Identifying Pleistocene animals from Malta, courtesy of

It's well known that the islands of the Mediterranean were formerly home to an assortment of island endemics, all of which are now extinct. Most of the best known ones are mammals like pygmy elephants, pygmy hippos, pygmy megacerine deer and giant dormice, but there were also large birds, tortoises and lizards. My excellent friend Bob Nicholls of has been kind enough to share this wonderful piece of art, featuring extinct and extant animals of Pleistocene Malta (close-ups below the fold). It's used with permission and is © Robert Nicholls.

One thing I particularly like about Bob's painting is that it shows various of the small herps known from the Maltese fossil record. In the foreground, you can see (left to right) a Mediterranean painted frog Discoglossus pictus, European pond terrapin Emys orbicularis and Green toad Pseudepidalea viridis. Ah, toads - have I mentioned them on Tet Zoo before? An unfortunate anuran is being eaten by another animal in the painting. I don't need to say, of course, that people worldwide are universally familiar with Bob's ability to juxtapose the anurans that we know and love with extinct megafauna: look at the cover of the late J. Alan Holman's Fossil Frogs and Toads of North America, previously shown here.

Anyway, identifying still-extant amphibians and turtles is easy, but what about the other animals shown in this scene? Test your skills and see if you can find and identify them. The fact that I've already identified the scene as a Maltese one will make some of the animals relatively easy to identify, and I'll give you one more clue: there are two proboscideans here. Part of the illustration might be a bit of a homage to ZdenÄk Burian.

Bob's excellent work has been featured on Tet Zoo before: go here for some Mesozoic marine reptiles, here for his illustrations of the giant Jurassic pachycormid fish Leedsichthys, and here for a fanciful take on the spinosaurid Irritator.

For previous articles on extinct, Pleistocene or Holocene island-endemic animals of the Mediterranean and elsewhere, see...

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I don't know about the scientific names, but the swan is the Giant Swan and the tortoise is the Malta analouge of the Galapagoas Tortoise and that elephant (the one by the storks, which I presume to be a fossil species related to the Wood Stork) is the famous Pygmy or rather Maltese Elephant. The rest I am alittle shady on.

I'll give you one more clue: there are two proboscideans here

Elephas falconeri (to the left) and two Elephas mnaidiriensis. But were those two ever actually contemporary on Malta?

The swan is the giant extinct Cygnus falconeri, and the two wading birds are probably supposed to be the extinct crane Grus melitensis. And the little furry animal sitting (dangerously exposed) on the dead tree branch to the right is surely one of those endemic dormice?

Elephas falconeri (to the left) and two Elephas mnaidiriensis.

Upon reflection, perhaps it's more likely the other way round though: the animal on the left is E. mnaidiriensis and the two on the right are E. falconeri. The animal on the left 'looks bigger' and E. mnaidiriensis was the larger-bodied taxon; besides, the Rules of Fossil Animal Reconstruction dictate that the most famous species (in this case, E. falconeri) should be featured the most prominently in these kinds of illustrations.

That lizard is surely the Ocelated lizard, right?

By Tim Morris (not verified) on 14 Jan 2011 #permalink

Sorry, Dave - I hadn't realised that you'd used the same pic as well. And there was me thinking that I had an exclusive...

Apparently, the giant tortoise is Geochelone robusta, the swan is Cygnus falconeri. The Maltese crane is Grus melitensis. Not included in this painting: the dwarf hippopotamus (Hippopotamus melitensis) and Maltese otter (Lutra euxena).

Is Elephas mnaidriensis definitively known from Malta? Was under the impression that it was Sicilian. And there's E. melitensis, which might be synonymous with E. falconeri...

I wonder if that big green lizard is supposed to be the extinct Lacerta siculimelitensis? Tortoise is Geochelone robusta I think.

The, erm, giant swan is obviously the Giant Swan (c. falconeri). I will say (based on tusk shape) that the leftmost elephant is E. falconeri (although I would've gone with Mammuthus lamarmorae had you not specified Malta) and the right ones E. mnaidiriensis. No idea on which dormouse though.

