Why Do Butterflies Puddle?

Puddling Butterflies - Why Do they Do It?

What is puddling?

The term "puddling" refers to the process in which butterflies 'sip' from puddles of water, mud, dung or carrion on the ground. Many species of butterflies are often encountered with their proboscis unfurled and probing into the ground to take in water and nutrients. At the muddy or sandy puddle (often tainted with animal urine or excreta), the butterfly sips water rich in mineral salts and other essential nutrients (mostly sodium chloride and nitrogen-rich solutions) that have leached from the surrounding soil and rocks. Male butterflies do more puddling than females.

Why do butterflies puddle?

The commonest explanation for this is that they are taking in water and nutrients. The dissolved salts and minerals may be used to make pheromones (that the male uses to attract females) and sperm. The main stimulant for puddling is sodium. The diet of adults (primarily nectar and fruits) lack sodium/salt, and the hostplants that they feed on in their younger larval stages do not provide enough sodium either. Sodium is vital for many physiological functions, including digestion, excretion, reproduction and flight. The males have to sustain high activity levels to be able to fly around and locate receptive females, so they have a need for higher energy and metabolism levels. Indeed, the males of many species are known to be more active and faster in flight than females.

So what is the function of sodium extracted during puddling?

Males seem to benefit from the sodium uptake through mud-puddling behaviour with an increase in reproductive success. Sodium is passed by the males to the females during mating. Males transfer this sodium and amino acids to the females together with the spermatophore during mating, as a nuptial gift - it enhances reproductive success. This nutrition also enhances the survival rate of the eggs.Each male can mate with multiple females, so with each copulation, the sodium/nutrient stock is depleted and he will have to puddle again.

Do butterflies of both sexes puddle?

It is usually the younger males that puddle. Females are known to puddle, but that is not the norm. There are various costs and risks associated with puddling, like predation or parasitism, and extra energy utilised. The females minimise these risks by not puddling. By not needing to puddle, they can focus their energies on obtaining more of the other kinds of nutrients, hunting out better oviposition sites, and on laying healthier, more fertile eggs.

How long is the puddling process?

A butterfly can puddle from anything between a few seconds to an hour or more, depending on a variety of factors. During the puddling process, butterflies suck the nutrient rich fluids through their proboscis, and "filter" it within their systems to extract the needed chemicals.

Where does the excess fluid go?

Obviously, fluid ejection results from a need to pass great quantities of soil moisture through the gut in order to extract sufficient useful, but dilute, chemicals. If a particular puddling butterfly is closely watched, one can often see sudden, regular ejections, even squirts, of fluid from the end of the abdomen. These regular ejections of fluid can sometimes be powerful enough to reach several body lengths away from the butterfly!

A close-up of the ejected fluids from the abdomen of a puddling Fivebar Swordtail

Are there any dangers associated with puddling?

Some species of butterflies observed during puddling appear to be in drunken stupor, especially those that have been puddling for a long time. The butterflies appear to be intoxicated on whatever they have been puddling on and become oblivious to their surroundings. This makes them very vulnerable to predatory or parasitic attacks.

Is puddling a social activity?

Probably not in the way we humans define a social activity. However, whenever damp patches of sand or soil is rich in animal excretions or fermenting organic matter, a lucky observer can often encounter a large congregation of various species of butterflies puddling at the same spot - often jostling with each other for the best positions to puddle. If disturbed, the entire congregation erupts in a cloud of colour as their liquid buffet party is interrupted, only to settle down very quickly again to puddle as soon as any perceived threat is over.

A single-species congregation of puddlers. These are Three Spot Grass Yellows

Butterflies of most of the families do puddle, although the Papilionidae, Pieridae, Nymphalidae and Lycaenidae are most often observed to puddle quite readily. Many species, usually powerful flyers which rarely offers an observer a chance to view or photograph them when they are in flight, allow a much better chance for close-quarters observation when they are puddling.

Text by Khew SK with contributions from Anthony Wong ; Photos by Khew SK, Terry Ong, Mark Wong & Anthony Wong

Life History of the Chocolate Royal

Life History of the Chocolate Royal (Remelana jangala travana)


Butterfly Biodata:
Genus: Remelana Moore, 1884
Species: jangala Horsfield, 1829
Subspecies: travana Hewitson, 1865
Wingspan of Adult Butterfly: 35mm
Caterpillar Host Plants: Eurya acuminata (Theaceae)



Physical Description of Adult Butterfly:
Above, the male is a deep lustrous purple, with broad brown to dark brown bordering on both wings; the female is paler, and differs from the male also in that the basal portions of spaces 2 and 3 on the forewing are entirely purple. The under surface is dark brown, with conspicuous end-cell bars and a narrow, darker brown, post-discal line on both wings. The two prominent black tornal spots (the one in space 1a larger than the other in space 2) on the hindwing are crowned with brilliant metallic green scaling. Each hindwing has two white-tipped tails of equal length at veins 1b and 2.

