Saturday, June 13, 2020

Life History of the Malay Staff Sergeant

Life History of the Malay Staff Sergeant (Athyma reta moorei)


Butterfly Biodata:
Genus: Athyma Westwood, 1850
Species: reta Moore, 1858
Subspecies: moorei Fruhstorfer, 1906
Wingspan of Adult Butterfly: 50-60mm
Caterpillar Local Host Plant: Glochidion zeylanicum (Phyllanthaceae).


A female Malay Staff Sergeant enjoying a good meal on the ripened fruits of the Singapore Rhododendron.

A male Malay Staff Sergeant puddling on wet ground.

A male Malay Staff Sergeant with closed wings giving a view of its underside.

Physical Description of Adult Butterfly:
Forewing cell is closed. Malay Staff Sergeant is sexually dimorphic with a black-and-white (white markings on black background) male and a black-and-orange female. On the forewing upperside, the cell streak is twice-constricted, and the arrowhead-shaped marking beyond the cell streak is well apart in the male. The cell streaks of the left and right forewings are connected via a dorsal band on the thorax. The post-discal spots in spaces 4, 5 and 6 are well separated in the male but confluent in the female. There is no post-discal spot in space 3 in the male but there is a small one in the female. The post-discal spot in space 2 is oval and well separated from that in space 1b in the male, but is oblong and adjoining the spot in space 1b in the female. On the hindwing upperside, there are sub-discal band, post-discal band and marginal band which run from the costa to the dorsum. On the underside, the spots and streaks are arranged as on the upperside but with the ground colour yellowish to dark brown.


Another male Malay Staff Sergeant puddling on wet ground.

A female Malay Staff Sergeant bathing in the sun on the leaves of its host plant Glochidion zeylanicum.

A female Malay Staff Sergeant with closed wings giving a view of its underside.

Field Observations of Butterfly Behaviour:
The Malay Staff Sergeant is a uncommon butterfly in Singapore. The strong and fast-flying adults can be sighted at the fringe of nature reserves, and several forested wastelands across the island. The adults visit flowers and ripened fruits for the sugary contents. The male has been sighted puddling on wet grounds.

A video record of the life history of the Malay Staff Sergeant.

A sun-bathing male Malay Staff Sergeant sighted in the western region of Singapore.

Another male Malay Staff Sergeant sighted in the western region of Singapore, resting on a flower of Bidens alba.

A newly eclosed female Malay Staff Sergeant.

Early Stages:
Only one host plant, Glochidion zeylanicum, has been recorded as larval hosts for the Malay Staff Sergeant in Singapore so far. It is likely that other Glochidion species could serve as larval hosts, as in the case of the closely related Athyma nefte (Colour Sergeant).

Local host plant: Glochidion zeylanicum.

The caterpillars of the Malay Staff Sergeant adopt the same feeding and frass management routines as described for other Athyma species in earlier blog articles. The caterpillars feed on mature leaves of the host plant in all five instars of its development. On each leaf, they feed systematically from the leaf tip towards the petiole, leaving the midrib intact. From the 1st to the 4th instar, they build an extension of the exposed midrib using frass pellets held together by silk. Frass pellets are also sewn together beneath the midrib at the base of the extension. The caterpillars rest on either the frass extension or the exposed midrib above the bundle of frass pellets there. In the 5th (and final) instar, the caterpillars abandon these habits, resting on the leaf upperside and feeding irregularly from the leaf margin.

A mating pair of the Malay Staff Sergeant. Top: male; Bottom: female.

The eggs of the Malay Staff Sergeant are laid singly at the tip of a leaf of the host plant. Each dome-shaped egg is yellow in colour, with its surface marked with polygonal ridges which are tipped with fine, translucent spines at intersections of these ridges. Basal diameter is approximately 1.0mm.

A female Malay Staff Sergeant ovipositing at the tip of a leaf of Glochidion zeylanicum.

A time-lapse sequence of a female Malay Staff Sergeant ovipositing at the tip of a leaf of Glochidion zeylanicum.

Two views of an egg of the Malay Staff Sergeant.

The egg takes about 3 days to hatch. The young caterpillar nibbles away a portion of the egg shell to exit and then proceeds to devour the rest of the egg shell. It has a cylindrical body in pale, dull brown, and an initial body length of about 2.2mm. The body is covered with fine setae, and dorso-lateral, lateral and sub-spiracular tubercles. The head is brown in colour.

Two views of a fully developed egg of the Malay Staff Sergeant.

A newly hatched caterpillar of the Malay Staff Sergeant eating its egg shell.

Two views of a newly hatched caterpillar, length: 2.2mm.

An early 1st instar caterpillar of the Malay Staff Sergeant, observed in the field, resting on the frass extension a few hours after hatching.

Soon after its emergence, it feeds on the leaf lamina around the midrib, and builds an extension of the exposed midrib using frass pellets sewn together with silk. The first instar lasts about 3 days with the body length increases to about 4.8mm.

