27 March 2010

Life History of the Pitcher Blue

Life History of the Pitcher Blue (Virachola kessuma deliochus)


Butterfly Biodata:
Genus: Virahola Moore, 1881
Species: kessuma Horsfield, 1829
Subspecies: deliochus Hewitson, 1874
Wingspan of Adult Butterfly: 26mm
Caterpillar Local Host Plants: Nepenthes gracilis (Nepenthaceae, common name: Slender Pitcher Plant), N. rafflesiana (Nepenthaceae, common name: Raffles' Pitcher Plant)


An adult Pitcher Blue perching on a wild flower in an open field in Southern Ridges.

A male Pitcher Blue on a perch.

A female Pitcher Blue on a leaf perch.

Physical Description of Adult Butterfly:
Above, the male has its forewing mostly black with a small basal patch of shining blue at the dorsum, its hindwing mostly shining blue with a black costal border and a pale brown dorsum. The female is duel pale blue with broad brown borders on both wings. Beneath, the wings are pale brown and marked with white striae in similar arrangement as for the Four-line blues (pavana group of Nacaduba spp.) but without basal striae in the hindwing. Each hindwing has a small tornal lobe, a prominent subtornal orange-crowned black spot in space 2, blue and green metallic scales in space 1b and a white-tipped filamentous tail at the end of vein 2.

The upperside of a female Pitcher Blue.

A female Pitcher Blue checking out a flower of the Common Asystasia.

Field Observations of Butterfly Behaviour:
The Pitcher Blue is rarely seen in Singapore due to the loss of habitat for its larval host plants to various industrial, commercial and housing development projects in the past decades. In recent times, the adults have been sighted rather infrequently in locations where small pockets of pitcher plants remain. Such sites include parts of the nature reserves, Southern Ridges and other scattered wastelands. The adults are fast fliers. They usually stay high up in the trees and only make occasional visits to flowers in the vicinity. With luck and perfect timings, a keen observer can find females making oviposition visits to seed pods of the pitcher plants.

A female Pitcher Blue sighted making a brief stop-over in a wasteland.

A female Pitcher Blue on a flower of Buah Cheri (Muntingia calabura).

A female Pitcher Blue perching on a flower.

Early Stages:
Of the two Nepenthes spp. recorded as local larval hosts, N. gracilis, is found to be more commonly utilized by the Pitcher Blue as it occurs in much greater numbers than N. rafflesiana in their common habitats. N. gracilis is shrubby with alternate leaves and climbs with twisting leaf-tendrils. A fully formed leaf composed of a petiole, a lanceolate or linear leaf blade, a tendril and a tubular pitcher which has a lightly swollen and shortly incurved base. The pitchers are typically green with occasional reddish tinge. The flowers occurs in racemes and the fruits in capsule form (seed pods).

Host plant: Nepenthes gracilis. Leaf and pitchers (left). A fructescence.

Host plant: Nepenthes rafflesiana. Leaf and pitchers (left). A fructescence.

The caterpillars of the Pitcher Blue feed on the maturing seeds within a seed pod after gaining entry through a hole bored on the side of the pod. When their body size allows, especially during first three instars, they will stay within the confine of the pod and devour all its contents. The act of boring into a pod and eating its contents is repeated many times during the larval stage. Moulting invariably takes place within a seed pod.

Several eggs of the Pitcher Blue found on one single fructescence of N. gracilis. Look for a white dot on the tip of a seed pod.

Eggs of Pitcher Blue are typically laid singly, and rarely in pairs or triplets, on the stigma remnant of a seed pod. At times, eggs are laid at the lower portion of the seed pod near the calyx. It is not uncommon to see multiple egg-bearing seed pods on one single fructescence.

Close-up on egg-bearing seed pods of N. gracilis (Left) and N. rafflesiana (Right).

Empty egg shells on the stigma remnant of seed pods. Up to 3 eggs have been found on a single pod.

The small egg is dome-shaped (about 0.8-0.9mm in diameter) with a depressed micropylar. The surface is covered with a reticulated pattern of intersecting raised ridges. When freshly laid, the egg is whitish with a yellowish green undertone.

Two views of an egg of the Pitcher Blue. Diameter: 0.8mm.

