The New World Leaf-Nosed Bat Radiation

I’ve said a few times here at TetZoo that bats have never really been given adequate coverage. This isn’t because I’m not interested in them: on the contrary, I think about bats more than I think about most other groups of mammals, and I see them and watch them more often than I do most other mammal groups. For a group that includes about 18% of extant mammalian species (using 2019 figures*), I can’t pretend to have ever given bats fair coverage. Having said all that, bats have actually been covered at TetZoo a fair bit: there was an entire 20-part series on vesper bats (properly Vespertilionidae) at ver 3, and I also published several ver 2 articles on the history and evolution of vampire bats, and on much else besides. The fact that all of these articles have been rendered worthless via the removal of their images is mightily dispiriting though, and essentially means that I need to start from scratch.

* c 6495 mammal species, c 1200 bat species.

Caption: TetZoo Towers bat library. The several boxfiles of reprints and photocopied articles are not shown. Image: Darren Naish.

Here, I want to talk about a group I don’t think I’ve ever covered at TetZoo before, namely the phyllostomids, or New World leaf-nosed bats, American leaf-nosed bats or spear-nosed bats. This is a large, American group that contains around 200 living species, making it the third largest bat family (vesper bats are the biggest group, followed by fruit bats). The group has sometimes been called Phyllostomatidae – the vernacular version of which is phyllostomatid – but this is less popular than Phyllostomidae. I have no idea which is really correct here and opt to merely follow majority usage on these sorts of things (insert quote from Gene Gaffney**). It’s not strictly true that I’ve never covered phyllostomids before, since vampires – once upon a time given their own eponymous family (Desmodontidae) – are now universally agreed to be nested within Phyllostomidae, and I have at least written about them.

Caption: Chrotopterus, a big spear-nosed bat. Notice how this bat has relatively broad, low-aspect wings and a large, deep uropatagium (the membrane between the legs). Contrast this with some of the images below. Image: George Henry Ford, public domain (original here).

Phyllostomids occur from Argentina in the south to the southern USA (Nevada being their most northerly occurrence) in the north, and they’re highly diverse ecologically and behaviourally. They include insectivores, frugivores, nectarivores, palynivores (that’s pollen-eaters), omnivores, animalivores and (of course) obligate sanguivores. Numerous different taxonomic subdivisions have been named. We don’t need to worry about any of this in detail but, in simplified terms, Macrotinae (big-eared bats), Micronycterinae (little big-eared bats) and Desmodontinae (vampires) are outside a much larger clade that includes Vampyrinae (false vampires and kin) and Phyllostominae (spear-nosed bats and kin) as well as the nectarivorous and frugivorous Glossophaginae (long-tongued and long-nosed bats) and Stenodermatinae (American fruit bats, fig-eating bats and kin) (Baker et al. 1989, 2003, 2012; but see Wetterer et al. 2000). Vampyrinae is a clade within Phyllostominae according to some studies, in which case it gets down-graded to Vampyrini (Baker et al. 2003). All of this is depicted in a cladogram below.

Caption: some phyllostomid portraits. At left: Big-eared woolly bat or Peters’s false vampire Chrotopterus auritus. At right: Hairy big-eyed bat Chiroderma villosum. Images: both Guilherme Garbino, wikipedia, CC BY-SA 4.0 (originals here and here).

Phyllostomids are mostly brownish bats with simple, narrow ears. A nose-leaf – typically simple and spear-shaped – is common but not present in all species, a tragus is always present, and many (but not all) of the species that lack nose-leaves have chin-leaves (or a series of chin ‘warts’) instead. Facial stripes are common, dark dorsal stripes are present in a few species, and such things as white patches at the wing tips and yellow rims to the ears and nose-leaves are present in some (Hill & Smith 1984). The tail is variously long, or short, and even absent altogether in some taxa, and similar variation is present in the uropatagium, or tail membrane.

Caption: the tail and uropatagia (the membranes joining the inner sides of the legs to the tail) are reduced, and sometimes highly reduced, in some phyllostomids. Here, we see this reduced condition in (at left) the Toltect fruit-eating bat Dermanura tolteca and (at right) in a Little yellow-shouldered bat Sturnira lilium. Images: M.H. de Saussure, 1860, in public domain (original here); Tobusaru, wikipedia CC BY 3.0 (original here).

