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Naish_profile_70_px.jpg With six years of phd work on theropod dinosaurs behind him, Darren Naish mostly spends long hours in the library, hunched over his laptop. But he gets out sometimes, and picks up litter and pursues exotic lizards across the British countryside, aiming all the while to publish his technical work on obscure Cretaceous dinosaurs. He also messes around with pterosaurs, swimming giraffes, British big cats and stuff like that. He has given up on the stupid idea of being a dedicated academic and ekes out a living as a technical consultant, editor and author. He can be contacted intermittently at eotyrannus (at) gmail dot com. For more biographical info go here.

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« One-eyed indri | Main | Giant killer pigs from hell »

Sleep behaviour and sleep postures

Category: mammalogyornithology
Posted on: September 15, 2008 5:39 AM, by Darren Naish

Sleeping_asian_elephant_wikipedia.jpg

For a long time now I have been, shall we say, gently encouraged by two of my friends to write about a subject that is both familiar, and yet also strangely alien and poorly understood. Sleep behaviour. We still know comparatively little about this subject: not only about the big stuff like its function, but even about its distribution within animals. I am not, by the way, about to tackle the big questions about sleep, nor am I going to discuss the different types of sleep (e.g., REM vs NREM sleep) and on how they differ from creature to creature. Instead I'm interested in the more superficial stuff, like how and where animals sleep, and on the postures they adopt.

What we do know about sleep is scattered widely in the literature and it isn't easy to obtain any sort of review of the subject. There is at least one book devoted to it though: Maurice Burton's Sleep and Hibernation in the Animal World (Odhams Books, 1969), one of twelve books written for the Animal World in Colour series. Burton is well known for amassing large amounts of anecdotal data on the natural world (he also wrote books covering cryptozoology and the emotional lives of animals), and while the books in the Animal World in Colour series include a lot of often bizarre and fascinating information, they are completely unreferenced and at least some of the content looks apocryphal and difficult to be confident about. Despite these problems I have borrowed heavily from his book on sleep here.

sleeping_cow_in_India.jpg

Needless to say, I'm only going to be looking here at sleep in tetrapods, though where relevant I will mention other animals. The first thing to mention is that some animals get by without virtually any sleep, or in fact may even go without sleep entirely. Among vertebrates, all identified non-sleeping species are fish (Kavanau 1998). The species concerned have little in common (they include blind cave-dwellers, various sharks including some triakids, lamnids, hammerheads and sevengills, and mackerels and scombrids), but all have lifestyles where 'visual input is greatly reduced or absent during lengthy periods', where 'schooling greatly reduces needs for sensory information, particularly visual', or where the animals lead a 'comparatively routine existence in essentially featureless, open waters' (Kavanau 1998, p. 269) [sleeping cow from here].

Perhaps because no tetrapod lives in this way, it seems that all indulge in at least some sleep activity. But some sleep very little, with one of the best examples being seabirds: some species are on the wing for weeks at a time and appear to be constantly active. It has been suggested that they might sleep on the wing, but it's considered equally plausible that they simply delay their sleep. And having mentioned 'sleep on the wing', while it's 'well known' that swifts sleep in flight, the evidence for this (which essentially consists of observations of swifts seen decreasing in altitude during the early hours of the morning) is poor and we still require detailed studies (Rattenborg 2006).

donkey_not_dead_just_lying_in_the_mud.jpg

Some ruminants also seem to engage in very little sleep. This might be because they need to remain constantly alert for predators, but it might also be because their biology requires a virtually continuous cycle of eating and ruminating. Burton (1969) wrote of observations made on goats which suggest that, during a rest period that lasts about 8 hours, they never really relax fully, and it is even stated that some ruminants (like cattle) sleep with their eyes open (this must be true: before indulging in the nefarious activity known as cow-tipping you're supposed to shine a torch into the cow's eyes to see whether it reacts or not. No reaction = asleep). Ruminants do sleep with their eyes closed at least sometimes however, as verified by the sleeping cow shown above. Passive stay apparatus (special locking mechanisms in the limbs) mean that some hoofed mammals can remain standing virtually indefinitely, and such mammals (which include deer, rhinos and horses) are therefore capable of sleeping while standing. Having said that, they will still lie down to rest when they want to. I photographed the donkey shown here as I initially thought it was dead.