Sorry, Bob had handed it over some time ago. Still, since it's not especially Mesozoic Archosaur-based this is probably a better place for it! And since you were (quite rightly) bigging up Bob's awesome work, it didn't seem totally inappropriate to self link! ;-) And I certainly have nothing to add when it comes to identifying Maltese mammals....

By Dave Hone (not verified) on 14 Jan 2011 #permalink

So wait, how large are we talking for a giant swan, and how small for a pygmy elephant? I'm having trouble wrapping my brain around the dimensions of these beasts.

Also, I love the swampy atmosphere of the painting, really gives it that hazy heavy moisture-laden air feel. But why does the moon always seem to be in the background of prehistoric paintings?

Yonk -

Elephas falconeri was about 0.9 meters (~3') at the withers whereas Cygnus falconeri had a 3 m wingspan and bill-tail length of ~2 m (around 1/3 larger than mute swan).

Geochelone robusta; the big tortoise
Lacerta siculimelitensis; the quite big lizard
Grus melitensis; the crane
Cygnus falconeri; the (too much, maybe)big swan
Hypnomys (?) or more probably Eliomys Maltamys(?) Gliridae (I dont'n know the english name of "ghiri" e "quercini").
Palaeoloxodon (or more corectly Elephans) falconeri; the very little elephant
Paleoloxodon (or Elephans) mnaidriensis; the bigger little elphant, (olso know in Sicily).



Ä. With a pointed, V-shaped háÄek, not a round breve.

Is Elephas mnaidriensis definitively known from Malta? Was under the impression that it was Sicilian.

With a name like that, it has to be from Malta.

By David MarjanoviÄ (not verified) on 14 Jan 2011 #permalink

Dormouse! Of course! Sorry

Many species once considered endemic to Malta are endemics shared with Sicily. The only problem is that, as I understand, E. falconeri was not contemporary with E. mnaidriensis, but I could be wrong.

Leithia melitensis (Adams, 1863) is realy big, the biggest dormous I think. Too big to be one of the picture

A bit of literature on the fossil mammals of Sicily (mostly in Italian)

Accordi B. (1972) - Gli elefanti nani del Quaternario in Sicilia. Le Scienze Quaderni. (1988) 49, 88-95.
Ambrosetti P. (1968) - The Pleistocene dwarf elephants of Spinagallo (Siracusa, South-Eastern Sicily). Geol. Rom. 7, 277-398.
Azzaroli A. (1990) - Lezioni di Paleontologia dei vertebrati. 375 pp., Ed. Pitagora, Bologna.
Azzaroli A. (1982) - Insularity and its effect on terrestrial vertebrates: evolutionary and biogeographic aspects. Proceedings of the 1st International Meeting on âPalaeontology, Essential of Historical Geologyâ, Fondazione Giorgio Cini, Venezia 2-4 June 1981, pp. 193-213, S.T.E.M. Mucchi.
Bonfiglio L., Marra A.c., Masini F. & Petruso D. (2001) - Depositi a vertebrati e ambienti costieri pleistocenici della Sicilia e della Calabria meridionale. Biogeographia 22, 29-43.
Caloi L., Kotsakis T. & Palombo M.R. (1986) - La fauna a vertebrati terrestri del Pleistocene delle isole del Mediterraneo. Geol.Rom. 25, 235-256.
Caloi L., Kotsakis T., Palombo M.r. & Petronio C. (1996) - The Pleistocene dwarf elephants of Mediterranean islands. In: Shoshani J. & Tassy P. Eds., The Proboscidea. Evolution and Palaeoecology of Elephants and Their Relatives,234-239,Oxford Science Publications.
Kotsakis T. (1996) - I micromammiferi. In: In Basile B. & Chilardi S. Eds., Siracusa. Le ossa dei giganti: lo scavo paleontologico di Contrada Fusco, 68-71, Lombardi Editore.
Masini F., Bonfiglio L., Petruso D., Marra A.c., Abbazzi L., Delfino M., Fanfani F. & Torre D. (2002) - The role
of coastals areas in the Neogene-Quaternary mammal island populations of the central Mediterranean. Biogeographia 23,
Musi Nicolussi C. (1971) - Biometria di molari elefantini di varie specie conservati nellâIstituto geologico universitario di Padova. Mem. Acc. Patavina SS. LL. AA ., Cl. Sc. Mat. Nat. 79, 204-223.
Petruso D. (2002) - Il contributo dei micromammiferi alla Stratigrafia e Paleogeografia del Quaternario continentale siciliano. Tesi di Dottorato di Ricerca in Geologia del Sedimentario XIV ciclo. Università degli Studi di Napoli âFederico II â. 315 pp.
Piccoli G. & Del Pup G. (1967) -I resti di elefante nano Elephas falconeri della grotta âLuparelloâ (Palermo). Mem. Acc. Patavina SS. LL. AA ., Cl. Sc. Mat. Nat. 79, 243-260.
Rinaldi L. (2002) - Dal mito alle faune insulari della collezione paleontologica del Museo di Geologia e Paleontologia dellâUniversità di Padova: due percorsi didattici. Tesi di Laurea inedita, 51 pp., 2 all., Padova.
Kotsakis T. (1986) - Vertebrati insulari e Paleogeografia: alcuni esempi. Boll. Soc. Pal. It. 24 (2-3), 225-244.