Skeletally, phyllostomids are robust, have a distinctive humerus where the distal end is angled relative to the shaft, and have a prominent secondary articulation between the large bony lump (properly termed the greater tuberosity) at the proximal end of the humerus and the scapula (Czaplewski et al. 2007). That’s right: a number of bat groups have an accessory peg-in-socket articulation involving the humerus and the body of the scapula. This means that the humerus and scapula are locked together during the upper part of the wing stroke (Hill & Smith 1984).

Caption: the most prominent exception to the ‘phyllostomids are mostly brown’ generalisation is the Honduran white bat Ectophylla alba, sometimes likened to a fuzzy ping-pong ball and well known for its habit of constructing tents by biting through leaf ribs such that the two sides of the leaf droop on either side of the central axis. Note the yellow ears and nose leaf! The individual at left is releasing a bit of urine. Images: Geoff Gallice, wikipedia, CC BY 2.0 (original here); Leyo, wikipedia, CC BY-SA 2.5. (original here)

Some phyllostomids are really exceptional as goes their anatomical and behavioural novelty. Perhaps the most remarkable are the long-tongued glossophagine flower bats, some of which have extraordinary tubular snouts, remarkably long tongues tipped with papillae, and a highly reduced dentition. The most extreme example of this sort of thing is the Banana bat, Trumpet-nosed bat or Colima long-nosed bat Musonycteris harrisoni of Mexico, an ‘extreme’ mammal as goes snout length. It’s fairly typical for people who aren’t that familiar with bat diversity to confuse glossophagines with the Old World flower-feeding megabats grouped together in Macroglossinae. There’s obviously a degree of evolutionary convergence here, though it hasn’t been that well explored in the literature, to my knowledge. Various glossophagines have symbiotic relationships with sympatric plants. Incidentally, Pallas’s long-tongued bat Glossophaga soricina is able to see UV light (Winter et al. 2003).

Caption: some distantly related (but broadly similar) members of the phyllostomid clade Glossophaginae. At left: a long-tongued champion (though not necessarily the longest-tongued of phyllostomids), Pallas’s long-tongued bat Glossophaga soricina. At right: Underwood’s long-tongued bat Hylonycteris underwoodii. Images: Betty Wills, wikipedia CC BY-SA 4.0 (original here); Karin Schneeberger/Felineora, wikipedia CC BY-SA 3.0 (original here).

Entirely different specialisations are seen in the short-faced, frugivorous phyllostomids included within Stenodermatinae. These have flattened, broad teeth, typically have white facial stripes (an aposematic warning of their powerful bites?), and are sometimes handsome or even cute, big-eyed bats. One of the strangest of bats – the Wrinkle-faced or Lattice-winged bat Centurio senex – belongs to this group. The naked, wrinkled faces of males are mostly concealed by massive skin flaps when the bat is roosting or sleeping. There are also neck glands that seem to secrete scent, and obvious transverse bands on the wing membranes.

Caption: resting Wrinkle-faced bats Centurio senex partially conceal their faces beneath thick skin folds. Translucent patches on the lower of these skin folds seem to allow these bats to detect light-level changes even when their faces are covered. Image: Jplevraud, wikipedia CC BY-SA 3.0 (original here).

My favourite phyllostomids are very different from tubular-snouted flower-feeders and short-face fruit-eaters: they are the robust, more generalised species traditionally lumped together in Phyllostominae (though the name Vampyrinae has also been used for some of them). These are mostly omnivores that eat insects, fruit and small vertebrates, and some are specialised predator bats that variously catch and eat amphibians, mammals (including other bats) and birds. They include Peters’s woolly false vampire Chrotopterus auritus, the Frog-eating bat Trachops cirrhosus – famous for eating frogs and selecting them on the basis of their calls – and the spectacular Linnaeus’s false vampire Vampyrum spectrum, a predatory giant that can, in cases, have a wingspan of over 1 meter.