It used to be thought that the biggest extant hoofed mammals - the giraffes - did not sleep at all, but this was shown to be incorrect by Grzimek (1956) who demonstrated that adult giraffes slept recumbently on the ground for short periods (2.5-6 minutes), resting the head on the hindquarters or ground and holding the neck in an arced posture. This posture is not unique to giraffes but is also practised by okapis and some bovids. The sleep behaviour of giraffes was looked at in detail by Tobler & Schwierin (1996) who showed that giraffes slept both while standing and while recumbent for a total of 4.6 hours per 24 hours.

sleeping_baby_giraffe.jpg

Evidently little known is that elephants lie down on the ground to sleep properly: even today it seems widely thought that elephants 'crush themselves under their own weight' if they lie down, but this is just not true. In the wild, elephants sleep for 1-4.5 hours (per 24) in a recumbent position, with individuals showing preferences for sleeping on their left or right sides. Tobler (1992) reported that captive Asian elephants slept in a recumbent position for an average of 72 minutes a night. Elephants have been photographed propping themselves up against termite mounds and other structures when lying down to sleep, and in captivity they have been reported to construct 'pillows' from straw. The photo at the top shows a sleeping captive elephant (from wikipedia): for images of African elephants sleeping recumbently in the wild go here.

Large birds are also now known to recline on the ground for at least short periods. Burton (1969) reported how a scientist at Frankfurt Zoo monitored the sleep behaviour of ostriches: for 7-8 hours per night, they slept lightly with the neck erect. But for an average of 9 minutes each night they would lie on their sides, stretch their necks and legs out on the ground, and sleep deeply, during which time they could not be roused by lights or noises (except the very loudest noises). Emus, incidentally, are said to sleep deeply with the head and neck resting on the back, rather than lying on the ground.

Moving to other birds, terrestrial, ground-foraging birds like pheasants roost on the ground or in the low branches of trees, and simply huddle up and tuck in their appendages. I have been interested to learn, however, that small passerines (and perhaps others) don't sit out on branches during the night, but secrete themselves into small spaces. I know this because, while creeping through the woods at night once (don't ask), I discovered a Blue tit Cyanistes caeruleus tucked deep within a crevice in the bark of a tree, only its tail tip sticking out. It seems that most small birds climb deep into foliage, into crevices or cavities to sleep, and don't sit out in the open. Remember this next time you unnecessarily tear down the ivy from a tree or fence. Some passerines, including wrens, weavers and bush-tits, form large huddles within cavities (Perrins 1976). Burton (1969) wrote that birds which climb vertically on substrates, like nuthatches and treecreepers, also sleep in vertical postures, with nuthatches resting head-down and treecreepers resting head-up. Woodpeckers are also reported to sleep clinging vertically to vertical trunks (Perrins 1976). A few passerine species have been reported sleeping on their nests even when the nests were empty of eggs.

King_penguin_sleeping.jpg

According to Burton (1969), birds do not put their heads under their wings when they sleep, but instead bury the bill into the scapular feathers. He also stated that penguins are the only birds that properly hide the bill under the wing: they can't submerge any part of their head within their scapular feathers, because their feathers are so short.
I wonder what the hesperornithines (which lacked long feathers and possessed strongly reduced wings) did [adjacent King penguin photo by J. P. Matth from here].