By valerio peverelli (not verified) on 14 Jan 2011 #permalink

Was Cygnus falconeri still able to fly, or was it flightless?

By gray Stanback (not verified) on 14 Jan 2011 #permalink

Maltese otter is Lutra euxena? Shouldn't that be a Black Sea otter? did the namer get really confused? am I really confused?

By David Houston (not verified) on 14 Jan 2011 #permalink

pygmy elephants, pygmy hippos, pygmy megacerine deer

But Darren, how do you explain all of these PYGMIES + DWARFS??


By Michael O. Erickson (not verified) on 14 Jan 2011 #permalink

Is the large bird in the center an ancestor of the swan? (Can't believe how huge they were...)

I'm amused by all these Maltese falconeri-s.

By Adam Fuller (not verified) on 15 Jan 2011 #permalink

@Adam Fuller FTW! Why didn't I see those two words juxtaposed?


By Noni Mausa (not verified) on 15 Jan 2011 #permalink

I miss the small goose-like swan Cygnus equitum, and also the "top predator" (top scavenger?) Gyps melitensis. Admittedly the latter was not a Malta endemic, but then neither was Cygnus falconeri, Elephas falconeri or Elephas mnaidrensis, since they all occurred on Sicily.
And where is Pitymys melitensis? Though you might argue that it was later than the other species in the picture, most of which may be approx. last interglacial in age.
Elephas mnaidrensis is named after Mnaidra gap, a well known maltese fossil site.

By Tommy Tyrberg (not verified) on 16 Jan 2011 #permalink

On further thought, the rodent on the right might be Maltamys, it seems too small for Leithia. The swan is of course falconeri, which was very large (3 meter wingspan) and flightless, or nearly so. It was a very large member of the whooper group, so the bill is probably correctly reconstructed.
The really small elephant would be Elephas falconeri and the somewhat larger Elephas mnaidrensis. However these are different ages, falconeri being older, and I don't think they have ever been found together in a stratigraphically secure context.

By Tommy Tyrberg (not verified) on 16 Jan 2011 #permalink

Gotta love island endemics.

One thing that intrigues me, given the pervasiveness of human-mediated megafaunal extinctions worldwide, is to what extent African/South Asian faunas may have been altered by humans/hominids. Comparing today's Africa to late Pleistocene N. America or S America- the latter, classic cases of hominid-mediate extinction- and Africa today still seems rather impoverished.
I tried google and didn't really find the right combo of keywords to find peer-reviewed reports, but did find this:
It is a nonscientific tract, but does compile a number of different natural history facts (and is fairly readable and not too crackpottish).
Also, the somewhat compelling case of the extinction of African giant tortoises- if you read this you must start halfway down page:

@#24: I think it was this blog that first alerted me to how tiny the present range of orangutans is compared to their range in the early Holocene (IIRC). Plus there's the issue of how obligatorily arboreal they really are.

