See you down the puddle! Puddling in butterflies.

Turquoise (Doxocapa agathina vacuna) puddling on my jeans!
Turquoise (Doxocapa agathina vacuna) puddling on my jeans!

‘Puddling’ or ‘mud-puddling’ is when butterflies, moths and other insects settle on moist substrates to absorb liquids. Butterflies – particularly in the tropics – exhibit this puddling behaviour when feeding on moist ground, mud, animal excrement, rotting fruits, carrion, dung, bird droppings, sweat, tears and so on.

Common Redring (Pyrrhogyra otolais) mud puddling on mud!
Common Redring (Pyrrhogyra otolais) mud puddling on mud!

One might think that puddling butterflies are just having a drink, but it turns out to be a bit more complicated than that. For one thing, in butterflies at least, most of the individuals seen puddling are young males; females are rarely seen puddling, although it has been recorded in some older individuals.

Mixed group of butterflies puddling on sweaty wooden hand-rail
Mixed group of butterflies puddling on wooden hand-rail where sweaty hands have been placed

What puddling butterflies are doing is making up for the chemicals or nutrients they lack as a result of deficiencies in their diet as larvae. So puddling is a way of imbibing the salts and micronutrients they need when they become adults and start mating.

An aggregation of Broad-banded swallow tails (Heraclides astyalus) mud-puddling
An aggregation of Broad-banded swallow tails (Heraclides astyalus) mud-puddling

Butterflies need to stock up on both sodium (the Na in NaCl; sodium chloride or salt) and nitrogen (in the form of amino acids; the building blocks of proteins) because when they mate they will transfer substantial amounts of sodium and protein to the females in the form of a ‘spermatophore’. A spermataphore is a large – up to 10% of body mass in some butterflies like the Monarch – capsule of sperm and essential nutrients. Female butterflies lose a huge amount of their body salt – up to 75% – as a result of egg laying, so the male is in effect providing her with a gift; a so-called ‘nuptial gift’ of nutrients which he has obtained via puddling.

Cloudless Sulphur (Phoebis sennae) and White-angled sulphur (Anteos clorinde) butterflies doing a bit of mud-puddling
Cloudless Sulphur (Phoebis sennae) and White-angled sulphur (Anteos clorinde) butterflies doing a bit of mud-puddling

Providing the female with resources for her to use for making eggs, can be seen as the male’s contribution towards the process of reproduction. A sort of division of labour between the sexes: he spends time and energy obtaining and then passing on to her what she needs to produce their fertilised eggs. The quality of the nutrients in the spermatophore which the male provides to the female may therefore, improve her fertility and give their offspring the best chance in life (i.e. increase their fitness).

An Eighty-eight (Diaethria clymena janeira) butterfly puddling on sweat left on a wooden hand-rail
An Eighty-eight (Diaethria clymena janeira) butterfly puddling on sweat left on a wooden hand-rail

So puddling is a very important activity in the life of an adult male butterfly to obtain sufficient salts and nutrients to pass on in this way.

A mixed group of butterflies puddling on a discarded workman's glove (presumably nice a sweaty!)
A mixed group of butterflies puddling on a discarded workman’s glove (presumably nice and sweaty!)

Butterfly species differ with respect to what particular substrates they prefer to puddle on! Some species are more attracted to mud, others prefer dung and so on. Some species become specialised on one type of substrate – such as carrion – and presumably become highly adept at finding what they need. A source of nitrogen is very important for many butterflies, so puddling on dung and carrion is a good way of obtaining proteins as well as salts.

Unidentified South American butterfly puddling on some fresh dung!
Unidentified South American butterfly puddling on some fresh dung!
Common nawab (Polyura athamas) absorbing nitrogen by puddling on dead fish
Common nawab (Polyura athamas) absorbing nitrogen by puddling on dead fish

Some aggregations of butterflies seen next to streams may simply be imbibing water; especially in the tropics where temperatures are so high. An aggregation of male swallowtail butterflies of the species Papilio polytes, was even seen drinking seawater at low tide, on the island of Guam.