So far I haven't mentioned the fact that some animals sleep with only 'half of the brain': this is called unihemispherical sleep, and it allows the animal to stay partly alert while in a restful state. Animals capable of this may be able to keep one eye open and alert for danger (as in, for example, wading birds that rest while exposed in open habitats like mudflats), or may be able to regulate movement while also resting. The latter ability is important to marine mammals, which doze at or near the water surface and still need to regulate their breathing. Cetaceans sleep unihemispherically, and a number of pinnipeds are also reported to sleep floating in the water, hanging in a vertical posture in the case of Grey seals Halichoerus grypus, Northern fur seals Callorhinus ursinus and walruses Odobenus rosmarus. Weddell seals Leptonychotes weddelli are reported to do this while submerged at their breathing holes in the ice. It has also been noted that some pinnipeds sleep underwater in cetacean fashion, 'rising periodically, while still asleep apparently, to take a breath and then sinking again to continue their slumber' (Burton 1969, p. 27). This is correct, as verified by this photo (below) of a Harbour seal Phoca vitulina sleeping underwater (from Rock Paper Lizard - thanks to Vasha for the heads-up).

sleeping_seal_from_Rock_Paper_Lizard.jpg

This has been nothing like a thorough review of the subject and, as I said at the start, it's been very superficial and I haven't looked at the big questions about sleep behaviour. For tetrapods at least, sleep would appear to be essential, even if members of some lineages can get by on very little. Speaking of which, at the time of writing it's gone 2 am. Good night.

Refs - -

Burton, M. 1969. Sleep and Hibernation in the Animal World. Odhams Books, London.

Grzimek, B. 1956. Schlaf von Giraffen und Okapi. Naturwissenschaften 17, 406.

Kavanau, J. L. 1998. Vertebrates that never sleep: implications for sleep's basic function. Brain Research Bulletin 46, 269-279.

Perrins, C. 1976. Bird Life: An Introduction to the World of Birds. Magna Books, Leicester.

Rattenborg, N. C. 2006. Do birds sleep in flight? Naturwissenschaften 93, 413-425.

Tobler, I. 1992. Behavioral sleep in the Asian elephant in captivity. Sleep 15, 1-12.

- . & Schwierin, B. 1996. Behavioural sleep in the giraffe (Giraffa camelopardalis) in a zoological garden. Journal of Sleep Research 5, 21-32.

Comments

Totally fascinating and, as far as I know, all correct information. It is funny how sleep researchers rarely pay attention to stuff like sleep position, as opposed to EEG data, sleep duration, timing and patterns.

Posted by: Coturnix | September 15, 2008 7:32 AM

On sleep in seals, see recent blog post at Rock Paper Lizard

Posted by: Vasha | September 15, 2008 7:35 AM

I kept having to stop reading this article to read bits out loud to my wife. I really, really hope it's part 1 in a series. Fascinating.

Posted by: Michael P. Taylor | September 15, 2008 8:08 AM

Very interesting post. Thanks.

Posted by: Casz | September 15, 2008 8:39 AM

How about amphibians? I've noticed my axolotls and xenopus toads are less active in the day, but as they have no eyelids how can I tell when they're asleep?

Posted by: Mo Hassan | September 15, 2008 9:55 AM

I work at a wildlife rehab center in the US and at night we have to catch songbirds in their aviaries to be released the next day. Once it gets dark they go sleep, usually concealed in foliage. Wrens can be very hard to find. Jays and thrushes perch up high and are conspicuous. With just a little light you can catch them easily by hand. Small birds will cluster and you can catch 4-5 with one hand. If you miss they quickly come to life and can only be caught with a net.

Greg

Posted by: tigerbird62 | September 15, 2008 11:37 AM

On the subject of small passerines sleeping in samll crevices, I think its come for blue tits etc to roost in nest boxes in groups up to 10 or 20, huddling together to preserves heat on cold winter nights

Posted by: Neil | September 15, 2008 12:19 PM

^^^ 'come' is meant to read common

Posted by: Neil | September 15, 2008 12:20 PM

Very interesting little overview. Especially with the pictures. Though I realize it's an unscentific reaction -- the giraffe sleep posture looks very uncomfortable.

I'm kind of interested in hooved animals' ability to sleep while standing. As well as a bat's ability to sleep hanging upside down (which you didn't mention), I'd be curious to know a little about how those strategies evolved.

Posted by: GAC | September 15, 2008 1:22 PM

Fascinating post, Darren. Of course, in some animals its impossible to tell. My lungfish seems to sit on the bottom of his tank without moving most of the time (and of course, no eyelids to close).