Each egg takes 3.5-4 days to hatch. The young caterpillar emerges after nibbling away sufficiently large portion of the egg shell. Measured at a length of about 1.1mm to 1.2mm, its pale yellowish brown body is cylindrical in shape, sporting long fine setae (particularly long for those anterior and posterior ones), a dark brown to black head capsule and a large and oval shaped prothoracic shield. A dark brown anal plate is also featured with a prominent black spot.

Two views of a newly hatched caterpillar boring into a seed pod, length: 1.1-1.2mm.

The newly hatched makes its way to the side of the egg-bearing seed pod and starts to bore into it. It stays within the seed pod and feeds on the developing seeds. After about 2 days of growth, it reaches about 3.5mm in length. It then lies dormant within the seed pod for its moult to the 2nd instar. One seed pod is usually sufficient to last the entire 1st instar.

Two views of a 1st instar caterpillar, day 2 in this stage, length: 2mm.

Two views of a 1st instar caterpillar in a seed pod with content nearly all eaten.
The pod was peeled opened to reveal the caterpillar. Length: 3.2mm.




Life History of the Pitcher Blue. Part 1 of 3: Egg to late 1st instar.

Covered with numerous dark setae, the body of the 2nd instar caterpillar is brown to reddish brown in color with moderately longer setae running laterally. The head capsule is yellowish brown in colour. The dark brown to black prothoracic shield is rather prominent, resembling a triangle with a long base and short height. The body is still more or less cylindrical with larger anterior segments. The growth in this stage brings the caterpillar to a length of about 5.5-6.0mm, and after about 2 days in this stage, it moults again.

Two views of a 2nd instar caterpillar boring into another seed pod, early in this stage, length: 4mm.

A 2nd instar caterpillar in a seed pod, late in this stage, length: 6mm.

The 3rd instar caterpillar has numerous short and fine body setae, again with longer setae laterally. The head capsule is yellowish brown. The large and prominent prothoracic shield is more angled and comes with a little tooth at its base. The base body color is reddish brown but with lighter coloration on the dorsum of middle body segments. The 3rd instar takes about 3 days to complete with the body length reaching about 9mm before the next moult.

Frontal views showing the head and prothoracic shield of caterpillars in
2nd instar (left) and 3rd instar (right).


Two views of a 3rd instar caterpillar boring into a seed pod. early in this stage, length: 5mm.

3rd instar caterpillars in cut-open seed pods, lengths: 7mm (top) and 9mm (bottom).

As in the 3rd instar, the 4th (and final) instar caterpillar has numerous body setae and long lateral setae. There are also two rows of long setae occuring dorso-laterally. The body color is reddish brown to wine red.

Two views of a 4th instar caterpillar boring into a seed pod, early in this stage, length: 8mm.

Two views of a 4th instar caterpillar boring into a seed pod, midway through this stage, length: 12mm.

One prominent change from the 3rd instar is seen in the prothoracic shield which has now become pale yellow in base colour with small black spots lying within and large black spot in its four corners. The dorsal nectary organ is rather prominent. It is functional as ants have been observed to attend to it for caterpillars found in the field.

The prothoracic shield (left) and the dorsal nectary organ (right) of a 4th instar caterpillar.

A 4th instar caterpillar attended to by an ant in its natural habitat.

On the last feeding day in this instar, the first two thoracic segments and 3rd to 5th abdominal segments becomes lighter in coloration (shades of yellow, orange and even beige), giving the caterpillar a distinct banded appearance.

4th instar caterpillars, late in this stage. Top: eating through a hole, length: 14mm.
Bottom: wandering for a pupation site, length: 15mm.




Life History of the Pitcher Blue. Part 2 of 3: 2nd to 4th instar.


After 4-5 days of growth and reaching a maximum length of around 15-16mm in the final instar, the body of the caterpillar gradually shrinks. The caterpillar ceases eating and wanders around for a pupation site. All bred specimens chose to enter their pre-pupatory phase on a spot of the leaf surface within a gape in a pile of leaf litter. At the chosen site, the caterpillar readies itself for pupation by spinning a silk girdle and a silk pad. In the wild, some caterpillars even choose to pupate within a pitcher.

Two views of an immobile pre-pupatory larva of the Pitcher Blue,
The silk girdle can be easily seen.