Field Observations of Butterfly Behaviour:
This species is is rather rare in Singapore in recent times and sightings have been very infrequent over the last few years. Its occurrences appear to be restricted to a few areas in the the Central Catchment Nature Reserve.

Early Stages: The host plant, Eurya acuminata, has been described in the life history article for Narrow Spark. Thus far, it has been recorded as the local host plant for Narrow Spark, Chocolate Royal and Ambon Onyx. The caterpillars of Chocolate Royal feed on flower buds of this plant almost exclusively in all four instars. The only exception being in the final instar when leaves are also eaten if flower buds become scarce. It is possible that Chocolate Royal has at least one other host plant locally. Once, a female was observed to oviposit an egg on an Ixora bush. Unfortunately the young caterpillar did not survive past its first instar then.


Host plant : Eurya acuminata
On a flowering host plant in the nature reserve, the mother butterfly was observed to lay eggs singly on the underside of a number of leaves. Each egg is white, but with a strong greenish tinge when freshly laid; circular and covered with rather large hexagonal pits. The diameter is about 0.7mm.


An egg of the Chocolate Royal. Left: fresh egg; Right: empty egg shell.

1st instar caterpillar, newly hatched, length: 0.9mm

It takes 2.5 to 3 days for the egg to hatch. The young caterpillar nibbles away the upper portion of the egg shell to emerge. With a length of about 0.9mm, it is cylindrical with very long setae (hairs) and pale yellow in base color. Yellowish brown bands run dorso-laterally along its body. As it grows, the body assumes the more typical onisciform (woodlouse) shape.

1st instar caterpillar, length: 2mm

The 1st instar caterpillar finds its way to nearby flower buds and feeds by boring a hole into a flower bud and eating the interior flower parts. After 3 days of growth, and reaching a length of about 2.5mm, its moults to the next instar. The second instar caterpillar has shorter hairs covering its body. The prothoracic shield is roughly diamond in shape and black to dark brown in colour. The prominent dorso-lateral yellowish brown bands were present initially but they fade away during this instar which lasts about 4 days. As the bands fade away, the entire body assumes a light green coloration. The inconspicuous dorsal nectary organ can be seen with a close-up examination.


2nd instar caterpillar, early in thist stage, length: 2.9mm


2nd instar caterpillar, length:5mm
2nd instar caterpillar, late in this stage, length: 7mm

The 2nd instar caterpillar reaches a maximum length of about 7mm. The moult to the 3rd instar brings few obvious changes. The entire body is now covered in very shot fine hairs, and a dorsal nectary organ becomes prominently marked in reddisk brown with a central white patch. The 3rd instar takes 3 days to complete with the body length reaching about 12-13mm.

3rd instar caterpillar, early in this stage, length: 7.5mm


3rd instar caterpillar, late in this stage, length: 13mm

The 4th (and final) instar caterpillar has the same general appearance as the 3rd instar caterpillar. The prothoracic segment is covered with light brown spot while the prothoracic shield is yellowish brown in contrast. This takes about 4 days to complete with the body length reaching 21-22mm. As it grows, the caterpillar becomes more greenish in coloration. Nearing the end of this instar, the caterpillar ceases feeding, and its body shrinks in length. It finds its way to a leaf and takes up a position on the surface to become an immobile pre-pupa.


Close-up views on parts of a 4th instar caterpillar.
Left: the first two horacic segment showing the diamond-shaped prothoracic shield;
Right: dorsal nectary organ

4th instar caterpillar, early in this stage, length: 13mm

4th instar caterpillar, late in this stage, 22mm

The pre-pupatory caterpillar prepares for pupation by spinning a silk girdle and a silk pad to which it attaches itself via claspers. Pupation takes place after about 1 day of pre-pupal stage. The pupa has the typical lycaenid shape, though somewhat wider in proportion, and a length of about 11.5-12.5mm. It is entirely jade green in coloration.

Two views of a pre-pupa of the Chocolate Royal with the silk girdle featured

Two views of a fresh pupa of the Chocolate Royal

Six days later, the pupa becomes darkened in color signaling the imminent emergence of the adult. The uppersides of the forewings become visible through the now transparent pupal skin. The next day the adult butterfly emerges from the mature pupa.

Two views of a mature pupa of the Chocolate Royal

A Chocolate Royal resting on the underside of a leaf


Another Chocolate Royal found in the nature reserve

References:

• The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, 4th Edition, Malayan Nature Society.