Posted by: Graeme | September 15, 2008 1:49 PM

I always wonder how my frog-eyed gecko sleeps without eyelids. The only way you can tell he's asleep is that his legs are sprawled out and his pupils are impossibly thin. I suppose that's akin to closing one's eyes--if you close off the pupil, you're achieving much the same result as closing your eyelids.

Posted by: Zach Miller | September 15, 2008 3:13 PM

My ferret curls up in a tight ball, even though hes so long. (he never flops out of one side like you see cats and dogs do) He sleeps up to 20 hours a day, and is active at random times.

I cant sleep at night unless I spend some time on my belly. sometimes I sleep in this position, sometimes not. Im told the optimal positions on my back, but I dont like it, and how many people do sleep that way??

Posted by: Tengu | September 15, 2008 3:41 PM

There is a good deal of information about the roosting (and other) habits of Blue Tits and Great Tits in "Birds as Individuals" by Len Howard.

Posted by: Don Cox | September 15, 2008 3:45 PM

I cant sleep at night unless I spend some time on my belly. sometimes I sleep in this position, sometimes not. Im told the optimal positions on my back, but I dont like it, and how many people do sleep that way??

I can only fall asleep on my belly. Except when I'm very tired; in that case I can fall asleep on my back -- but then I always soon get a nightmare and wake up! I do occasionally turn around, but falling asleep only works well on the belly.

Sleeping on a side is impossible -- leads to squeezing the blood off in the lower shoulder.

What's so uncomfortable about lying on my back is that I have to lift all inner organs to breathe in. I'd need gastralia to prevent abdominal collapse. :o) (Wait -- that can't be it. Crocodiles have gastralia and faint when turned on their backs.)

Two probable reasons:
- People with Asperger's "syndrome" like pressure. I have a few symptoms, and I like pressure on the belly.
- When I was a baby, all doctors believed laying babies on the belly was best, so that's what was done with me. Later, laying babies on the back became fashionable (apparently there's even a connection between lying on the belly and sudden infant death); and indeed, like the rest of the family, my little sister much prefers lying on her back...

Posted by: David Marjanović | September 15, 2008 6:35 PM

I do occasionally turn around

while asleep, that is. I sometimes wake up on my back. I should try to find out if that's correlated with waking up from a dream -- though probably it isn't strongly correlated.

Posted by: David Marjanović | September 15, 2008 6:37 PM

"I can only fall asleep on my belly. Except when I'm very tired; in that case I can fall asleep on my back -- but then I always soon get a nightmare and wake up!"

This is weird, I ususally can't sleep on my back for the same reason. I tend to have nightmares in this position, I don't know why. My mom was one of those people who believed that babies should sleep on their stomachs. She thought we would end up with flat heads if we slept on our backs...Seriously.

Posted by: Susan | September 15, 2008 7:01 PM

I have a friend who is a very committed vegetarian. If we eat out together I usually avoid ordering meat just to prevent an arguement. She grew up on a farm and told me that whenever you see a cow or a horse lying in a field it means that they're sick. According to her, it's unnatural for these animals to sleep lying down. I know she's wrong but I wonder if this is a common belief among vegetarians? I usually don't bother discussing it because she lived on a farm and I didn't so therefore she knows more then me. But still...Has anyone else heard of this before?
I also wanted to extend my condolences to Darren, I was sorry to read about your loss in a previous post.

Posted by: Susan | September 15, 2008 7:16 PM

There is a good deal of information about the roosting (and other) habits of Blue Tits and Great Tits in "Birds as Individuals" by Len Howard.

In somewhat the tradition of Walking With Dinosaurs, I think there should be a show that examines the sleeping behavior and posture of birds, and it should be titled Sleeping With Great Tits (And Other Nocturnal Adventures With Various Birds).

Posted by: Stevo Darkly | September 15, 2008 7:22 PM

"The king sleeps on his back, a wise man on his side, a businessman on his stomach."