Two views of an early 1st instar caterpillar, resting at the base of the frass extension, length: 2.5mm.

Two views of a late 1st instar caterpillar, length: 4.8mm.

In the 2nd instar, the dorso-lateral, lateral and sub-spiracular tubercles have short, branched setae. An obscure, pale brown diamond-shaped dorsal patch stretches from the 5th abdominal segment to the 4th abdominal segment, and a similar coloured triangular dorsal patch extend from the 3rd abdominal segment to the 4th abdominal segment. The dark brown head is adorned with short pale brown tubercles and peripheral spines. The 2nd instar lasts about 2 days with the body length reaches about 7.0mm.

Two views of a 2nd instar caterpillar, newly moulted, length: 4.2mm.

Two views of a 2nd instar caterpillar, length: 5.8mm.

Two views of a late 2nd instar caterpillar, dormant prior to its moult, length: 7mm.

A 2nd instar caterpillar of the Malay Staff Sergeant, observed in the field, resting on the exposed midrib.

Another 2nd instar caterpillar of the Malay Staff Sergeant, observed in the field, resting on the exposed midrib.

The 3rd instar caterpillar mostly resembles the the second instar caterpillar, but with the body tubercles, setae and the head spines longer and more prominent. The body and the head are now much darker, and pale brown dorsal patches more distinct. Another pale brown triangular dorsal patch appears from the 3rd thoracic segment to the 1st abdominal segment. There are also whitish sub-spiracular processes from the 7th to 9th abdominal segments. This instar takes about 2 days to complete with body length reaching about 10.5mm.

Two views of a 3nd instar caterpillar, early in this stage, length: 7mm.

Two views of a 3nd instar caterpillar, length: 10.5mm. Inset: head.

Two views of a late 3nd instar caterpillar, dormant prior to its moult, length: 10.3mm.

A 3rd instar caterpillar of the Malay Staff Sergeant, observed in the field, resting on the exposed midrib.

The 4th instar caterpillar has distinctly longer dorso-lateral branched setae in the 2nd and 3rd thoracic segments than the other setae. Peripheral head spines are longer than those in the previous instar. The anterior two dorsal patches are initially brown, but turning green in the latter part of the instar. The dorsal patch in the 5th abdominal segment is reddish brown. The 4th instar lasts about 3 days with body length reaching about 15mm.

Two views of a 4th instar caterpillar, early in this stage, length: 10.5mm.

Two views of a 4th instar caterpillar, length: 10.5mm. Inset: head.

Two views of a late 4th instar caterpillar, length: 15mm.

The next moult brings the caterpillar to its 5th and final instar. Now the dorso-lateral setae are alongated and branched distally and peripheral spines on the head becomes much more pointed. The body colour is initially brown, but turning green within a day. At the same time, the dorso-lateral and lateral processes turn reddish, and the anterior two dorsal patches disappear. The dorsal patch on the 5th abdominal segment is rectangular, black initially but turning purplish to reddish brown as the body turns green. The head is dark brown to black, with off-white or pale pinkish front stripes flanking the adfrontal suture. Tubercles present on the head are of the same colour as the frontal stripes.

The heads of two 5th instar caterpillars, showing the two different colorations of stripes and tubercles.

Two views of a newly moulted 5th instar caterpillar.

An early 5th instar caterpillar of the Malay Staff Sergeant sighted in the field.

Two views of an early 5th instar caterpillar, with the body colour transiting to green.

Two views of a 5th instar caterpillar, with the body colour transition completed, length: 27mm. Inset: head.

Two views of a 5th instar caterpillar, length: 32mm.

Towards the end of the 5th instar, the body ground colour changes to yellow and then pale beige, with pale blue to dark blue markings appearing on all body segments. The tubercles one the head also turn bluish. The 5th instar lasts about 4-5 days, with the body length reaches up to 32mm. On the last day, the caterpillar ceases feeding and seek out a spot on the underside of a leaf (usually on the midrib) or rarely a stem/branch to spin a silk pad. It then anchors itself there via its posterior end, and assumes its upside-down pre-pupatory pose.

Two views of a 5th instar caterpillar, late in this stage, length: 32mm.

Two views of a prep-pupa of the Malay Staff Sergeant. Left: early; Right: late.

After one day of the immobile pre-pupatory phase, the transformation process to the pupal phase takes place. The pupa has cephalic protuberances with their apices extended laterally. There are also dorsal protuberances on the first thoracic segment and the first abdominal segment which are long and keeled, and pointed towards each other. The pupa is initially pale biege brown but turning orange-brown with golden patches a few hours later. Length of pupae: 22-23.5mm.

Three views of a pupa of the Malay Staff Sergeant.

After about 6 days of development, the pupa becomes darkened in color, and the spots/streaks on the forewings are visible through the pupal skin in the wing pads. The next day the eclosion event takes place with the adult butterfly emerges to start the next phase of its life cycle.