Pupation takes place after one day of the pre-pupal stage. The lightly hairy pupa has the typical lycaenid shape, light brown in base colour with variable number of black to dark brown patches on the pupal surface. The pupa of Pitcher Blue has silk girdles and is equipped with cremastral hooks to secure it to a silk pad on the substrate. The pupa has a length of about 9-10mm.

Two views of a pupa of the Pitcher Blue, length: 9.5mm.

Eight days later, the pupa becomes darkened in color signaling the imminent emergence of the adult. Patches of green and blue can be seen in the wing pads through the now transparent pupa skin. The next day the adult butterfly emerges from the mature pupa.

Two views of a mature pupa of a male Pitcher Blue.



Life History of the Pitcher Blue. Part 3 of 3: Pupation and Eclosion.

A newly eclosed male Pitcher Blue drying its wings on a young fern frond.

References:
  • [C&P5] The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, G. and N.  van der Poorten (Eds.), 5th Edition, Malayan Nature Society, 2020.
  • Butterflies of Thailand, Pisuth Ek-Amnuay, 2nd Edition, 2012.
  • A guide to the carnivorous plants of Singapore, edited by Hugh T. W. Tan, Singapore Science Centre, 1997.
Text by Horace Tan, Photos by Sunny Chir and Horace Tan.

21 March 2010

Mergers, Partnerships & Betrayals

Mergers, Partnerships and Betrayals
The Miletinae - The Harvesters


A multi-tasking pair of Bigg's Brownie (Miletus biggsii biggsii) - mating & feeding amongst their protector ants

In the real corporate world, the title of this blog would probably not elicit much interest and excitement, other than offering a good read about yet another unfortunate company being swallowed up, or employees being thrust into a chasm of anxiety about their future.


A Bigg's Brownie (Miletus biggsii biggsii) feeding off what appears to be a "herd" of mealy bugs that are tended by their protector ants

In the world of Lepidoptera, however, the partnership between butterfly and ants is already a well-studied topic by entomologists. Myrmecophily, or the positive interspecies association between ants and butterflies (or rather, their caterpillars) has been widely studied. This symbiotic association occurs with some Lycaenidae caterpillars evolving specialised organs that exude honeydew in return for protection by the ants. These caterpillars then go about their daily business of feeding on the leaves of their host plants, whilst enjoying 'armed' guards' protection from predators.



However, there exists a subfamily within the Lycaenidae family - the Miletinae, that is unique in the sense that their caterpillars are predatory or "carnivorous" in the butterfly world. Whilst it is generally well-known that most caterpillars are "herbivorous" and generally feed on leaves, buds and flowers of plants, the Miletinae (or commonly referred to as the Harvesters) species' caterpillars feed on a variety of Homoptera like aphids, coccids, mealy bugs and one species even feeds on ants' larvae!



In Singapore, the Miletinae is represented by five genera - Allotinus, Logania, Spalgis, Miletus and Liphyra. Other than Liphyra, the other genera have caterpillars that feed mainly on a variety of Homoptera.



The aphids, coccids and mealy bugs, which form the principal diet of the caterpillars of the four genera mentioned, tend to feed on young shoots of various plants. These insects have adapted piercing mouth parts that pierce and suck the sap of plants - usually soft-stemmed ones. The Homoptera must suck large amounts of sap from the plant to get their nutrients. Much of the sap is a sugary fluid that cannot be digested. This extra liquid is passed through their body and excreted as "honeydew." Ants and other insects "farm" the Homoptera and collect honeydew for their own food. In return for their food, ants will protect the Homoptera that provide their colony with a food source.



In the field, observations have been made of the Miletinae adults feeding off the secretions of the aphids, mealy bugs and coccids. The butterflies do this brazenly, without any fear of being attacked, in the presence of the protective ants that are also 'milking' the aphids or mealy bugs of their honeydew.

Walking gingerly in a minefield of ants!

Besides feeding off the honeydew, the Miletinae also lay their eggs where the aphids and mealy bugs are present, and the caterpillars of these species feed on the aphids and mealy bug community.

A Pale Mottle (Logania marmorata damis) amidst mealy bugs and their "farmers"

It is interesting to note that the ants neither extract honeydew from, nor tend to the caterpillars in this case. They go about their business of tending to the aphids or mealy bugs, but leave the caterpillars alone - despite the caterpillars devouring their "herd"! Why then, don't the normally ferocious ants attack and kill the predatory caterpillars?


