14 January 2012

Butterfly Courtship Rituals

Butterfly Courtship Rituals
Success strategies in finding the right mate



A large part of a butterfly's lifespan is spent directly or indirectly to achieve a particular objective - that of procreation. A newly eclosed butterfly spends most of its day feeding to build up its reserves and energy so that it can survive and search for a mate and to quickly ensure the continued survival of the species before it dies or succumbs to predation.



For a butterfly to be able to fulfil its objective of mating and producing viable eggs for the next generation, the males and females have to first locate each other. Males have to correctly identify a female of his own species by visual cues. A variety of characteristics, from colour, pattern, size, flight and other behavioural attributes that may not be apparent to the human eye, are used to assess the butterflies that they encounter.



Locating a Mate
So where does a fertile male go to look for potential females to mate with? Males generally have two main strategies. The first is perching or hill-topping. Basically, this method requires the male to search out a high vantage point from which he can have a clear view of an area. The moment an intruder or any movement is detected, the male will dart down and check out the newcomer into his territory.



If the intruder is a male, or a butterfly of another species, the male dives into the path of the newcomer and "attacks" it. An aerial dogfight ensues, until the intruder leaves the incumbent's territory. The resident male then returns to one of several favourite perches, and then continues to monitor his domain for new activity.



Hilltops or tall trees with the surrounding vegetation lower are ideal places for the 'perching' strategy. Hence such places function as a "singles' club" for both males and females of a species to find each other.



The second strategy employed by males of butterflies is what is referred to as "patrolling". The male flies continuously along tracks, forest edges and streams to look out for females. As it detects females by visual contact, it will check out anything that is in motion and even if it vaguely resembles a female of its own species.



Some males also patrol near its caterpillar host plants, with the objective of finding a newly-eclosed female of the same species. A well-known phenomenon amongst the Heliconiinae of South America is "pupal rape" where males of a particular species tear through the pupal shell of a female of the same species to mate with her even before she ecloses!



When a male locates a female of his own species, it will switch to courtship mode and track the female persistently. He then begins the rituals of courtship. For the male, finding the prospective mate is just the beginning. There are still no guarantees that the female will considering him worthy of fertilising her offspring.



The female being courted will assess and choose the male depending on age, general appearance and health and potential ability to provide viable spermatophore for her reproductive success. Females of many species tend to mate only once so her choice is crucial. Indeed, many ButterflyCircle members have observed an ovipositing female, collected the eggs for breeding, only to find that the eggs were infertile and did not hatch at all! Could that be due to her eggs not being successfully fertilised by the male?







Courtship for the male commences with trying to get the female's attention, via a series of signals, ranging from flapping around the female to gain her attention (she may be going around her own business of feeding for example), sending chemical signals to her, in the form of pheromones. A typical movement is where the male flaps in position, usually upwind from the female, and release his pheromones so that she gets a clear signal of his intention to mate.

In the male, pheromones are usually found in the specialised scales on various parts of the wing - called androconial cells, specialised organs like the hair pencils in Danaianae species, or hair brushes on the undersides of the wings brushing against a special scent organs on the wings.



The female which picks up the signals then decides on her next course of action. She may continue with her own business and fly away, totally ignore the male. Of course, the male usually being persistent in his courtship attempts (like some men do in our human world!), will take the initial rejection in his stride and continue to re-send the physical and chemical signals to the female repeatedly.



Once again, the female may fly off, indicating that she is not interested. This can go on for several attempts before the female responds, or continues to reject the male until he gets the message and gives up.



In some cases, the males are more aggressive and flies in such a way as to obstruct the female's ability to fly off, or corners her. If the female is still not interested, she will usually stop with her wings opened, and adopt her "rejection posture".



This is usually done with her wings opened either fully flat or half-opened, but with her abdomen thrust upwards and elevated such that it would be all but impossible for the male to clasp the end of her abdomen. A persistent male may still flap violently above the female and try to push her abdomen down, but he will usually fail, and fly off after a few attempts.



If the female responds positively and decides to accept her suitor's advances, she will close her wings and remain still. This signals the male to now approach her. He will land beside her, flick his wings a few times, moves beside her facing the same direction, and curls his abdomen towards hers and grasps hers using his claspers. The receptive female will extend and offer her abdomen for coupling.







Once engaged, the coupled butterflies will face away from each other in the usual mating pose. Usually, if disturbed, the larger wingspan of the two (usually the female) will do the flying, whilst the other partner remains still. It would be unimaginable for both to fly at the same time, and in opposite directions!