Caption: Vampyrum, the False vampire or Spectral bat (see comments for a hot take on the term ‘false vampire’), has to be considered one of the most awesome of all bats. It’s convergently similar to the distantly related megadermatid bats of Africa, Asia and Australasia, also (confusingly) often called false vampires. Image: Marco Tschapka, wikipedia, CC BY-SA 3.0 (original here).

Caption: the very impressive skull of Vampyrum. It is robust, with big, strong teeth, especially prominent upper canines (which have an additional internal cusp) and a prominent sagittal crest. The skull can be 5.1 cm long in total (which is big for a bat). Image: Naturalis Biodiversity Center, wikipedia, public domain (original here).

When this variation in feeding ecology is mapped onto a phylogeny, it would appear that the earliest phyllostomids were insectivorous, that omnivory, nectarivory (or nectivory, take your pick) and palynivory evolved from among these insectivores, and that frugivores evolved from among nectarivores and palynivores (Baker et al. 2012). The highly specialised vampires appear – according to phylogenetic data – to have evolved directly from insectivores (which is a surprise in view of some models proposed to explain vampire evolution) and at least some members of the main frugivorous clade appear to have reverted to insectivory (Baker et al. 2012; but see Wetterer et al. 2000). Of the various evolutionary events that must have occurred here, it’s the transition to obligate frugivory that seems to have been the most successful, since the frugivorous clade is the largest (as in, most species-rich) within Phyllostomidae, containing about 70 species in 20 genera.

Caption: a few more vertebrate-eating phyllostomids. At left: California leaf-nosed bat Macrotus californicus, the most northerly occurring phyllostomid. At right: Fringe-lipped bat Trachops cirrhosus, a widespread species of Central and South America that eats seeds, fruits, arthropods and lizards in addition to frogs. Images: National Wildlife Service, wikipedia, public domain (original here); Karin Schneeberger/Felineora, wikipedia CC BY 3.0 (original here).

This is also the radiation that’s seemingly resulted in the greatest, most rapidly evolved amount of morphological variation, since everything here seems to have happened within the last 10 million years and has given rise to taxa that are among the most divergent and specialised of phyllostomids. Also of interest here is that some lineages within this frugivorous clade appear to have evolved in the Antilles before invading the mainland (Dávalos 2007), a case of ‘upstream colonisation’ that contradicts traditional scenarios whereby continental animals give rise (via ‘downstream colonisation’) to island-dwelling forms.

Caption: substantially simplified phyllostomid cladogram, based mostly on Baker et al. (2003), and using their nomenclature (though they regarded false vampires - as Vampyrini - as nested within Phyllostominae). Images (top to bottom): Macrotus = National Wildlife Service, wikipedia, public domain (original here); Desmodus = Uwe Schmidt, wikipedia, CC BY-SA 4.0 (original here); Vampyrum = Marco Tschapka, wikipedia, CC BY-SA 3.0 (original here); Phyllostomus = Karin Schneeberger/Felineora, wikipedia, CC BY 3.0 (original here); Platalina = Juan A. Malo de Molina, wikipedia, CC BY-SA 3.0 (original here); Sturnira = Burtonlim, wikipedia, CC BY-SA 3.0 (original here).

Where in the bat tree? What sort of bats are phyllostomids, and what do we know about their evolutionary history? On the basis of anatomical characters, bat experts have generally thought that phyllostomids are close allies of naked-backed, moustached or ghost-faced bats (Mormoopidae) and bulldog bats and kin (Noctilionidae), the whole lot being grouped together in a clade termed either Phyllostomatoidea or Noctilionoidea (and it’s the last of those terms that should be preferred, so I understand). In turn, this group was thought – again, on the basis of anatomical characters – to be closely related both to vesper bats and their kin (Vespertilionoidea), and to a clade that includes both sheath-tailed bats and kin (Emballonuroidea) and horseshoe bats and kin (Rhinolophoidea) (Smith 1976).

Caption: prior to recent (post-2000-ish) molecular studies, noctilionoids were thought to be close kin of rhinolophoids as well as emballonuroids and vespertilionoids. Rhinolophoids are now known to belong elsewhere. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project, progress on which can be seen here. Image: Darren Naish.