Miletinae caterpillars amongst their "sheep" which provides food as they grow

Researchers have postulated that the caterpillars (and even adult butterflies) have some kind of chemo-mimicry that render themselves 'invisible' to the ants. In other words, as far as the ants are concerned, the Miletinae adults and caterpillars are perceived as just another ant in the colony.


Hello? Are you my brother? An ant checks out a Lesser Darkie (Allotinus unicolor unicolor).

Therefore, although the caterpillars and adult butterflies of the Miletinae derive 'protection' from the ants, they give nothing back in return. On top of that, the adult butterflies partake of the honeydew that are secreted by the aphids and mealy bugs, and their caterpillars feed on the ants' source of food. Hence the "betrayal" part of the whole situation, where the ants have been hoodwinked into a win-lose relationship with the Miletinae.

A Lesser Darkie feeds on honeydew produced by some aphids that are tended by their farmer ants

An even more extreme situation occurs with the Moth Butterfly (Liphyra brassolis abbreviata). The caterpillar of this species feeds on the larvae of the Weaver Ant (Oecophylla smaragdina). It is interesting to note that the caterpillar feeds and grows inside the nest of the ants, and has evolved a clever way to escape the wrath of the ants when it ecloses. Our earlier blog article describes the Moth Butterfly.

A newly-eclosed Moth Butterfly (Liphyra brassolis abbreviata) whose caterpillar feeds on ant grub

This unique characteristic of the sub-family Miletinae in the butterfly world is amazing, and runs contrary to the conventional belief that all caterpillars are herbivorous and feed only on plant material. The adult butterflies of the five genera of the Miletinae found in Singapore are therefore very widespread, having little dependence on the location of host plants, as their "host" food is actually other living insects! Hence wherever there is a colony of aphids, coccids or mealy bugs, the adult butterflies of this sub-family will seek them out, for food for themselves as well as their young.

An Apefly (Spalgis epius epius) one of the Miletenae whose caterpillars also feed on Homoptera

So do look out for these butterflies and special behaviour when you are out in the field!


Text by Khew SK ; Photos by Sunny Chir, James Foong, Federick Ho, Khew SK, Loke PF, Bobby Mun, Horace Tan & Tan Tze Siong

13 March 2010

Life History of the Mottled Emigrant

Life History of the Mottled Emigrant (Catopsilia pyranthe pyranthe)



Butterfly Biodata:
Genus: Catopsilia
Hübner, 1819
Species: pyranthe Linnaeus, 1, 1758
Subspecies:
pyranthe Linnaeus, 1, 1758
Wingspan of Adult Butterfly: 65mm
Caterpillar Local Host Plants:
Senna alata (Leguminosae, common name: Seven Golden Candlesticks)


A Mottled Emigrant perching on a grass blade.


Another Mottled Emigrant sighted on its perch in an urban park where its host plant is found.

Physical Description of Adult Butterfly:
Above, the wings are greenish white and the forewing has a black apical border and a black cell-end spot. This black border and the cell-end spot are respectively broader and larger in the female than in the male. Beneath, the wings are of a darker and greenish-ochreous hue and there are many rather faint and short transverse striae in reddish brown, giving it a "mottled" appearance.

A newly eclosed female Mottled Emigrant resting near its pupal case.


Another newly eclosed Mottled Emigrant.

Field Observations of Butterfly Behaviour: The Mottled Emigrant is described in C&P4 as being the commonest and most widespread species of Catopsilia in Malaya. However, the reverse is true in Singapore as it is the rarest of the three Catopsilia species locally. The fast flying adults are typically found at the fringes of nature reserves or in a few wastelands where its host plant, Senna alata, is growing. In addition, where this plant is cultivated, it is likely that Mottled Emigrant will make its appearance as well.

Early Stages:
The host plant, Senna alata, is a shrub up to 3m high with flowers growing in columns resembling fat candlesticks. The leaves are simple pinnate with large leaflets, each of which is 50-80mm long. The caterpillars of the Mottled Emigrant feed on the relatively young to middle-aged leaflets.



Host plant: Senna alata (Seven Golden Candlesticks).


A mating pair of the Mottled Emigrant.