By Jenny Islander (not verified) on 16 Jan 2011 #permalink


In any case, I used to fantasize about having my own personal super-powerful doohickey (found in an alien space pod or on a beach somewhere or in a dead wizard's pocket) that could blink me from our Earth to the present potential Earth generated by the extinction of H. sapiens circa 70,000 BP. If I actually had one, I would most definitely save up for tickets to Malta. I just wouldn't be taking the same tours as everyone else. Go to the beach during siesta time (because beaches are most likely to be at the same level and everything as their counterparts in the other universe). Duck around a handy rock, blink to the beach in the other universe, and see whether the local elephants were naive enough to approach. Resist the temptation to collect some feathers from molting giant swans . . .

By Jenny Islander (not verified) on 16 Jan 2011 #permalink


I'm having trouble wrapping my brain around the dimensions of these beasts.

As Cameron said, the smaller elephant, Elephas falconeri, had a shoulder height of 0.9 m. In other words, IRL it was about as tall as the very tallest domestic dogs (e.g., Great Danes or Irish Wolfhounds).

David H.:

Maltese otter is Lutra euxena?


Shouldn't that be a Black Sea otter?

What's that? I've never heard of such a species.


the "top predator" (top scavenger?) Gyps melitensis

For the record, there were also large mammalian terrestrial predators on Malta in the Pleistocene. Fossil remains of both wolves Canis lupus and bears Ursus cf. arctos (as well as foxes Vulpes sp.) have been found at the Ghar Dalam site. In other words, these carnivores were contemporary with at least some of those 'pygmy' elephants - some fossil elephant bones that have been discovered from these deposits even have carnivore tooth marks on them (Hunt & Schembri, 1999).

That pygmy elephants could evolve/survive in the presence of large predators may sound surprising, but perhaps it isn't really all that remarkable. It's probably a mistake to think that these small proboscideans were easy prey and pushovers. Even though they were tiny by modern elephant standards, they were still fairly sizeable beasts - and as they were proboscideans, we may be reasonably certain that they were both highly intelligent and sociable. The Maltese wolves and bears might very well have preferred easier prey (the tooth marks on the fossil elephant bones are not conclusive evidence of predation; they could just as well represent scavenging).


Hunt, C.O. & Schembri, P.J. 1999. Quaternary environments and biogeography of the Maltese Islands. In: Mifsud, A. & Savona Ventura, C. (eds.) Facets of Maltese Prehistory, The Prehistoric Society of Malta, pp. 41-75.

Those predators only invaded Malta/Sicily at the end of the last interglacial and seem to have exterminated the large endemic species. Pitymys is just about the only endemic found together with them in the upper layers of Ghar Dalam.

By Tommy Tyrberg (not verified) on 17 Jan 2011 #permalink

@Jenny Islander- just a note, I think it's interesting to consider what hominid-mediated extinctions may have occurred in the early Pleistocene or even Pliocene, such as the aforementioned African giant tortoises or (later, after Acheulean technology, in early Pleistocene) some of Africa's diverse elephant species...
@Tommy Tyrberg-

Those predators only invaded Malta/Sicily at the end of the last interglacial and seem to have exterminated the large endemic species.

Really? how/why would they get there only at the END of the last interglacial? That's when sea level is higher...confused...
Also, when did humans arrive? I believe there are very early dates- at least predating last interglacial- for when Homo species, possibly H erectus, H neandertalensis, or H sapiens- necessarily crossed significant water bodies in Europe- possibly by boat.…
Although as Darren has previously posted, there are many examples of large mammals crossing a lot of water to get to islands without boats...such as Madagascar carnivores, Falklands fox, Flores Stegodons.


Those predators only invaded Malta/Sicily at the end of the last interglacial

Maybe so, but I wasn't making any claims about when they arrived or for how long they persisted; I just pointed out that they were present on Malta (if only temporarily).

seem to have exterminated the large endemic species

Perhaps they did (although I'm fairly sceptical of the idea of elephant-killing wolves). But it's worth noting that the predators themselves (or the Maltese bear at least) were apparently also reduced in size.

Pitymys is just about the only endemic found together with them in the upper layers of Ghar Dalam.

According to Hunt & Schembri, the Sicilian-Maltese endemic 'pygmy' red deer Cervus elaphus siciliae is also found in the same deposits.