In the field, we have seen impatient male butterflies hovering around pupae from which females are about to eclose, and immediately mate with them when the female breaks out of her pupa. Copulation occurs even before the female is able to dry her wings properly. In one case, the male was so insistent on 'doing it' that he damaged the wings of the female, leaving her crippled and unable to fly!



In other cases, we have also observed males of various species coupling in flight in an instant, dispensing with the long courtship rituals and avoid suffering the fate of rejection. One example that I have personally witnessed, was when a male, perched on a high vantage position, swooped down on a passing female, engaged her in mid-air, and both fell downwards, coupled into the bushes and stayed mated for quite some time! Now that, I would call precision engagement!



And so we see how in the world of butterflies, males have to woo their mates and work hard before they can successfully mate with a female of their choice. Males will often have to suffer the 'indignity' of rejection as there are no guarantees that the female that they have targeted, would accept them, no matter how hard they try. But like all things in life, success always comes to those who are persistent in the face of failure and work hard for it.



Text by Khew SK : Photos by Sunny Chir, Federick Ho, Khew SK,Terry Ong & Anthony Wong

References :
  • Handbook for Butterfly Watchers : Robert M Pyle, 1984, 1992; Houghton Mifflin Company, New York
  • A World for Butterflies - Their Lives, Behavior and Future : Dr Phil Schappert, 2000; Firefly Books
  • Butterflies : Dick Vane-Wright, 2003; The Natural History Museum, London

07 January 2012

Life History of the Malay Dartlet

Life History of the Malay Dartlet (Oriens paragola)


Butterfly Biodata:
Genus: Oriens Evans, 1932
Species: paragola 
de Nicéille, 1896
Wingspan of Adult Butterfly: 22-27mm
Caterpillar Local Host Plant: Ottochloa nodosa (Poaceae).





Physical Description of Adult Butterfly:

Above, both sexes are dark chocolate brown with amber-coloured post-discal bands on both fore- and hindwings, and amber-coloured cilia on the hindwing termen. The forewing band stretches from the dorssm to almost touching the costa, and is deeply excavate at vein 5. The lower cell spot fuses with this band at space 2. The hindwing post-discal band is shorter, stopping well short of the costa.   Underneath, the wings are dark brown  with  post-discal bands mirroring those on the uppersides. These bands are sharply defined and not edged by black spots (as is the case of the Oriens gola, the Common Dartlet).




Field Observations of Butterfly Behaviour:
This species was not listed by early researchers as extant in Singapore. It was  discovered about one year ago and added to the Singapore checklist as species #297. According to C&P4, the Malay Dartlet frequents  lowland forests. However, adult sightings in Singapore  have so far been very infrequent and only confined to one grassland adjacent  to a forested area. The species has a preference for perching in semi-shady areas, and visiting flowers in the grassland for nectar. Given that the larval host plant is a common grass species, there is a possibility that Malay Dartlet exists elsewhere in Singapore. Even if this is not so now, this species  has the potential of spreading beyond the currently known location.



Early Stages:
The host plant, Ottochloa nodosa, is a common grass species which can be found in many locations in Singapore and is utilized by several other common skipper spp. (Common Dartlet, Chestbut Bob, and one Caltoris sp.) as larval host plant. The caterpillars of the Malay Dartlet feed on leaves of this plant, and live in shelters made by joining edges of a grass  blade together with silk threads.


Host plant: Ottochloa nodosa.

A female Malay Dartlet resting between oviposition runs.

A female Malay Dartlet laying an egg on the underside of a grass blade.

The eggs are laid singly on the underside of a leaf of the host plant. Between each oviposition run, during which one or two eggs are laid on separate leaves, the mother butterfly has the habit of  perching on a sunny spot  to re-energise  This could go on for a few rounds before the mother decides to retire altogether.  Each tall hemispherical egg is milky white and is finely recticulated.  The base diameter is about 0.75mm.

 
An egg of the Malay Dartlet.

A maturing egg of the Malay Dartlet.

A mature egg with the young caterpillar already nibbled away the polar part of the egg shell.

It takes about 5 days for the egg to hatch. The young caterpillar eats just enough of the shell to emerge, and will devour the rest of the egg shell after emergence. The newly hatched has a length of about 1.7-1.8mm. Its has a cylindrical body shape, very short dorso-lateral and sub-spiracular setae and a tuff of long setae at the posterior end. The body is creamy white with a prominent black collar on the dorsum of the prothorax. The head capsule is entirely black.