Posted by: Nathan Myers | September 15, 2008 8:01 PM

@Susan: I have no idea about your question, but why should it have anything to do with her being vegetarian? I think that belief would much more likely to be connected with growing up on a farm than with being a vegetarian.

Posted by: GAC | September 15, 2008 8:29 PM

I think what Susan means is that people who adopt vegetarianism for primarily emotional reasons may have a tendency to assert any odd factoid they may have come across that seems to support the idea that primates can't digest meat properly, or that hunting is cruel and unsustainable, or that farmed animals are all sick and miserable. And so on. I'm not taking issue with the generalisations, just noting the prevalence of confirmation bias in those committed to a position one way or the other, as on any question.
Growing up on a farm makes it likely this person has seen farm animals sleeping lying down (but probably not often), and is also familiar with the same animals sleeping while standing. Also, some farm animals get sick, and many people understandably feel icky about the idea of eating sick animals. Makes sense to err on the side of caution, not to eat animals that sleep lying down? But there's only one way to be sure...

Posted by: John Scanlon FCD | September 16, 2008 12:06 AM

@John: Makes sense. I'd think it's definitely plausible that she doesn't eat meat because she grew up on a farm and got attached to the animals (or, conversely, saw firsthand how dirty and disgusting they can be). But I was just pointing out that, since we didn't know precisely WHY she was a vegetarian (plenty of people do it for perceived health benefits, independent of any feelings toward the animals), we can't necessarily see how that would affect her ideas about animals.

Besides that, I've heard the "only sick horses/cows lie down to sleep" meme from sources that didn't seem to be advocating animal rights craziness. I dunno, though.

Posted by: GAC | September 16, 2008 1:22 AM

Large seals sleep during deep descending dives, and while ashore sleep in apnea (breath hold). Anadder blog had a bit on sleep today too: Anadder
where I commented that sleep is "anti-photosynthetic" or so. Eyelids are primarily sleep/burrowing related, I'd guess, as opposed to dust / foreign-object protection.

I wonder if the pupillary light reflex of the oculomotor cranial nerve is derived from primarily sleep or from focusing, I'd guess sleep came first.

Possible correlation to the photic (solar powered) sneeze and dark adaptation in the trigeminal cranial nerve in humans?

Posted by: DDeden | September 16, 2008 2:06 AM

How did you know Ive got aspergers, David?

Don, I have that book, `Birds as individuals` its a fastinating read. how did they get all those birds to live with them? Darren, you should read that book.

Posted by: Tengu | September 16, 2008 3:20 AM

This thread has reached a new low for TetZoo. I am very disappointed in those people who posted disparaging speculations on why certain people have chosen not to eat other animals.

Vegetarians have many different, often entirely rational reasons for their choice. You cannot evaluate how rational their choice is without determining how rational is your own, a task that you cannot (by definition) do with any objectivity.

"If we weren't meant to eat other people, why do they taste like pork?"

Posted by: Nathan Myers | September 16, 2008 5:39 AM

How did you know Ive got aspergers, David?

Do I know that? Interesting. :-)

where I commented that sleep is "anti-photosynthetic" or so.

What, if anything, does that mean?

Eyelids are primarily sleep/burrowing related, I'd guess, as opposed to dust / foreign-object protection.

Eyelids are obviously related to keeping the eye from drying out. That's why you blink when you're awake, why fish and larval amphibians lack eyelids, and why permanently aquatic amphibians never grow eyelids.

(Geckos, snakes and other have fused eyelids with a transparent window in them.)

Posted by: David Marjanovi?, OM | September 16, 2008 8:43 AM

On the opposite side, many primates sleep very long. Wild gibbons fall asleep long before dusk. I also seen chimps and monkeys in a zoo sleeping already in afternoon.

Apparently, ground squirrels who hibernate and periodically wake up for short periods, also have episodes of normal sleep between hibernation.

Off-topics, I cannot fall asleep without turning a few times to lay on left and right side. I guess it is some innate behavior for stretching muscles. I also find very interesting how people pick beds in a room. When you have several people in a dormitory, always the most distant (and hidden) places are picked first. I suppose, its about picking most safe place.

