevailing use in popular

literature is more or less what I described, and I dont think its
worth fighting with. As for scientific literature, I wouldnt
recommend using it at all
Link to this

27. llewelly
11:46 pm 06/12/2012

Zombie Gorgosaurus: The original living fossil.

Link to this

28. Dartian
1:59 am 06/13/2012

Darren:

Tet Zoo ver 2 has now been deleted in entirety

The most appropriate response to that rhymes with clucking

bell! Is the Shiny Digital Future here yet?

On-topic: How ecologically similar or dissimilar is the

mugger to the Nile crocodile*? A priori, one might naively
assume these species to be each others opposite numbers in
their respective continents, but their morphological
differences (e.g., regarding snout shape) seems to suggest
otherwise. Am I right to suspect that the Nile crocodile is,
ecologically speaking, more of a generalist than the mugger is?

* Sensu lato; Im aware of recent suggestions to split the

extant Nile crocodile into two different species but I
suppose youll be covering that ground in some future part of
this article series.
Link to this

29. Heteromeles
10:15 am 06/13/2012

Now that its a few days later, perhaps we can start The Legend
of TetZoo 2, The Lost Library.

Personally, I think that the best way to do this is (as with the
Library of Alexandria project) to rebuild TetZoo 2 somewhere
else, all the while wailing about how the comments on the new
version just werent as good as the comments on the original.
But then again, Im not nearly as busy as Darren.

Maybe someone can start a fan site, with bootleg essays?

Link to this

30. naishd
10:31 am 06/13/2012

Re: comment 28 (how similar is the Mugger to the Nile

crocodile, in ecological terms) the Mugger seems to be a
generalist, with everything from insects, molluscs and fish to
frogs, birds, snakes and such mammals as dogs, leopards,
deer, goats, and antelopes being recorded as prey items. Fish
and insects might make up the bulk of its diet, and large
animals are of course over-represented in popular accounts
since the eating of such prey is more interesting. They do
attack and kill people on occasion: Vyas (2010) reported 14
attacks, six of which were fatal, between 1987 and 2007.

In general terms, it seems that Mugger and Nile crocs are

similar overall in terms of ecology and diet. Nile crocodiles
are of course covered in the series.. but I have to get through
the members of the Indopacific assemblage (and its hangerson) first.

Darren

Ref -

Vyas, R. 2010. Mugger (Crocodylus palustris) population in

and around Vadodara City, Gujarat State, India. Russian
Journal of Herpetology 17, 43-50.
Link to this

31. vdinets
11:58 am 06/13/2012

I wonder if muggers more massive head compared to Nile

croc(s) has something to do with non-softshell turtles being
historically more abundant in Asia than in Africa.
Link to this

32. Andreas
Johansson
1:16 pm 06/13/2012

A semantic question: which came first etymologically, the

mugger crocodile, or the human mugger who grabs purses?

According to the Online Etymological Dictionary, the later is

etymologically someone who punches people in the face, from
mug (as in mug shot), in turn, by some bizarre turn of
slang, from mug = drinking vessel.

Mugger the crocodile OTOH is from an Indic word for

crocodile.
Link to this

33. Heteromeles
2:31 pm 06/13/2012

Another beautiful etymology, slain by cold reality. Oh well.

Thanks Andreas!
Link to this

34. Hai~Ren
12:25 am 06/14/2012

The mugger crocodile is the species featured in The

Undertakers, one of Rudyard Kiplings stories in the Second
Jungle Book:

It was a twenty-four-foot crocodile, cased in what looked

like treble-riveted boiler-plate, studded and keeled and
crested; the yellow points of his upper teeth just overhanging
his beautifully fluted lower jaw. It was the blunt-nosed
Mugger of Mugger-Ghaut, older than any man in the village,
who had given his name to the village; the demon of the ford
before the railway bridge, camemurderer, man-eater, and
local fetish in one. He lay with his chin in the shallows,
keeping his place by an almost invisible rippling of his tail,
and well the Jackal knew that one stroke of that same tail in
the water would carry the Mugger up the bank with the rush
of a steam-engine.
Link to this