Yes, but the red deer came in together with the predators and then differentiated subspecifically. The faunal dynamics of Malta/Sicily have been driven by the fact that it has mostly been an isolated island (or islands), that have occasionally been invaded by waves of immigrants when low sea levels and/or tectonics has permitted. Usually five quaternary "faunal complexes" are recognized:

Monte Pellegrino
Elephas falconeri
Elephas mnaidriensis

Grotta San Teodoro

Monte Pellegrino is early Pleistocene (Villafranchian). Elephas falconeri mostly early Middle Pleistocene. Both these are very unbalanced insular faunas that were presumably recruited by sweepstake dispersal. Elephas mnaidriensis which is probably late Middle Pleistocene to early Late Pleistocene is a much richer and more balanced assemblage with large predators and many mainland forms. It must have been recruited through some kind of filter dispersal (e. g. a very narrow/short-lived landbridge or a strait narrow enough to be swimmable). However it must have been followed by a new period of isolation (Last interglacial, MIS 5?) since there is a moderate degree of differentiation (i. a. Elephas mnaidriensis, Hippopotamus pentlandi). A few of the smaller taxa from the older faunas also survive e. g. Leithia, Maltamys.
San Teodoro is Late Pleistocene (Devensian in Great Britain). It is essentially identical to the mainland fauna and indicates that there was now definitely a land bridge (even horses and partridges occur, both are notoriously unable to cross water barriers). Castello finally is a depauperate late glacial version of San Teodoro. Homo only occur in the two latest faunas (on Sicily, not in Malta). Malta seems to have become separated from Sicily before the San Teodoro Faunal Complex arrived.

And actually there is very little evidence that any hominid before H. sapiens had any appreciable capacity for crossing water barriers. There were no definite pre-sapiens hominids on any Mediterranean islands (though early stone-tools have been claimed for Crete). Admittedly there is Homo floresiensis, but since they seem to have remained isolated on Flores for half a million years and evolved into a separate species they probably got there by sweepstake dispersal (e. g. swept out to sea on a raft of vegetation by a hurricane or tsunami).

By Tommy Tyrberg (not verified) on 18 Jan 2011 #permalink

In any case, endemic faunas of Mediterranean lived in isolation long after humans sailed to Australia.

BTW, remember the dwarf elephant on Egyptian sculpture? Is it possible that it was Elephas falconeri imported from one of Meds island?

An alternate history group I belong to had fun discussing that very idea.

What if one of the dwarf elephant species had been preserved, not only until the days of the Pharoahs, but right up to the present day? How might history have been affected if a donkey-sized elephant had been available for use as a beast of burden? Would it have given some people an advantage that they didn't have in the real past? Or would it just have been a curiosity traded among kings?

By Jenny Islander (not verified) on 18 Jan 2011 #permalink

David H.:
Maltese otter is Lutra euxena?
Shouldn't that be a Black Sea otter?
The classical name of the Black Sea was the Euxine, with an i in the second syllable. I don't know what "euxena" means, but with an "e" there i suspect it's a different word.

(Hmm... If the "xen" is from xenos (foreign), i suppose "euxena" could mean "really strange," but given my liomited Greek and the really strange way Paleontologists sometimes concoct names, I'm not going to speculate.)

By Allen Hazen (not verified) on 18 Jan 2011 #permalink

In any case, endemic faunas of Mediterranean lived in isolation long after humans sailed to Australia."

Really? Humans arrived in Australia 50 kya, Crete, perhaps >130 kya; other islands, who knows. I don't know it's nailed down yet; but just because humans crossed some water barriers, doesn't necessarily mean all water barriers would be crossed at or near that point.

remember the dwarf elephant on Egyptian sculpture?

Being too lazy to google that myself, how do they know it's a dwarf vs a baby or even just a misrepresented size? I mean, they also depicted people with the heads of alligators and birds...

