Tetrapod Zoology

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Hooray: another of those articles that I’ve been promising to publish for weeks and weeks.

Thanks mostly to the importance of the species in the international pet trade, the Green iguana Iguana iguana is typically imagined as a rather uninspiring lizard that sits around on branches all day long, occasionally munching on salad or sitting in its water bowl. It’s true that some captive individuals become remarkably charismatic and idiosyncratic, but for the most part the Green iguana is generally thought of as a rather dull animal that doesn’t really do much of interest. Today we’re going to change all that (I hope). Field studies stretching back over three decades have demonstrated beyond question that the social life of the Green iguana is remarkable and complex, and if you’re unaware of the sorts of behaviours that have been reported for these lizards, you might be surprised…

Firstly, Green iguanas can be described as leading fairly active social lives, at least during the breeding season. They are territorial lizards with a lek-style breeding system: males choose exposed arboreal display sites, often selecting trees that are dead or sparsely vegetated. They proclaim their territory with lots of head-bobbing and displaying of the large dewlap. Any males that try to move in to the area are chased away, but between four and eight females move into the territory, where they compete among themselves for access to high-quality territory-holders (Burghardt 2002). Males go without eating during this territorial breeding phase, they have to put energy into changing their appearance (they change colour from greenish to orangish and increase the size of their jowls), and they also have to chase off rival males, and woo and mate with females. But as has just been revealed by a new study of another lekking iguana, the Galapagos iguana Amblyrhynchus cristatus, females may incur high energetic costs in discriminating among potential mates (Vitousek et al. 2007), so they don’t necessarily have an easy time either.

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Incidentally, not all male Green iguanas are big, showy, territorial animals. Some are small, superficially female-like, and more cryptic. Rather than attracting females, they try to mate with them forcibly. Cryptic males that morphologically mimic females have now been documented in quite a few tetrapod species: we’ve previously encountered this sort of thing when covering newts and sheep.

During January and February, mated females migrate to favoured nesting areas. And they really do migrate – in the case of the well-studied Panamanian iguanas that breed on the island of Slothia, in Gatun Lake, the females travel up to 3 km in order to reach their nesting area (as demonstrated by radio tracking: Montgomery et al. (1973)). They walk across land and then swim to the island. They clearly come to Slothia to nest and not much else, and don’t live on the island during the rest of the year. While Green iguanas in parts of their range nest individually, those that come to Slothia nest colonially and therefore synchronize their nesting. As many as 150-200 female Green iguanas gather together at favoured clearings, and here they compete with one another for access to the best nesting areas.

On Slothia, iguanas shared a nesting area with an American crocodile Crocodylus acutus over three successive nesting seasons. It’s possible that the iguanas did this so that their nests and/or juveniles might receive protection from the crocodile, but it seems most likely that the two species were brought together because of similar nest requirements, and neither seemed to benefit from the proximity: the crocodile both interrupted iguana nesting activities and killed and ate some of the iguanas, and the iguanas sometimes dug up the crocodile’s eggs (Dugan et al. 1981, Bock & Rand 1989).

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Iguana nests are not just scrapes in the ground: rather, the iguanas build complex burrow systems that become deeper and increasingly complex over the lifespan of a nesting colony (Bock & Rand 1989). While female iguanas do not nest-guard as crocodiles do, they may stay with the burrow for a day or two after laying, defending it from other females. Presumably this is an attempt to stop later-nesting females from digging up the clutch when creating their own nests.