35. vdinets
2:06 am 06/14/2012

Hai-Ren: I used the opening lines of that story as an epigraph

to my dissertation on croc songs
Link to this

36. Dartian
3:22 am 06/14/2012

Darren:
the Mugger seems to be a generalist, with everything from
insects, molluscs and fish to frogs, birds, snakes and such
mammals as dogs, leopards, deer, goats, and antelopes being
recorded as prey items

I knew that (the source for leopard as a prey item is Dunbar

Brander from the 1920ies, right?), but those are anecdotal
data. Are there any modern, systematic comparative studies
of the dietary habits of these crocodilian species?

it seems that Mugger and Nile crocs are similar overall in

terms of ecology and diet

But similar, of course, does not necessarily equal same. (A

jaguar is very similar to a leopard in many respects, but its
different from it in others.) Let me re-phrase my original
question:

We know for a fact that there are morphological differences

(notably regarding snout shape) between palustris and
niloticus. Are these morphological differences related to
differences in ecology or behaviour between these crocodile
taxa?

Diet is, of course, an obvious possible explanation here. Thus,

its interesting that there are notable differences between the
respective large freshwater predator guilds in Africa and the
Indian subcontinent. In the former, there is, for the most part,
just the Nile crocodile (true, there are other extant crocodile
species in Africa too, but they are much smaller than niloticus
and please correct me if Im wrong they typically do not
occur in sympatry with it). In India, by contrast, the mugger at
least locally occurs in sympatry with the fish-eating gharial
(and also with the equally fish-eating river dolphins
Platanista). To me, that seems like a textbook case of a
situation where evolutionary character displacement that is,
specialisation might occur. In order to avoid competition
with gharials and river dolphins, there might therefore
conceivably be selection against fish-eating in the mugger; are
there dietary studies suggesting that this is, in fact, the case?

Finally, if the mugger and the Nile crocodile do not

significantly differ ecologically despite occupying different
positions in morphospace, what are the implications of that
observation for the reconstruction of the lifestyles of

dinosaurs and other extinct archosaurs? If we observe cranial

shape differences of similar magnitude between (say)
Albertosaurus and Daspletosaurus, must we then conclude
that those differences do not inform us anything about their
possible differences in ecology?
Link to this

37. naishd
4:42 am 06/14/2012

Dartian (comment 36): great questions, but I dont really know

the answers as I havent seen ANY detailed work on Mugger
ecology/dietary preferences, partly because not that much
seems to be published (I mean, as goes detailed study). If
anyone knows of such work, please let us know. Actually,
there is

Kumar, V., Choudhury, B. C. & Soni, V. C. 1995. Dietary habits

of the mugger (Crocodylus palustris) in Andhra Pradesh,
south India. Hamadryad 20, 8-12.

Whitaker, R 1976. Status and ecology of the marsh crocodile

in south India. Madras Snake Park Publication. Field Report
1976, 1-5.

but I havent seen them.

The impression I get is that, as noted above, invertebrates and

fish form the bulk of the diet as they do in many other
Crocodylus crocodiles, but this doesnt necessarily mean
overlap with gharials or river dolphins since both presumably
feed on smaller fish. Muggers and Gharials are truly
sympatric, even occurring along the same river stretches in
some regions.

Darren
Link to this

38. Dartian
5:22 am 06/14/2012

Darren:
this doesnt necessarily mean overlap with gharials or river
dolphins since both presumably feed on smaller fish

Good point; there are, of course, different kinds of piscivory,

too.

I wonder if muggers tend to scavenge more often than other

species of crocodiles? They certainly seem to come fairly
often into conflict with tigers over this (at least if nature
documentaries are anything to go by).
Link to this

39. Brad McFeeters

3:17 pm 06/14/2012

It seems little known that the Muggers distribution extends

east into southern Afghanistan and Iran.

Very little known, because those countries are west of India.