A newly hatched 1st instar caterpillar in the process of nibbling away its egg shell.

Two views of a newly hatched 1st instar caterpillar, soon after devouring the egg shell.

The body color changes to pale yellowish green after a few feeding sessions on the leaf. The newly hatched constructs its leaf shelter near the tip of a grass blade soon after it is done with the egg shell. Between feedings on the nearby leaf lamina, the caterpillar retreats to its shelter for rest and security. By the time the caterpillar lies dormant in its shelter for the moult to the 2nd instar, its length has reached about 4-4.5mm. The 1st instar takes about 4-4.5 days to complete.

Two views of a 1st instar caterpillar, early in this stage, length: 2.4mm.

Two views of a 1st instar caterpillar, late in this stage, length: 4mm.

The 2nd instar caterpillar is again whitish, but appears to be yellowish green with its "stomach" contents showing through the translucent cuticle.  The anal plate still carries a few setae but these are not as long and prominent as in the 1st instar. Although the prothoracic collar is still present, it appears to lose lighter in its black coloration. The head capsule is still black in colour. This instar lasts about 3-4 days with the body length reaching about 6.5-7mm.

Two views of a 2nd instar caterpillar, early in this stage, length: 4.2mm.

Two views of a 2nd instar caterpillar, length: 5.2mm.

The 3nd instar caterpillar resembles the 2nd instar caterpillar closely in body markings except for the diminishing black prothoracic collar.  Two faint whitish dorso-lateral bands, first appears in the 2nd instar but less obvious, are now discernible. This instar lasts about 2-3 days with the body length reaching about 8.5-9mm.

A newly moulted 3rd instar caterpillar, the head capsule is yet to turn black.

Two views of a 3rd instar caterpillar,  length: 7.2mm.

 
Two views of a 3nd instar caterpillar, later in this stage. length: 8.2mm.

The 4th instar caterpillar resembles the 3rd instar caterpillar closely but with the black prothoracic collar almost completely  diminished. As in the earlier instars, the head capsule is completely black and there is no markings on the anal plate.  This penultimate instar lasts 3-4 days with the body length reaching up to about 15.5-16mm.

Two views of a 4th instar caterpillar, early in this stage, length: 12mm.

 
Two views of a 4th instar caterpillar, late in this stage, length: 15.8mm.

The 5th (and final) instar caterpillar loses the black coloration of its head capsule seen in the earlier instars. The head is  now pale neutral yellow to neutral green. The body colour has a strong tone of yellowish white, with two broad and whitish dorso-lateral bands and two narrow pale yellowish lateral bands.  This instar takes about 6-8 days to complete with the body length reaching up to about 24-25mm.

Two views of a  newly moulted 5th instar caterpillar.

 
Two views of a 5th instar caterpillar, late in this intar, length: 24mm.

Towards the end of the 5th instar, the the body of the caterpillar shortens and its base coloration changes to a translucent pale green. Tyipcally it picks the underside of a grass glade for its pupation site. It becomes dormant in this shallow leaf shelter and enters the prepupatory phase which lasts for one day.  During this time period the caterpillar spins a silk pad with a transverse band for the grapers in its anal prolegs to attach to, and a silk girdle across the dorsum of the 2nd-3rd abdominal segments.

Two views of a pre-pupa of the Malay Dartlet.

The pupa  secures itself to the substrate with a cremastral attachment to the silk pad and the previously constructed silk girdle. It is yellowish green throughout and has  dorsal, dorso-lateral and lateral whitish to yellowish bands. There is a long and pointed snout, and the abdomen is relatively long.  Length of pupae: 18.5-21.5mm.

A Malay Dartlet caterpillar molts to its pupal stage.

Two views of a pupa of the Malay Dartlet.

On the last day of the pupal period, the pupa becomes mostly dark brown. Yellowish orange  markings against a dark brown background are now visible in the wing cases. Finally after about 7-8 days of pupal phase, eclosion takes place with the adult emerging from the pupal case.

Two views of a mature pupa of the Malay Dartlet.

A Malay Dartlet emerges from its pupal stage.

A newly eclosed Malay Dartlet.

References:
  • [C&P4] The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, 4th Edition, Malayan Nature Society.
  • Butterflies of Thailand, Pisuth Ek-Amnuay, 1st Edition, 2006
Text by Horace Tan, Photos by Koh Cher Hern, Loke PF, Chng CK, Sunny Chir, Khew SK and Horace Tan