Molecular studies, mostly published since 2000, have substantially revised our view of the bat family tree, however, and it’s now clear that rhinolophoids are not close to the other groups listed here at all (they are, instead, close relatives of megabats). Noctilionoids are still close kin of vespertilionoids, however. It also now seems that Mystacinidae and Myzopodidae are part of Noctilionoidea (Jones et al. 2002, 2005, Teeling et al. 2005, 2012). I’ll be talking more about ideas on bat phylogeny in a future article.

Caption: simplified cladogram depicting the affinities of several of the bat groups shown - via morphological and molecular studies - to belong together within Noctilionoidea. The illustrations here are among the many, many bat drawings I’ve done for my in-prep textbook project, progress on which can be seen here. The Vampyrum representing Phyllostomidae, incidentally, is a placeholder which needs replacing (the existing illustration was copied directly from the work of another artist). Image: Darren Naish.

What does the fossil record say about phyllostomid history? The pre-Pleistocene phyllostomid record is not great but it’s still at least good enough to show that the extinct phyllostomids of the Miocene – most notably those from La Venta in Colombia – were superficially much like living ones, and that the extinct species concerned were doing the sorts of things that phyllostomids do today. The group had almost certainly, therefore, undergone its main flowering and diversification by around 20 million year ago. The Pleistocene phyllostomid record, in contrast, is good and numerous extant taxa are known from sediments of this age. 

Caption: beautiful illustration of Salvin’s big-eyed bat Chiroderma salvini, a stenodermatine phyllostomid that has a wide range across South and Central America. The facial stripes are not normally this pronounced in life, though it should be noted that populations are variable as goes stripe thickness. Image: Joseph Smit, in public domain (original here).

And that’s where we’ll end things for now. I’d like to say a lot more about these bats, so we’ll be returning to them in time. And in fact I need to say a lot more about bats in general, so stay tuned for that too.

If you enjoyed this article and want to see me do more, more often, please consider supporting me at patreon. The more funding I receive, the more time I’m able to devote to producing material for TetZoo and the more productive I can be on those long-overdue book projects. Thanks!

For previous TetZoo articles on bats (concentrating here on articles that haven’t been stripped of images, as is the case for all ver 2 articles and the vast majority of ver 3 articles)…

• Chewed bones and bird-eating microbats, June 2006

• Greater noctules: specialist predators of migrating passerines, June 2006

• We flightless primates (on the Pettigrew ‘megabats are close kin of primates’ model), August 2006

• Books of the TetZooniverse: of Palaeoart, Bats, Primates and Crocodylians, August 2015

• Fossil Bat Stories, Part 1, February 2018

• Fossil Bat Stories, Part 2: What Are Noctilionids? What Are Noctilionoids?, February 2018

• Fossil Bat Stories, Part 3: Bulldog Bats, February 2018

Refs - -

Baker, R. J., Bininda-Emonds, O. R. P., Mantilla-Meluk, H., Porter, C. A. & Van Den Bussche, R. A. 2012. Molecular time scale of diversification of feeding strategy and morphology in New World leaf-nosed bats (Phyllostomidae): a phylogenetic perspective. In Gunnell, G. & Simmons, N. (eds). Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Cambridge, pp. 385-409.

Baker, R. J., Hoofer, S. R., Porter, C. A. & Van Den Bussche, R. A. 2003. Diversification among New World leaf-nosed bats: An evolutionary hypothesis and classification inferred from digenomic congruence of DNA sequence. Occasional Papers, Museum of Texas Tech University 230, 1-32.

Baker, R. J., Hood, C. S. & Honeycutt, R. L. 1989. Phylogenetic relationships and classification of the higher categories of the New World bat family Phyllostomidae. Systematic Zoology 38, 228-238.

Czaplewski, N. J. 1997. Chiroptera. In Kay, R. F., Madden, R. H., Cifelli, R. L. & Flynn, J. J. (eds) Vertebrate Paleontology in the Neotropics: The Miocene Fauna of La Venta, Colombia. Smithsonian Institution Press (Washington and London), pp. 410-431.

Dávalos, L. M. 2007. Short-faced bats (Phyllostomidae: Stenodermatinae): a Caribbean radiation of strict frugivores. Journal of Biogeography 34, 364-375.