Link to this

40. vdinets
3:24 pm 06/14/2012

Dartian: Nile crocs (especially C. suchus) are sympatrical with

fish-eating, narrow-snouted Mecistops over much of their
range. There isnt much habitat separation, and Mecistops
isnt much smaller than C. suchus. The size and habitat
differences with Osteolaemus are more pronounced, although
recently the latter have moved into areas where Nile crocs
have been exterminated (i. e. mangroves of Cameroon).
However, to my knowledge there is no difference in snout
morphology between Nile crocs in Western/Central Africa
(the area of sympatry) and Eastern/Southern Africa.
Also, all crocs (even gharials) would scavenge given a chance,
and I would be surprised if muggers are more enthusiastic
about it than Nile crocs.
Link to this

41. David Marjanovi

5:44 pm 06/14/2012

Response to himi on the fate of Tet Zoo ver 2: apart from

whatever is (temporarily) cached online, there seems to be no
record of any of it anywhere, not even behind the scenes at
wordpress.

What

the fuck.

Perhaps thats some bizarre temporary byproduct of

overhauling the site and restoring the comments. If not, raise
a fucking stink.

I didnt know that a certain level of diversity was anything to

do with living fossil status

If its diverse now, its not a living fossil. Thats a criterion Ive
seen again and again and again.

And I think there was also a lineage of marine gavialids in the

South Pacific in the Late Pleistocene

What, that late!?!?!

the crododylian total group, which I hesitate to try to assign

a name to. (Does it really have to be Pseudosuchia?)

As Ive said before, the PhyloCode is against Pseudosuchia

(scroll down to the bottom of the page) and for an unrestricted
emendation of Crurotarsi.

the dragons of the Ishtar Gate had to come from somewhere

You think so? Take a lizard, blow it up in your imagination,

add a few body parts from other animals

the face, from mug (as in mug shot), in turn, by some

bizarre turn of slang, from mug = drinking vessel

Its not particularly bizarre to liken the head to an empty

vessel. See noggin and German Kopf, which is from Latin
cuppa, which is also where cup comes from. Further see
French tte and Italian testa, from Latin testa shell.
Link to this

42. David Marjanovi

5:47 pm 06/14/2012

an unrestricted emendation of Crurotarsi

er, what would be an unrestricted emendation if the

PhyloCode were already in effect and Crurotarsi had already
been established (with its current definition, which includes a
phytosaur or maybe the group as a whole as an internal
specifier).

Dont hold back. Just use Crurotarsi and specify once how
youre using it.
Link to this

43. naishd
7:28 pm 06/14/2012

Comment 39

Very little known, because those countries are west of India.

Dammit, why did no-one point this out earlier? Thanks.

Darren
Link to this

44. BilBy
9:31 pm 06/14/2012

@Hai-Ren although it is not up there with Kipling, Harry

Flashman meets some crocodiles when running away from

Indian Mutiny trouble in Flashman and the Great Game he

rather testily reports that he couldnt identify the species.
Also, Darren, I hope when you get on to C. niloticus you will
be mentioning the book Eyelids of Morning?
Link to this

45. Dartian
4:59 am 06/15/2012

If were listing appearances of palustris in Western fiction,

lets not forget those, ahem, slightly American alligator-like
mugger crocodiles seen briefly in Indiana Jones and the
Temple of Doom
Link to this

46. Mythusmage
9:20 pm 06/15/2012

David,

The dragons look a lot like what a crocodile would look like if
you didnt know what a crocodile looked like. Besides, the
marshes of Mesopotamia looked like prime crocodile ground.

In any case, you didnt answer the question.

Link to this

47. David Marjanovi

5:38 pm 06/16/2012

The dragons look a lot like what a crocodile would look like if
you didnt know what a crocodile looked like.

Good. How about tales of crocodiles from Egypt? Contact with

Egypt was established pretty early on.

In any case, you didnt answer the question.

Thats because I cant, not knowing the answer and all.

However, if crocodiles had lived there in historical times, Id

probably know it. Maybe the ice ages drove them out, and
then they couldnt come back across the desert or the sea?
Link to this

Add a Comment
You must log in or register as a ScientificAmerican.com member to submit a comment.

Scientific American is a trademark of Scientific American, Inc.,

used w ith permission

Advertise

About Scientific American

Subscribe

Special Ad Sections

Press Room

Renew Your Subscription

 2012 Scientific American, a Division of Nature America, Inc.

All Rights Reserved.

Special Ad Sections

Press Room

Renew Your Subscription

Science Jobs

Site Map

Buy Back Issues

Partner Netw ork

Terms of Use

Products & Services

International Editions

Privacy Policy

Subscriber Customer Service

Use of Cookies

Contact Us