A group of Pale Jezebels (Delias sanaca perspicua) drinking or puddling? Northern Thailand
A group of Pale Jezebels (Delias sanaca perspicua) drinking or puddling? Northern Thailand

Many butterflies – including papilionids (swallowtails) and pierids (yellows and whites) – form large aggregations when puddling, which might be a way of minimising their exposure to predation, via safety in numbers.

Aggregarion of pierid butterflies: Cloudless Sulphurs (Phoebis sennae) and White-angled sulphurs (Anteos clorinde) and Large orange sulphurs (Phoebis agarithe)
Aggregarion of pierid butterflies: Cloudless Sulphurs (Phoebis sennae) and White-angled sulphurs (Anteos clorinde) and Large orange sulphurs (Phoebis agarithe)

There is no doubt an awful lot more to learn about what individual species are obtaining by puddling on different substrates and how this contributes to their success in reproduction. It comes down to insects getting what they need from their environment to best survive and perpetuate themselves.

Butterfly puddling on my sweaty finger!
Butterfly puddling on my sweaty finger!
The author as a sweaty substrate source for puddling butterflies!
The author as a sweaty substrate source for puddling butterflies!
  1. Boggs, C. L., & Jackson, L. A. (1991). Mud puddling by butterflies is not a simple matter. Ecological Entomology16(1), 123-127.
  2. Scriber, J. M. (2002). A female Papilio canadensis(Lepidoptera: Papilionidae) puddles with males. The American midland naturalist 147(1), 175-178.
  3. Otis, G. W., Locke, B., McKenzie, N. G., Cheung, D., MacLeod, E., Careless, P., & Kwoon, A. (2006). Local enhancement in mud-puddling swallowtail butterflies (Battus philenor andPapilio glaucus). Journal of Insect Behavior 19(6), 685-698.
  4. Oberhauser, K. S. (1988). Male monarch butterfly spermatophore mass and mating strategies.Animal behaviour 36(5), 1384-1388.
  5. Molleman, F. (2010). Puddling: from natural history to understanding how it affects fitness.Entomologia experimentalis et applicata134(2), 107-113.
  6. Beck, J., MuÈhlenberg, E., & Fiedler, K. (1999). Mud-puddling behavior in tropical butterflies: in search of proteins or minerals?Oecologia 119(1), 140-148.
  7. Pola, M., & García-París, M. (2005). Marine puddling in Papilio polytes (Lepidoptera: Papilionidae). Florida Entomologist 88(2), 211-

Tame me a Tegu!

Argentine black and white tegu (Salvator merianae)
Argentine black and white tegu (Salvator merianae)

I came across quite a few tegus – large lizards – when visiting Iguazu National Park in northern Argentina. As far as I can tell, they are all the Argentine black and white tegu (Salvator merianae), also called the Argentine giant tegu (1). They wander round quite happily, despite the presence of large numbers of people coming to see the waterfalls.

Argentine black and white tegu (Salvator merianae)
Argentine black and white tegu (Salvator merianae)

Tegus seem to eat just about everything! That is why are called omnivores: omni means ‘all’ or ‘every’. They will apparently consume invertebrates (millipedes, arachnids, insects and molluscs), vertebrates (birds, fishes, amphibians, lizards and small mammals), bird and turtle eggs, fruits, carrion and mushrooms (2). It’s not surprising given this diet that they grow so large! Over four feet long.

Argentine black and white tegu (Salvator merianae)
Argentine black and white tegu (Salvator merianae)

Tegus look a lot like monitor lizards (an Old World group) but they are not closely related, so must be assumed to have evolved to fill a similar niche (convergent evolution). There are 7 species of tegus, which were originally placed in a single genus, Tupinambis. But tegu taxonomy seems to have gone through some changes recently and the Argentine black and white, or giant tegu, was recently reclassified as Salvator merianae (3). For the record, the genus Salvator now consists of three species: S. duseni (The Yellow tegu), S. merianae and S. rufescens (The Red tegu), which mainly occur in open ecosystems (4). The genus Tupinambis now contains four species: T. longilineus (The Rondonia tegu), T. palustris (The Swamp tegu), T. quadrilineatus (The Striped or Four-lined tegu) and T. teguixin (The Golden tegu); and these tegus are mainly to be found in forest ecosystems (4).