During the first week of May, the baby iguanas start to hatch. Emerging from the central chamber of the buried nest, they dig their way to the surface. This takes up to seven days. However, they don’t simply emerge and then dash off into the forest, alone. On emerging, they sit with just their heads poking out, sometimes for as long as 15 minutes, and sometimes repeatedly disappearing and reappearing from the nest entrance. It seems likely that the iguanas are looking out for predators, but what’s particularly interesting is that the iguanas don’t just look out for predators, they also spend a lot of time observing other baby iguanas emerging from other nest holes (Burghardt 1977, Burghardt et al. 1977). By observing the behaviour of other hatchling groups, iguanas seem to decide whether or not it is safe to leave the nest. Burghardt (1977) reported cases where baby iguanas belonging to four different clutches all emerged synchronously, an observation which led him to conclude that ‘nest emergence seems socially facilitated by visual cues’ (p. 183). This is a far cry from the stereotypical image of the baby reptile crawling from its nest and immediately dashing off headlong into cover.

Juveniles form pods, usually consisting of about four individuals. They indulge in a tremendous amount of social behaviour of the sort typically regarded as unique to mammals and birds, rubbing their bodies and heads against one another, displaying their dewlaps, nodding their heads and wagging their tails at each other. They engage in allogrooming (grooming other members of the social group). The young iguanas stay associated at night, when they sleep in close physical contact with other pod members, sometimes even lying on top of them. Baby iguanas definitely recognise their own kin, apparently using olfactory cues (Werner et al. 1987), and continue to stay with them for many months after hatching (Burghardt 2002).

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While these pods obviously consist of siblings, they seem to exhibit some sort of structure, with one iguana acting as leader. Burghardt (1977) described and photographed cases where juveniles followed one another in a line through vegetation and across the ground, with the iguana or iguanas in the lead often looking behind to, apparently, check on the progress of the followers. Those iguanas that hatched on Slothia have to, at some point, leave the island and swim to the mainland, and to do this, they have to make their way through a reed-bed before setting out across the water. Prior to departing, the juveniles were seen to engage in lots of head-rubbing and other physical contact, and the individual that appeared to lead the group was the one that engaged in the greatest amount of these activities. Invariably this was the first animal to enter the water and start swimming. If its companions failed to follow, it would return to shore. The iguana identified as ‘leader’ was also reported to disappear into the reeds and reappear with additional recruits [unfortunately I couldn't find any available photos of wild groups of juveniles, hence the captive group shown in the adjacent image].

It is perhaps tempting to think from this that iguanas ‘look out’ for each other, or at least for their siblings. Such an interpretation might seem anthropomorphic, but it isn’t necessarily, because we know from studies on diverse species that kin selection can lead members of some species to exhibit behaviours that might favour the survival of their brothers and sisters. In Green iguanas, the idea that individuals really do ‘look out’ for siblings has received robust experimental support from studies of anti-predator behaviour. Noting that male and female young Green iguanas exhibited quite different types of anti-predator behaviour, Rivas & Levín (2004) flew model hawks at both lab-based and wild groups of Green iguana siblings. They showed that, while females tended to hide, stay motionless, or run away from the potential predator, males exhibited far more interesting and unusual behaviours: they ran in front of the model hawk, appeared from beneath cover (rather than hiding within it), and – most interestingly – covered their smaller female siblings with their own bodies, thereby concealing them from view.

It’s possible that these behaviours might be selfish: the unusual responses of the males ‘might surprise a searching predator and give the escapee more time to escape at the expense of the remaining animals’ (Rivas & Levín 2004). But it appears more plausible that this ‘covering behaviour’ is a hitherto undocumented form of fraternal care, where males are actually protecting their female siblings. If you want to know more about this study, you can read it yourself here (from Rivas’ iguana page).

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So there we have it. The significance of this behavioural complexity will not, I’m sure, have been lost on you. The stereotyped idea that lizards and other reptiles are far ‘simpler’ in behaviour and social life than birds and mammals is demonstrably false, and many forms of behavior long imagined as uniquely avian/mammalian are actually far more widespread. Other complex behaviours that I haven’t discussed have also been documented in iguanas, including appeasement behaviour for example. Recall that we now think that play behaviour is practiced by reptiles including turtles, crocodilians and monitor lizards (for more on this subject see the ver 1 article Dinosaurs come out to play). So go forth and spread the word.