Egyptian art is both highly stylized and very good at depicting the distinguishing features of whatever is being depicted--I could go on for a couple of paragraphs about why, but I won't. Anyway, the art in question clearly shows a very small elephant that has long tusks, so it isn't supposed to be a baby, and a back that slopes downward toward its hindquarters. If it had just been shown as smaller relative to a human than an adult elephant should be, it might have been chalked up to the way Egyptian artists adjusted the scale of the figures in a composition according to rules about who was most important and so forth. However, it was very small and it was not drawn like the classic elephant shown elsewhere in Egyptian art. These factors suggest that the artist had seen something that wasn't an African elephant.

By Jenny Islander (not verified) on 18 Jan 2011 #permalink

@Tommy Tyrberg

Do not underestimate the ability of navigation (or simply swimming) of primitive man. During the glacial peaks of the many islands in the Mediterranean are just a few kilometers from the mainland.

For example, Sardinia 500.000 years ago was probably reached by the first human migration, during glacial Tuscan archipelago became a peninsula, Corsica and Sardinia was connected.
There is no evidence of human bone (the oldest human remains is years ago), but some stone tools and another important clue.
In fact just 500.000 years ago was a fairly radical and sudden change wildlife.
"Nesogarale" wildlife (Nesogoral melons, Macaca maiori, Sus sondaari, Mammuthus lamarmorae)was replaced with "Tyrrenicola" association(Tyrrenicola Hensel Megaceros cazioti, Cynotherium sardous ) with a few survivors of the first (Prolagus sardus).
In between there are some caves, such as "Sa pedrosa-pantalinu" and "Sa Coa de Sa Multa," with stone tools dated as of the final phase of the Mindel glaciation.
It was not found any human artifact between 450,000 and 22,000 years ago.
In the south of Sardinia, around 22,000 years ago remains, including bone remains reliably attributed to Sapiens sapiens, with unusual features in the jaw and teeth.

Tyrrenicola fauna survived up to 12,000 years ago, when a new culture stone reached Sardinia from the mainland. Only Prolagus survived until the last remaining population, in Maddalena island, was hunted to extinction by sailors of Nelson's fleet at the beginning of the Trafalgar campaign.
If the man survived in Sardinia (and Corsica) between 450,000 and 22,000 years ago is a mystery.

Sorry if my English is difficult to understand.


The classical name of the Black Sea was the Euxine, with an i in the second syllable.

With ει, actually.

By David MarjanoviÄ (not verified) on 19 Jan 2011 #permalink

Correct me:
a) there are other sites with traces of human occupation in Sardinia in the middle of the "vacuum period". One is the Ziu Santoru cave, where wos found a firebox and bone remains Megaceros cazioti (to eat the broken bone) which may date back to 150,000 to 120,000 years ago.
Another is between Riu Codrovulos and Pantallinu and where there are clearly material paleolithic perhaps 300,000 or 150,000 years ago.
b) The survival of Prolagus to finish in the table Nelson was a hoax by an amateur cryptozoologist, though perhaps on the island of Tavolara still existed in the middle of '700.

[from Darren: sorry, delayed by spam filter]

I'm currently introducing a bunch of Korean biologists to english idioms. Yesterday, we were having difficulty with "damning with faint praise." The room finally lit up with comprehension when I refered them to this comment on my book (which most of them had read)in your blog:

" a nonscientific tract, but does compile a number of different natural history facts (and is fairly readable and not too crackpottish).
Also, the somewhat compelling case of the extinction of African giant tortoises- if you read this you must start halfway down page:"

I've been aware of, and a reader and fan of, your blog for a while. You're not so crackpottish yourself. In fact you're downright a downright rigorous, sensible and fascinating scientist.

Anyway, this evening I'm going try and explain to my group how one "gives the devil his (or her) due."

" give a person who one regards with dislike or condescension proper credit (for something). (She's very messy in the kitchen, but I have to give the devil her due. She bakes a terrific cherry pie.)" (

Perhaps you could supply me with another apposite example of this particular idiom? In reference, say, to my megafauna site?

You may want to look, in this regard, at Paul S Martin's Foreword to my online book:

And thanks again for the excellent and informative material which you're offering free online.

Baz - thanks for your comment. Just to clear things up, it wasn't me who commented on your site. Thanks for the compliments, much appreciated.

Ah, I see why you thought I was the author of comment 24 (they signed themselves as "me"). It's another commenter, using a pseudonym (see also comment 29).