Argentine black and white tegu (Salvator merianae) with a more golden pattern
Argentine black and white tegu (Salvator merianae) with a more golden pattern

Some of the tegus I photographed, like the individuals shown above and below, had a more yellow or golden colour and I wondered if these could have been Golden tegus (Tupinambis teguixin). Tegus are separated on the basis of a large number of characters, one of which is the number of loreal scales they have (3, 4). These are the large scales between the eyes and nostrils. All of the individuals shown here seem to have two large scales, so they cannot be T. teguixin, as that species only has one large loreal scale (4).

Argentine black and white tegu (Salvator merianae)
Argentine black and white tegu (Salvator merianae)

Surprisingly, tegus make good pets! Apparently, they are quite docile and can be house trained; they also get quite attached to their owners and get aggressive if they are not handled regularly! (1, 2). There are a number of Facebook sites (e.g. ‘The Tegu-phile’ and ‘Tegus from Around the World’) for people who love tegus! Unfortunately, not everyone appreciates their good nature and many are hunted for food and skins (5). Despite this fact, their numbers seem to be holding up.

There were also some very nice waterfalls as well as lizards!

Iguazu Falls in full spate from the Argentinian side
Iguazu Falls in full spate from the Argentinian side
  1. http://www.inaturalist.org/taxa/318758-Salvator-merianae
  2. http://carnivoraforum.com/topic/9508642/1/
  3. Harvey, M.B., G.N. Ugueto and R.L. Gutberlet Jr. 2012. Review of Teiid Morphology with a Revised Taxonomy and Phylogeny of the Teiidae (Lepidosauria: Squamata). Zootaxa 3459: 1-156.
  4. Passos, D. C., Lima-Araujo, F., Melo, A. C. B., & Borges-Nojosa, D. M. (2013). New state record and distribution extension of the golden tegu Tupinambis teguixin (Linnaeus, 1758)(Squamata: Teiidae) to the Caatinga biome, northeastern Brazil. Check List, 9(6), 1524-1526.
  5. Embert, D., Fitzgerald, L. & Waldez, F. 2010. Salvator merianae. The IUCN Red List of Threatened Species 2010:e.T178340A7526681. http://www.iucnredlist.org/details/178340/0

A fruit loving moth

Eudocima cocalus (Cramer) female
Eudocima cocalus (Cramer) female

This wonderful looking moth is in fact a bit of a pest, on account of the fact that it likes to make holes in fruits and suck out the juices! Hence it has acquired the name Cocalus Fruit Piercing Moth, which sounds very grand for such a little insect. This is the female shown here (above); the male does not have such nice white blotches. However, they can both pierce ripening fruits with their proboscises – penetrating the skin and pulp of fruit – to get at the juices. Unfortunately, this can cause crop losses of more than 50% in many crops – by allowing microorganisms to enter – such as Lychee and Carambola (1).

Seen in profile like this, the moth looks like it would blend in well on a lichen covered branch, but the hindwings (tucked away) have prominent yellow patches (2), which might be used to startle would-be predators? There is also a prominent ‘snout’ at the front end. I wonder what that is for? The antennae are long and thin and are tucked away in this photo; as is the fruit stabbing proboscis.

Also amazing to think that this noctuid moth was first described by Pieter Cramer in 1777! (3) I photographed it near Khao Yai, Thailand.

  1. Leong, S. C. T., & Kueh, R. J. H. (2011). Seasonal abundance and suppression of fruit-piercing moth Eudocima phalonia (L.) in a citrus orchard in Sarawak. The Scientific World Journal, 11, 2330-2338.
  2. http://www.projectnoah.org/spottings/37867071
  3. http://www.biodiversitylibrary.org/item/95175#page/142/mode/1up