PS – this article was originally going to be titled ‘Sorry Sarah Connor: that iguana really doesn’t like you’, but I couldn’t get a screen-capture of the appropriate scene in time. Extra points if you know what the hell I’m talking about.

PPS – I’ve just learnt that Discovery News did a piece yesterday about the Galve paper I recently published with Barbara Sánchez-Hernández and Mike Benton: the news article is here.

Refs – -

Bock, B. C. & Rand, A. S. 1989. Factors influencing nesting synchrony and hatching success at a green iguana nesting aggregation in Panama. Copeia 1989, 978-986.

Burghardt, G. 1977. Of iguanas and dinosaurs: social behavior and communication in neonate reptiles. American Zoologist 17, 177-190.

- . 2002. Walking with iguanas. BBC Wildlife 20 (5), 60-65.

- ., Greene, H. W. & Rand, A. S. 1977. Social behavior in hatchling green iguanas: life at a reptile rookery. Science 195, 689-691.

Dugan, B. A., Rand, A. S., Burghardt, G. M. & Bock, B. C. Interactions between nesting crocodiles and iguanas. Journal of Herpetology 15, 409-414.

Montgomery, G. G., Rand, A. S. & Sunquist, M. E. 1973. Post-nesting movements of iguanas from a nesting aggregation. Copeia 1973, 620-622.

Rivas, J. A. & Levín, L. E. 2004. Sexually dimorphic anti-predator behavior in juvenile green iguanas Iguana iguana: evidence for kin selection in the form of fraternal care. In Alberts, A. C., Carter, R. L., Hayes, W. K. & Martins, E. P. (eds) Iguanas: Biology and Conservation, pp. 119-126.

Vitousek, M. N., Mitchell, M. A., Woakes, A. J., Niemack, M. D. & Wikelski, M. 2007. High costs of female choice in a lekking lizard. PLoS ONE 2 (6): e567. doi:10.1371/journal.pone.0000567 [available here]

Werner, D. I., Baker, E. M., Gonzalez, E. del C. & Sosa, I. R. 1987. Kinship recognition and grouping in hatchling green iguanas. Behavioral Ecology and Sociobiology 21, 83-89.

Comments

  1. #1 John Scanlon, FCD
    July 12, 2007

    Amazing. I have a back yard full of iguanian lizards (Lophognathus gilberti, Agamidae) and they can be quite entertaining during the warmer months, but nothing as complex as these big, boring herbivores you’re talking about.
    Can’t guess exactly what scene of ‘Terminator’ you’re referring to, but I think the iguana was only on screen briefly two or three times – hiding in a kitchen cupboard, and maybe doing an open-mouth display at one point – was that it?

  2. #2 Darren Naish
    July 13, 2007

    … hiding in a kitchen cupboard, and maybe doing an open-mouth display at one point – was that it?

    Hi John. Yeah, it’s the open-mouth display scene I had in mind. Sarah says that Pugsley – or whatever his name is – still loves her, but his hissing, open-mouth display, and extended dewlap indicate otherwise. By the way, this doesn’t stop Terminator from being one of the best films of all time. If I opened a nightclub I’d call it Tech Noir.

  3. #3 David Marjanovi?
    July 13, 2007

    There’s an iguana in Terminator? I totally missed it. :-]

  4. #4 Sordes
    July 13, 2007

    Those habits of juvenile iguanas are really highly interesting. I really had not imagined this. But it is really so that social behavior is something most people never associate with reptiles or even amphibians. But if you take a closer look, you will find many amazing types of social behavior among reptiles or amphibians. Just think about the Solomon Island skink, which take care to their youngs for a very long time. Or those family-living Egernia saxatilis or the side-blodged-lizards.

  5. #5 jackd
    July 13, 2007

    They proclaim their territory with lots of head-bobbing and displaying of the large dewlap.

    We’ve been enjoying watching Carolina anoles (Anolis carolinensis) do the same thing. Until reading this I hadn’t realized anoles and iguanas were so closely related, which makes the similar behavior seem less surprising.