Hill, J. E. & Smith, J. D. 1984. Bats: A Natural History. British Museum (Natural History), London.

Jones, K. E., Bininda-Emonds, O. R. P. & Gittleman, J. L. 2005. Bats, clocks, and rocks: diversification patterns in Chiroptera. Evolution 59, 2243-2255.

Jones, K. E., Purvis, A., MacLarnon, A., Bininda-Emonds, O. R. P. & Simmons, N. B. 2002. A phylogenetic supertree of the bats (Mammalia: Chiroptera). Biological Reviews 77, 223-259.

Smith, J. D. 1976. Chiropteran Evolution. Texas Tech University, Lubbock.

Teeling, E. C., Dool, S. & Springer, M. S. 2012. Phylogenies, fossils and functional genes: the evolution of echolocation in bats. In Gunnell, G. & Simmons, N. (eds). Evolutionary History of Bats: Fossils, Molecules and Morphology. Cambridge University Press, Cambridge, pp. 1-22.

Teeling, E. C., Springer, M. S., Madsen, O., Bates, P., O’Brien, P. & Murphy, W. J. 2005. A molecular phylogeny for bats illuminates biogeography and the fossil record. Science 307, 580-584.

Wetterer, A. L., Rockman, M. V. & Simmons, N. B. 2000. Phylogeny of phyllostomid bats (Mammalia: Chiroptera) data from diverse morphological systems, sex chromosomes, and restriction sites. Bulletin of the American Museum of Natural History 248, 1-200.

Winter, Y., López, J. & von Helversen, O. 2003. Ultraviolet vision in a bat. Nature 425, 612-614.

** In a technical article on fossil side-necked turtles, Gaffney said of a very similar nomenclatural disagreement: “it’s true, I don’t give a rat’s ass which is used”.

The Life Appearance of the Giant Deer Megaloceros

Eurasian Pleistocene megafauna are among the most familiar and oft-depicted of prehistoric animals. And among these grand, charismatic and imposing animals is the giant deer Megaloceros giganteus, an Ice Age giant that occurred from Ireland and Iberia in the west to southern Siberia in the east. It persisted beyond the end of the Pleistocene, surviving into the Early Holocene on the Isle of Man (Gonzalez et al. 2000) and western Siberia (Stuart et al. 2004)*. It is often erroneously termed the Irish elk, though it certainly wasn’t restricted to Ireland, nor should it really be termed an ‘elk’ (ugh… we’ll avoid that whole hornet’s nest for the time being). It’s been termed the Shelk by others [UPDATE: but see comments!!]. It could be 1.8 m tall at the shoulder and weigh somewhere around 600 kg, the antlers spanning 3.5 m in cases and weighing 35-45 kg (Geist 1999).

Caption: a very conventional, traditional image of Megaloceros giganteus: it's depicted looking like a giant red deer, basically. Males and females are not that different in size, but males are often shown as maned. Most interest in this deer has, of course, concerned the spectacularly antlered males. This image is from Hutchinson's Extinct Monsters (published several times over the 1890s). Image: Hutchinson (1892).

* In a previous edit of this article, I said that M. giganteus also survived into the Holocene in central Europe, as demonstrated by Immel et al. (2015). I missed the fact that this research concerns specimens dated to the Upper Pleistocene, not the Holocene. Furthermore, I’ve also been told that the Isle of Man data proved incorrectly dated. Am chasing confirmation on this.

While big, M. giganteus was not the biggest deer ever, since it seems that the extinct, moose-like Cervalces latifrons was even bigger. I promise to talk more about that species when I get round to discussing moose and kin at length. And while the antlers of M. giganteus were obviously very big, they weren’t especially big relative to its body size: proportionally, they were about similar in size to those of large Fallow deer Dama dama, and well exceeded in proportional size by the antlers of reindeer and caribou.

Caption: a fine Megaloceros skull on show at London's Grant Museum. I seem to recall hearing or reading - possibly in one of Stephen J. Gould's papers - that this is one of the largest specimens in existence. Image: Darren Naish.