Posted by: Jerzy | September 16, 2008 9:29 AM

Do any tetrapods other than humans ever have sleep apnea? Specifically, the kind induced by malformations of the back of the palate, smooth muscle in the back of the throat closing inappropriately, etc.? Even any other primates? It seems like this would be the kind of thing natural selection would eliminate...except in humans, apparently (says the sufferer of said condition)...

Posted by: Jerry D. Harris | September 16, 2008 9:46 AM

Re; vegitarianism, a lot I have met are `not` rational, (any more than certain religious food prohibitions are rational) and are the sort of silly, holier than thou types that need poking fun at.

All food fads are ghastly, particularly those inflicted upon others.

At the end of the day, this could mean simply we have too much to eat. (and no, Im not one of those well fed people who think austerity is A Good Thing.)

Posted by: Tengu | September 16, 2008 10:37 AM

I cannot fall asleep without turning a few times to lay on left and right side.

I have to do that with my head.

Posted by: David Marjanović | September 16, 2008 11:11 AM

"The king sleeps on his back, a wise man on his side, a businessman on his stomach." Where does that remarkable quotation come from? No sign of it on Google.

"I also seen chimps and monkeys in a zoo sleeping already in afternoon." Many people have a siesta in the early afternoon.

Posted by: Don Cox | September 16, 2008 2:22 PM

No sign of it on Google? Enter

king "sleeps on" back "wise man" "on his side" stomach
and hit "I'm feeling lucky" and see what shows up.

Seriously, it does appear on three other pages, but with no more attribution. I recall it from decades ago, but cannot guess where. Variations that Google finds are "The poor man", "rich man", and "fool" for "businessman".

Posted by: Nathan Myers | September 16, 2008 5:06 PM

I too find it difficult to fall asleep when lying on my back. If I do, and am not wakened too quickly, I still wake up in some other orientation.

Posted by: Andreas Johansson | September 16, 2008 5:49 PM

This is weird, I ususally can't sleep on my back for the same reason. I tend to have nightmares in this position, I don't know why.

I'm the same. My guess is that in that position my airflow eventually gets blocked. If I'm dreaming, the fear that arises from being unable to breathe turns the dream into a nightmare. I distinctly recall certain dreams going from benign to terrifying right before waking up with a snort.

My pet conure sleeps in a little tent in her cage, with her tail sticking out. She'll also nap on a branch with one leg up, but that's usually only for short periods of time. Her preference by far is to fall asleep on my leg with my hand over her like a blanket. Terribly difficult to get any work done when she wants to nap.

Posted by: pough | September 16, 2008 8:14 PM

On pets: Tigger Mamum-Ra the tabby cat sleeps during the day in a pose that I call 'recumbent gorgosaur'. Ten points to the person who can explain what the hell I'm talking about.

Posted by: Darren Naish | September 16, 2008 8:20 PM

Its a reference to Laurence Lambe's restoration and description of Gorgosaurus. i.e. lying straight out on the stomach with the legs folded parallel to the body.

Google turns up a sketch by John Sibbick, and there's a partial quote out there about "hunger impelled wanderings", but I'm sure there was a fuller quote in Adrian Desmond's "Hot-Blooded Dinosaurs". Which of course I don't have to hand.

Oh, and I'm another person who sleeps on his stomach.

Posted by: Dave Godfrey | September 16, 2008 9:08 PM

Translated to the African mythos, it might be, "Lion sleeps on his back, rabbit on his side, jackal on his stomach" -- all of which seem a little unlikely.

Posted by: Nathan Myers | September 16, 2008 10:25 PM

Neat post. Too bad there was no reptile love in the sleep survey, but then I'd imagine herp related sleep studies are probably few and far between in the literature (though I think I remember reading about unihemispherical sleep existing in sea turtles).

Posted by: Jura | September 17, 2008 1:58 AM

A good follow up study would be on animals sleeping on other animals. Like cats on people, or passerines on cattle.