• The Butterflies of Hong Kong, M. Bascombe, G. Johnston, F. Bascombe, Princeton University Pres 1999

Text and Photos by Horace Tan

Butterfly of the Month - November 2008

Butterfly of the Month - November 2008
The Dark Blue Jungle Glory (Thaumantis klugius lucipor)

The Dark Blue Jungle Glory (Thaumantis klugius lucipor) belongs to the sub-family Morphinae of the family Nymphalidae (Brush-Footed Butterflies). It is a forest-dependent species which is rather rare. Although it is observed from time to time in the nature reserves, very little is known of its life history in Singapore, nor the host plant its caterpillars feed upon. Often, only individuals are observed, displaying its typical sulking behaviour, feeding on rotting fruit and other organic plant waste amongst the forest litter.

A Dark Blue Jungle Glory is well camouflaged amongst the dead leaf litter on a forest trail in the nature reserves in Singapore

The Dark Blue Jungle Glory, which is the sole representative of the Thaumantis genus in Singapore, has its upper sides an iridescent deep ultramarine blue in colour. The beauty of this tropical butterfly is often compared to the Morphos butterflies of the Amazonian forests.

As it flaps its wings whilst feeding on overripe figs on the forest floor, this individual flashes a glimpse of its deep ultramarine blue upperside

The undersides of this species bears crypic patterns of various shades of brown. This camouflage renders the butterfly invisible when feeding on the forest floor amongst the dead leaves and other organic clutter. This explains why the butterfly is more often seen with a flash of deep blue as it makes its escape when disturbed. Most of the time, the butterfly has a sulking habit, preferring deep forested areas, and keeping close to the ground. It is usually seen early the morning or in the later hours of the day.

Although it has been written that the species is generally not attracted to rotting fruit bait, it has been observed to feed on overripe fig fruits in the nature reserves in Singapore. It may well be that the Dark Blue Jungle Glory does not share the same preference to the rotting fruits that other species of the Morphinae are attracted to.

The iridescent deep blue colour of the wings above extends from the base to the black submarginal border in the male and is much reduced in the female, where the blue is restricted to the outer half of the wings. The butterfly is relatively large, and attains a wingspan of up to 100mm in the females.

Text by Khew SK ; Photos by Khew SK, Anthony Wong, Mark Wong & Sunny Chir

Life History of the Ancyra Blue

Life History of the Ancyra Blue (Catopyrops ancyra)

Butterfly Biodata:
Genus: Catopyrops Toxopeus, 1929
Species: ancyra C. Felder, 1860
Wingspan of Adult Butterfly: 24mm
Caterpillar Host Plants: Pipturus argenteus (Urticaceae), Trema tomentosa (Ulmaceae)

Physical Description of Adult Butterfly:
Above, the male is dull blue with two black and subequal tornal spots on the hindwing. The female is fuscous black with basal parts shot with iridescent blue on the forewing and duller blue on the hindwing, and it has black submarginal spots on the hindwing with those spots in spaces 1b and 2 crowned with orange. Underneath, the dark interneural submarginal spots on the hindwing are separated from the rest of the wing by a narrow lunulate line, and the orange areas crowning the large dark subequal spots in spaces 1b and 2 are inwardly defined by a narrow dark line. There is a pair of white-tipped tails at the end of vein 2 on the hindwings.


One male Ancyra Blue pondering on his next move


Another Ancyra Blue ready to take flight under the hot midday sun.

Field Observations of Butterfly Behaviour:
This species was not recorded as being extant in Singapore by early researchers. It was first sighted in 2004 in Pulau Ubin by ButterflyCircle members, and since then, sightings have been made at multiple locations on the Singapore main island, including the Central Catchment Reserves, the Southern Ridges and western areas of Singapore, where the host plants are found to be thriving. Typically, a small colony of the adults can be found in one location. Encounters with the adults usually take place in brightly-lit condition with them engaging in activities such as sunbathing, dog-fighting or making oviposition visits to the host plants.

Early Stages:

Outside Singapore, recorded host plants are found in the plant families Urticaceae, Fabaceae (Caesalpiniaceae) and Euphorbiaceae. The local host plants recorded todate are Pipturus argenteus (Urticaceae) and Trema tomentosa (Ulmaceae), the latter being a new addition attributed to Sunny Chir, an active ButterflyCircle member, who sighted a small colony with females ovipositing on the host plant in an urban park. According to Prof. Konrad Fiedler, this is the first confirmed record of Trema-feeding in C. ancyra.

Trema tomentosa is a shrub or small tree with ovate or ovate-elliptic leaves of length 8-15cm. The acuminate leaves are toothed and three-nerved from the base with undersides covered in tiny silvery hairs. Though not common locally, this plant can be found in various waste grounds such as the abandoned farmland in the west, the Central Catchment Nature Reserve and the Southern Ridges. The early stages of the Ancyra Blue mainly feed on the flower buds. However, when flower buds are in short supply, the caterpillars have been observed to munch on young leaf shoots. Furthermore, the final instar caterpillar has no trouble dealing with the tougher lamina of mature leaves.