The eggs of the Mottled Emigrant are laid singly on the upperside of a leaflet of the host plant. At times, more than one egg can be found on the same leaflet. The long spindle shaped egg is laid standing at one end with a length of about 1.7-1.8mm. It is white in color with vertical ridges and numerous indistinct and intermittent horizontal ridge lines. The micropylar sits at the tip of the standing egg.

Left: fresh egg. Right: mature egg showing the head at the hole near the top.
The egg takes about 2 days to hatch. The newly hatched has a length of about 2.7mm and has a pale white to yellow head capsule. It has a cylindrical and pale yellowish green body featuring dorsal, dorso-lateral and lateral rows of small tubercles running lengthwise. Each tubercle has a short seta emerging from the middle of it.


Two views of a 1st instar caterpillar, early in this stgae, length: 2.8mm.

After hatching, the young caterpillar eats the empty egg shell for its first meal, and then moves on to eat the leaf lamina. The growth is fast paced and the body length reaches about 4.5mm in this 1st instar which lasts just 1.0-1.5 days.

Two views of a late 1st Instar caterpillar hours before its moult, length: 4mm.

The 2nd instar caterpillar is yellowish green in color on both body segments and the head. The body is covered in rings (about six for each body segment) of numerous small black tubercles from which very short setae emerge. The head capsule also features numerous similar tubercles.
This instar lasts about 1-1.5 days with the body length reaching 8-9mm.


Two view of a 2nd instar caterpillar, length: 4.1mm

The 3rd instar caterpillar resembles the 2nd instar caterpillar closely except for the appearance of a faint whitish lateral lines. Towards the end of this instar, a faint yellow line also appears on the upper edge of this white line. This instar takes about 2-2.5 days to complete with body length reaching about 12-14mm.


3rd instar caterpillars. Top: 9mm. Bottom: 12mm.

The white-yellow stacked lateral line on the 4th instar caterpillar is much more prominent. Furthermore, the first or first 2 rows of dark tubercles above this lateral line are larger with a bluish shine, thus forming prominent outline to the white-yellow line. This penultimate instar lasts 2-2.5 days with body length reaching about 23-25mm.


Two views of a 4th instar caterpillar, early in this stage, length: 14mm.


Two views of a 4th instar caterpillar, late in thie stage, length: 24mm


A close-up view of the stacked white-yellow line and row/rows of larger black tubercles above.

The 5th and final instar caterpillar resembles the 4th instar caterpillar closely with no obvious changes. The 5th instar lasts for 2.5-3 days, and the body length reaches up to 41-45mm.


Two views of a 5th instar caterpillar, newly moulted, length: 26mm.

5th instar caterpillars. Top: 39mm. Bottom: 45mm.

On the last day of the 5th instar, the body of the caterpillar shortens and changes to a dull shade of pale green. It ceases feeding and comes to rest on the midrib on the underside of a leaflet or the stem/stalk on the host plant. Here the caterpillar spins a silk pad and a silk girdle to secure itself and then becomes immobile in its pre-pupatory pose.


A pre-pupatory larva of the Mottled Emigrant on the leaf underside.


Pupation takes place about 0.5 day later. The pupa secures itself with the same silk girdle as in the pre-pupal stage, but with cremaster replacing claspers in attaching the posterior end to the silk pad on the stem. The pupa has a pointed head, and its yellowish green body has a yellow lateral line on each side. There is a dorsal protrusion with a yellow ridge line on the thorax. The keel formed by the tapering wing case is much less pronounced than those seen in the Eurema spp.
Length of pupae: 28-29mm.


The pupation event of a Mottled Emigrant caterpillar.



Two views of the pupa of a Mottled Emigrant.


Two views of the mature pupa of a female Mottled Emigrant.

The now transparent wing pad shows the broad black border and cell spot.

After about 5 days of development, the pupal skin turns translucent as the development within the pupal case comes to an end. The yellow coloration and black borders on the forewing upperside are now discernible. The following day, the adult butterfly emerges from the pupal case.


The eclosion sequence of a male Mottled Emigrant.


A newly eclosed Mottled Emigrant drying its wings on its pupal case.


References:

  • The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, 4th Edition, Malayan Nature Society. [C&P4]
  • Butterflies of Thailand, Pisuth Ek-Amnuay, 1st Edition, 2006
Text by Horace Tan, Photos by Jason Ng, Benedict Tay, Bobby Mun, Anthony Wong and Horace Tan