  6. #6 Hai~Ren
    July 13, 2007

    Really, really fascinating information. Who’d have thought? Turns out that the extant reptiles have more complex social lives than we once thought.

    I wonder if anyone has tried to compare the brain anatomy of different lizard species, or to see which lizards had a higher EQ or were more intelligent. I won’t be surprised if the teiids, varanids and the larger iguanids rank among the most intelligent lizards.

  7. #7 Mike Keesey
    July 13, 2007

    “Cryptic males that morphologically mimic females have now been documented in quite a few tetrapod species.”

    Not to mention our own (“metrosexuals”). :)

  8. #8 The Carrot
    July 13, 2007

    Darren,

    Thanks for the article! I’ve been keeping (and breeding) green iguanas for the past fifteen years. They truly are fascinating animals that are much more interesting than the casual observer would believe.

    It’s a shame that iguanas are so popular in the pet trade. When people see my large male iguana due to his size they often think he’s a freak of nature, when in fact most people have no idea how large adult iguanas can become due to the fact that most of the cute baby iguanas seen in pet stores never survive long enough to become fully grown. Sad, really.

  9. #9 Darby
    July 13, 2007

    This is fascinating, but can you tell me why 10 – 15% of my webtraffic lately is coming from Google image searches for a jpg image I link to?

    They are cute, but the searches are originating from all over the world.

    I didn’t really figure you’d know, but it is iguanas, social, and weird, so it relates.

    I should mention, too, that my personal experience with captive iguanas suggests that their personalities are about as varied as cats, whatever that means.

    [from Darren: is that photo for real??]

  10. #10 rajita
    July 14, 2007

    This is a nice article– I must re-read it in leisure. It is good that you bring up “interesting” behavior in lizards — many people see the low EQ of lizards to mean a lack of interesting behavior. Is there any information whether these green iguanas are susceptible to diseases due to their social activity?

  11. #11 Tristram Brelstaff
    July 15, 2007

    Darren, it is nothing to do with iguanas, but have you seen this:
    Found: the giant lion-eating chimps of the magic forest

    [from Darren: yup, I saw the news when it broke. Cool stuff: Groves' mention of possible new subspecies status is totally contra to what has been announced before. More later]

  12. #12 Darby
    July 15, 2007

    I was doing a section on the evolution of the Galapagos species, and just wanted a nice picture of the green ancestors. And for picture links, this one is dramatic and shows the animals well, just what I look for when choosing one. The originating site seems to be just jpegs, so I can’t tell you anything about the image itself, but you’ve just written about iguanas relating to each other, and from personal experience I can tell you that they can get into some odd positions and they love to lick things. My suspicion is that it’s just a great captured moment.

  13. #13 nemo ramjet
    July 15, 2007

    Sorry to interrupt so needlessly, but did you receive my illstration for Quemisia? Cheers!

  14. #14 Steve Bodio
    July 15, 2007

    Wow!I know that more complex behavior is constantly being found in ‘lower’ taxons (Pianka and Vitt’s newish lizard book has some stuff on this and Carel somewhere mentions the playfulness of softshell turtles). But I had no idea that boring old iguanas had such complex lives.

    Though come to think of it even the little Sceloporus and Cnemidiphorus in the yard can be pretty ‘interactive’…

  15. #15 johannes
    July 17, 2007

    Ah, you mean the Sarah Connor from Terminator, not the german singer (sighs with relief). For a few minutes of horror I thought that the eurotrash virus had crossed the channel… :-)

  16. #16 Tim Morris
    August 18, 2009

    I read this, and other similar things about reptiles and their “love”, and I almost cry for how I was decieved as a child :(

    People go to such lengths to defend “fluffies” from cruelty, when “lower” animals get it just as bad.

    But then again, many mammals, equally fluffy get subjected to predjudice regardless, bats and lemurs.

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