The host plant, Trema tomentosa.


Clusters of flower buds of Trema tomentosa.

Eggs of Ancyra Blue are laid singly on the underside of a leaf. The egg is small (about 0.4-0.5mm in diameter) and light green in colour, circular with a slightly depressed micropylar area and a reticulated pattern of intersecting ridges.


Left: an egg of the Ancyra Blue, diameter: 0.4mm-0.5mm. Right: ready for hatching.

An animated hatching sequence of an Ancyra Blue caterpillar.
The actual process lasts about 2 minutes.

Two to three days later, hatching takes place after the young caterpillar has nibbled away sufficiently large upper portion of the egg shell to emerge. Measured at a length of about 0.8mm-0.9mm, its pale yellow body is cylindrical in shape, sporting long setae (hairs) and a dark brown head capsule.


1st instar caterpillar feeding on a flower bud, length: 1.2mm

The 1st instar caterpillar finds its way from the leaf underside to a nearby cluster of flower buds to start its feeding frenzy. As it feeds and grows, it gradually takes on a yellowish to light greenish coloration.

Another feeding 1st instar caterpillar with frass pellets nearby, late in this stage, length: 1.8mm

After about 3 days of growth and reaching a length of about 2mm, the caterpillar moults to the next instar. The 2nd instar caterpillar has numerous short setae covering its body which has taken on a woodlouse shape. The body is yellowish green overall with the head capsule now in pale yellow.

Two views of a 2nd instar caterpillar, early in this stage, length: 2.5mm

The 2nd instar caterpillar reaches a length of about 4mm, and after 2-3 days in this stage, it moults again. The 3rd instar caterpillar has proportionately shorter but still numerous body hairs, a green body color and a wider body compared to the earlier instars. This instar takes 3 days to complete with the body length reaching about 5.5mm.


Two views of a 3rd instar caterpillar, late in this stage, length: 5.5mm

The 4th (and final) instar caterpillar is more striking in general appearance as there are a fair number of prominent white specks doting the green body surface, in addition to other smaller brown specks which are less discernible. Zooming in on these specks, one could see that they occur at the base of setae and have an "asterisk" structure. These structures could serve as mechano-receptors or might be glandular in nature. At a larger scale, the typical lycaenid nectary organs (both the dorsal nectary organ on the 7th abdominal segment and tentacular organs on the 8th abdominal segment) are now distinguishable.

Two views of 4th instar caterpillars in varying degrees of development,
Lengths as stated in the pictures.

The white "asterisk" structures at base of setae on the body of a 4th instar caterpillar.

A 4th instar caterpillar showing the presence of the dorsal nectary organ and tentacular organs.

After 5 days of growth and reaching a maximum length of around 12mm in the final instar, the body of the caterpillar gradually shrinks. Soon the caterpillar seeks out a spot on the surface of an adjacent leaf to station itself. At this site, it enters the pre-pupatory phase where it readies itself for pupation by spinning a silk girdle and a silk pad. The caterpillar secures itself to the silk pad via claspers on its posterior end.


Two views of a pre-pupa of the Ancyra Blue, showing a finished silk girdle
securing the pre-pupa to the leaf surface.

An animated pupation sequence for an Ancyra Blue caterpillar.

Pupation takes place after 1 to 1.5 days of the pre-pupal stage. The pupa has the typical shape for most Lycaenid species. It is greenish in color with a few brown and black specks scattered on the surface. The pupa has a length of about 7-8mm.


Two views of a fresh pupa of the Ancyra Blue.

5 to 5.5 days later, the pupa becomes darkened in color signaling the imminent emergence of the adult. The next day the adult butterfly emerges from the mature pupa in the morning hours. Out of a group of 8 adults successfully bred in the last month, 3 were male and the rest female.


Mature pupae: male (top), female (bottom). Note the difference in the wing pad area as
the forewing uppersides are different for the two sexes.


A short eclosion sequence of one male Ancyra Blue


A newly eclosed Ancyra Blue resting next to its pupal case.


Another newly eclosed Ancyra Blue

Acknowledgments:

I would like to express my gratitude to Prof. Konrad Fiedler for his generous advice on host plant information for the Catopyrops spp. and the functions of the "asterisk" structures found on the body surface of the mature larvae.

References:

• The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, 4th Edition, Malayan Nature Society.
• Butterflies of Borneo, Vol.2, No.1 Lycaenidae, Y. Seki, Y Takanami & K. Otsuka, Tobishima Corporation 1991.

Text and photos by Horace Tan