I should add that M. giganteus was not the only Megaloceros species. Several others are known, differing in how palmate or slender and branching their antlers were, and not all were as large as M. giganteus (some were island-dwelling dwarves). There are other genera within this deer lineage (Megacerini) as well. Also of relevance to our discussion here is the position of these deer within the cervid family tree. Some experts have argued that megacerines are close to deer like the Red deer Cervus elaphus (Kuehn et al. 2005), while others point to genetic and morphological data indicating a close relationship with the Fallow deer Dama dama (Lister et al. 2005, Hughes et al. 2006, Immel et al. 2015, Mennecart et al. 2017). I have a definite preference for the latter idea, and right now it's a far better supported relationship than the alternative.

Caption: male M. giganteus skulls in the collections of the National Museum of Ireland, Dublin, examined in 2008. Yes, there is indeed a preponderance of males. Image: Darren Naish.

Like most European people who’ve been lucky enough to visit museums and other such institutions, I’ve seen Megaloceros specimens on a great many occasions – there are a many of them on display. I’ve also seen and handled a reasonable number of the Irish bog specimens during time spent in Dublin. There does appear to be a preponderance of big, mature males. Maybe this reflects collecting bias (in that people were more inclined to extract the skulls and skeletons of big, prominently antlered males), but it also seems to be a valid biological signal: it has been argued that the calcium-hungry males were likely attracted to calcium-rich plants like willow at the edges of lakes and ponds, and were thus more prone to drowning, miring or falling through ice in such places than females (Geist 1999). Oh, we also know that male mammals across many species are more inclined to take stupid risks, be reckless, and even display deliberate bravado more than their female counterparts.

Here we come to the main reason for this article: what, exactly, did M. giganteus look like when alive? I’ve surely mentioned this topic on several occasions over the years here; I’m pretty sure I threatened to write about it after producing similar articles on the life appearance of the Woolly rhino and Ice Age horses. M. giganteus has been illustrated a great many times in works on prehistoric life, and the vast majority of reconstructions show it a near-monotone dark brown or reddish-brown. It’s very often depicted with a shaggy neck mane. In short, it’s usually made to look like a big, shaggy Red deer, and the tradition whereby this is done – it extends back to Zdenek Burian, Charles Knight and other founding palaeoartists – seems to me to be another of those palaeoart memes I’ve written about before. I’ve taken to calling this one the ‘Monarch of the Glen’ meme (see my palaeoart meme talk here). I will add here that we're generally talking about males of the species (since people mostly want to see depictions of specimens with those awesome antlers), though virtually all that I say below applies to females too.

Alas, this view of M. giganteus is almost certainly very wrong. Why do I say this?

Caption: note the many obvious external features of this male Fallow deer: a throat bulge corresponding with the larynx - an 'Adam's apple' - is obvious, and this is a boldly marked deer overall, with prominent spots (including some that have coalesced into stripes), a white rump patch, and pale ventral regions. If megacerines are close kin of Dama deer, we might predict a similar ancestral condition for Megaloceros and its relatives. Image: Dave Hone.

Firstly, if we look at the colours and patterns present across cervine deer as a whole, we see quite a bit of variation and no strong and obvious reason why a ‘Red deer look’ should be favoured. Secondly, we have that data indicating that M. giganteus is phylogenetically closer to Dama deer than to Cervus, in which case we would predict that it descended from ancestors with prominent spotting, pale flank stripes, and dark markings on the tail, all features typical of modern Dama populations. If the ‘Dama hypothesis’ is correct, there is again no reason to favour a ‘Red deer look’ for M. giganteus. Thirdly, body size, limb proportions, antler size and habitat choice all indicate that M. giganteus was an open-country (Clutton-Brock et al. 1980), cursorial specialist, and in fact the most cursorial of all deer (Geist 1999). Cursorial, open-country artiodactyls are often pale, with large white areas across the rump, legs and belly (examples include addax, some Arctic caribou and some argali). Again, no reason here to suspect that ‘Red deer look’.

And... fourthly, we have direct eyewitness data on the life appearance of this animal. Members of our own species saw it in life and drew it, seemingly to a very high degree of accuracy. What did they show?