Posted by: Alan Kellogg | September 17, 2008 4:12 AM

My guess is that in that position my airflow eventually gets blocked. If I'm dreaming, the fear that arises from being unable to breathe turns the dream into a nightmare. I distinctly recall certain dreams going from benign to terrifying right before waking up with a snort.

That is definitely not the case in my case (er... yeah). When my nose closes due to excess production (I'm allergic to dust mites), and I happen to be dreaming, I dream that my nose closes; it doesn't get reinterpreted, it enters the dream directly. And when I have a cold, I often try to sleep on my back because that way it's easiest for liquids to leave the nasal cavity. Tends to work.

Posted by: David Marjanović | September 17, 2008 8:46 AM

Wow. This really has become the David Marjanovic Show :)

I tried sleeping on my front last night (at 03-40, the earliest I could get to bed). I cannot do it, have to be on my side. Well done Dave (Godfrey)... the 'recumbent gorgosaur' thing refers to Lawrence Lambe's inferred resting posture for Gorgosaurus (depicted in Fig. 48d in Lambe (1917)), later re-drawn by John Sibbick. Lambe wrote 'This position of rest, and particularly the recumbent one of repose at full length were probably those [sic] most frequently assumed by a reptile having the form, and the supposed sluggish disposition of Gorgosaurus' (p. 83). Yes, whenever I look at a tyrannosaur I think 'sluggish' (I don't: that was a joke).

Incidentally, my hard-copy of Lambe (1917) is copied from the personal copy that once belonged to Alick Walker. It's covered with annotations where he compared the tyrannosaur's anatomy with that of Ornithosuchus.

Ref - -

Lambe, L. M. 1917. The Cretaceous theropodous dinosaur Gorgosaurus. Memoirs of the Geological Society of Canada 100, 1-84.

Posted by: Darren Naish | September 17, 2008 9:26 AM

Great summary! I also hope that it is only the first of several posts on the subject.

Alexander Skutch wrote a wonderful overview of sleeping behavior in birds:

Skutch, A.F. (1993) Bird asleep, University of Texas Press,

220p.

Posted by: Hector Gómez de Silva | September 17, 2008 1:02 PM

Ducks (mallards) exhibit unihemispheric slow-wave sleep (USWS).

Ref:
Rattenborg, N et al (1999)Facultative control of avian unihemispheric sleep under risk of predation. Behavioural Brain Research 105, 163-172.

Posted by: Rose | September 17, 2008 2:25 PM

OK, I'm shutting up already :o)

Posted by: David Marjanović | September 17, 2008 4:55 PM

When I look at the old restorations (including skeletal ones) most dinosaurs tend to look "sluggish", with the dragging tail, you end up with a "stunned kangaroo" look. This sort of thing is one of the few times when post-modernism actually works (i.e. different ways of looking at the same evidence, influenced by different paradigms.) Mantell initially reconstructed Iguanodon with the proportions of a lizard, and these days we're disappointed if a maniraptoran doesn't get feathers. Charles Knight et al fall somewhere between these two extremes. (OK I know one is closer to the truth than the other two, but that's not my point).

Posted by: Dave Godfrey | September 17, 2008 7:52 PM

One thing we can all agree on: sleeping animals are cute. Especially sleeping baby animals. Awwwwwww.

Posted by: susan | September 18, 2008 8:27 AM

Thanks, nice article.

Posted by: NaUm | September 18, 2008 11:12 AM

Regarding the poikilotherms, there's a somewhat applicable quote from Charles Elton:

All cold-blooded animals... spend an unexpectedly large porportion of their time doing nothing at all, or at any rate, nothing in particular.

Perhaps they're sleeping?

I don't know who Charles Elton is / was; this quotation comes from the front of Pennak (1978), a reference book about the aquatic invertebrates of eastern North America.

Another person here hoping for a follow-up article with more odd sleeping animals. Thanks for putting this up.

Posted by: TheBrummell | September 20, 2008 9:34 AM

Is that penguin sleeping? Also do all girrafes sleep like that or is that just that one? How do elephants get up?

Posted by: seanupshaw | November 10, 2008 1:21 PM

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