Caption: the famous panel at Cougnac, southwest France, showing M. giganteus males and females. This part of the cave is also interesting in depicting a short-horned bovid (at upper right) sometimes interpreted as a tahr. There are also ibex here too. I'm uncertain of the exact origin of the photo shown here: I took it from Fabio Manucci's blog Agathaumus. Numerous additional photos of the same cave can be seen at Don's Maps.

Virtually all cave art depicting M. giganteus shows a rounded, tall shoulder hump that’s sometimes shown as if it had a crest of raised hairs. Guthrie (2005) termed this a ‘hackle tuft’. There’s no obvious indication from the skeleton that a hump like this was present (indeed, fatty humps in mammals very often do not have an underlying skeletal correlate), so this is a neat thing that we wouldn’t know from skeletons alone. A protruding lump on the throat that seems to correspond to the larynx is also shown in images at Lascaux, Roucadour and elsewhere (Guthrie 2005). This feature is very reminiscent of Fallow deer.

Caption: cave art depicting M. giganteus is not all that numerous (most ancient depictions of deer are of reindeer or red deer), but what does exist shows several details worthy of note, here emphasised in illustrations produced by R. Dale Guthrie. The shoulder hump is a consistent feature. Image: Guthrie (2005).

Some of the art provides information on pigmentation. A collar-like band is depicted encircling the neck in images from Chauvet and Cougnac, the shoulder hump is shown as being dark in images from Cougnac and elsewhere (Lister 1994), and some of the Chauvet and Roucadour images show a dark diagonal line that extends across the side of the body from the shoulder to the edge of the groin, and sometimes across the leg as far as the hock (ankle). An especially detailed image at Cougnac, partially illustrated on a stalactite, shows what looks like a dark vertical stripe descending from the shoulder hump and forming a division between the deep neck and the rest of the body. The same image also shows dark near-vertical markings around what might be a pale rump patch (Guthrie 2005).

Caption: other people have taken the same evidence I've discussed here and produced very similar reconstructions. This piece - which I hadn't seen until after producing my own illustrations (on which, see below) - is by Pavel Riha. Image: Pavel Riha, CC BY-SA 3.0.

If these details have been interpreted correctly, M. giganteus was boldly marked, with obvious dark striping across its neck, shoulders and torso, and on its rump too. R. Dale Guthrie proposed that the vertical shoulder stripe formed a boundary between a near-white neck and head region and the rest of the body, with the latter being pale just posterior to the stripe but darker across the legs, rump and flank (Guthrie 2005). I’m not absolutely convinced by the evidence from cave art for a near-white neck and head or for a white rump patch but these things are consistent with what I said above about the open-country lifestyle and cursoriality of this deer. Geist (1999) was a fan of this idea, and his reconstruction of M. giganteus – shown here – is meant to show the animal as being quite pale apart from its obvious striping and other dark markings.

Caption: M. giganteus as reconstructed by Valerius Geist, and shown to scale with the extant Dama dama. Geist was (and presumably is) a strong advocate of the idea that megacerines (yes: megacerines, not 'megalocerines') are part of the same lineage as Dama. Image: Geist (1999).

Guthrie produced a very striking illustration depicting all of these details, but his drawing, as reproduced in his book (Guthrie 2005), is less than 4 cm long. Here it is (below), but note that I’ve produced a larger illustration here (scroll down) that shows the same details.

Caption: at left, the best of the M. giganteus images from Cougnac in France, as re-drawn by Guthrie (2005). At right, Guthrie's reconstruction of the animal's life appearance. Image: Guthrie (2005).

And that just about brings us to a close. Over the years, I’ve been perpetually dismayed by the fact that most people illustrating this animal aren’t aware of the information I’ve discussed here – I mean, we have direct eyewitness data that should be pretty much the first thing we take account of when reconstructing this animal. Alas, the usual problem here is that the people who provide advice on reconstructions of fossil animals to artists are virtually never that interested in or knowledgeable about the life appearance of the animals concerned (sorry, palaeontologists). That’s an unfair generalisation though, and there are of course exceptions. Indeed, I should note that accurate, informed reconstructions of M. giganteus have appeared here and there over the years: the Megaloceros depicted in the Impossible Pictures TV series Walking With Beasts, for example, includes most of the features I’ve discussed here and obviously benefitted from the input of an informed consultant.

Anyway, my hope for the article you’re reading now is that it will inspire the current generation of palaeoartists to start illustrating Megaloceros in a way that’s more in accord with the data from prehistoric art, all of which has been out there in the literature for years now (Lister 1994, Guthrie 2005).

I have further articles of this sort in mind and hope to get them published here eventually. On that note, here’s your reminder that I rely on your kind support at patreon, and that the more such support I receive, the more time and effort I can devote to Tet Zoo, and to my various book projects.

For previous Tet Zoo articles on Pleistocene megafauna, see...

• Tet Zoo picture of the day # 3 (Elasmotherium), May 2007

• The remarkable life appearance of the Woolly rhino, November 2013

• Spots, Stripes and Spreading Hooves in the Horses of the Ice Age, February 2015

And for articles on deer, see...

• Oh deer oh deer oh deer, October 2006

• Deer oh deer, this joke gets worse every time I use it, November 2007

• Britain: wildlife theme-park, December 2007

• At the 56th SVPCA - hello Dublin! (Megaloceros), September 2008

• The plasticity of deer, October 2008

• Pouches, pockets and sacs in the heads, necks and chests of mammals, part IV: reindeer and a whole slew of others, October 2010

• The seemingly endless weirdosity of the Milu, October 2011

• Eld's deer: endangered, persisting in fragmented populations, and morphologically weird... but it wasn't always so, April 2012

• South America's Many Remarkable Deer, November 2014

• Confrontational behaviour and bipedality in deer, December 2014

Refs - -

Clutton-Brock, T. H., Albon, S. D. & Harvey, P. H. 1980. Antlers, body size and breeding group size in the Cervidae. Nature 285, 565-567.

Geist, V. 1999. Deer of the World. Swan Hill Press, Shrewsbury.

Gonzalez, S., Kitchener, A. C. & Lister, A. M. 2000. Survival of the Irish elk into the Holocene. Nature 405, 753-754.

Guthrie, R. D. 2005. The Nature of Paleolithic Art. The University of Chicago Press, Chicago and London.

Hughes, S., Hayden, Th. J., Douady, Ch. J., Tougard, Ch., Germonpré, M., Stuart, A., Lbova, L., Garden, R. F., Hänni, C. & Say, L. 2006. Molecular phylogeny of the extinct giant deer, Megaloceros giganteus. Molecular Phylogeny and Evolution 40, 285-291.

Hutchinson, H. N. 1892. Extinct Monsters, 2nd edition. London: Chapman & Hall.

Immel, A., Drucker, D. G., Bonazzi, M., Jahnke, T. K., Münzel, S. C., Schuenemann, V. J., Herbig, A., Kind, C.-J. & Krause, J. 2015. Mitochondrial genomes of giant deers suggest their late survival in Central Europe. Scientific Reports 5: 10853.

Kuehn, R., Ludt, C. J., Schroeder, W. & Rottmann, O. 2005. Molecular phylogeny of Megaloceros giganteus - the Giant deer or just a giant red deer? Zoological Science 22, 1031-1044.

Lister, A. M. 1994. The evolution of the giant deer, Megaloceros giganteus (Blumenbach). Zoological Journal of the Linnean Society 112, 65-100.

Lister, A. M., Edwards, C. J., Nock, D. A. W., Bunce, M., van Pijlen, I. A., Bradley, D. G., Thomas, M. G. & Barnes, I. 2005. The phylogenetic position of the ‘giant deer’ Megaloceros giganteus. Nature 438, 850-853.

Mennecart, B., deMiguel, D., Bibi, F., Rössner, G. E., Métais, G., Neenan, J. M., Wang, S., Schulz, G., Müller, B. & Costeur, L. 2017. Bony labyrinth morphology clarifies the origin and evolution of deer. Scientific Reports 7: 13176.

Stuart, A. J., Kosintsev, P. A., Higham, T. F. G. & Lister, A. M. 2004. Pleistocene to Holocene extinction dynamics in giant deer and woolly mammoth. Nature 431, 684-689.