Three views of a fully developed pupa of the Malay Staff Sergeant.

The eclosion events of both male and female Malay Staff Sergeant.

A newly eclosed female Malay Staff Sergeant.

A newly eclosed male Malay Staff Sergeant.


References:
  • Discovery of the previously overlooked female of Athyma reta, and its taxonomic implications, L.G Kirton, H. Tan, C-K Phon adn S.K. Khew, Raffles Bulletin of Zoology, vol. 68, pp.249-266.
  • [C&P4] The Butterflies of The Malay Peninsula, A.S. Corbet and H.M. Pendlebury, 4th Edition, Malayan Nature Society, 1992.
  • Butterflies of Thailand, Pisuth Ek-Amnuay, 2nd Edition, 2012.
  • A Field Guide to the Butterflies of Singapore, Khew S.K., Ink On Paper Communications, 2nd Edition, 2015.
Text by Horace Tan, Photos by Sunny Chir, Khew SK and Horace Tan

Sunday, June 7, 2020

Oh Crab!! I Didn't See It Coming!

Oh Crab!! I Didn't See It Coming!
Crab Spiders - Butterfly Killers


A Crab Spider (Thomisus sp.) lying in wait on a Purple Snakeweed flower

Despite the rather melodramatic title, it is well known that butterflies and their caterpillars form part of the food web in any study of animal ecology. Mostly seen as "prey", our flying jewels are almost at the base of the food pyramid and are often targeted as a source of nutrition by a wide range of predators from birds to lizards to spiders.


A Chocolate Demon that appears to be feeding on a flower, but is actually dead - a victim of a fatal spider bite

Butterfly watchers and photographers often encounter dead butterflies on flowers and sometimes wonder how the poor butterfly met its demise. Often, the "murderer" is hidden in plain sight, and still holding tightly onto its prize. One needs to look a little closer and the culprit is very likely to be a  well-camouflaged Crab Spider (Family : Thomisidae   Genus : Thomisus).


A Great Eggfly immobilized by a Crab Spider whilst it was feeding on the flower of the Spanish Needle

This article explores the observations of some of our Singapore photographers who have come across such fatal encounters between various species of Thomisus spiders and their unfortunate butterfly preys. In quite a few of the cases, it would initially appear that the butterfly is still feeding on the flower, but the absence of any movement for some time would suggest that the butterfly may be dead.


A Crab Spider well camouflaged amongst the white flowers of the Stringbush attacked this Leopard

A Brown Awl makes a juicy meal for the Crab Spider hidden amongst the flowers

The generic English name Crab Spider is used to describe over 2,100 species in 175 genera of spiders of the family Thomisidae. Most of these species of spiders in this family are not regularly encountered by our butterfly watchers. However, of particular interest are the species in the genus Thomisus and what we usually also refer to as "Crab Spiders" or "Flower Crab Spiders".


A Crab Spider (Thomisus sp.) with its aggressive "crab-like" stance when it is disturbed

According to online sources, there are globally about 150 described species in the genus Thomisus. The genus includes species that vary widely in their behaviour and ecology. However, like several other genera in the family Thomisidae, their English common name of Crab Spiders probably came from their crab-like motion and their way of holding their much-longer front legs, reminiscent of a crab spreading its claws as a threat.


Danger lurks above! An unsuspecting Orange Emigrant feeding on the flower of the Purple Snakeweed.  Its next stop may be its last!

Crab spiders are considered hunters (as opposed to spinning a web to capture their prey). However, the vast majority of crab spiders are not active hunters that vigorously chase down and kill their prey. The Thomisus crab spiders prefer to sit camouflaged amongst flowers until an unsuspecting prey wanders too close whilst feeding. The strong clawed front legs of the crab spider then quickly reach out and grab the butterfly and inject them with immobilizing venom, hardly moving their body or shorter back legs at all.


A Crab Spider (Thomisus sp.) waiting on the flower of the Singapore Daisy for its next prey

The bushy flowers of Syzygium sp. are ideal for the Crab Spider's concealment as it strikes at the 3-Spot Grass Yellow


A close up of a Crab Spider holding tightly on to its Skipper dinner

Another interesting point about these Crab Spiders is that quite a few are brightly coloured, and are even able to change their colour to match that of the flower that it is hiding on! Talk about chameleon colour-changing capabilities in a spider! The small ones that we see in Singapore are not harmful to humans, but is able to bite if threatened. There are about 45 species of Thomisidae species in Singapore, according to local spider expert Joseph Koh.



A meal fit for a king - Crab Spiders with large prey like the Cruiser - Top : Female   Bottom : Male

A Dark Grass Brown falls victim to a Crab Spider whilst feeding on grass flowers

A Crab Spider finishing off its tiny meal of a Lesser Grass Blue

When there are no butterflies for a meal, a moth will also do! 

From the variety of butterfly species that have been killed, the Crab Spider does not pick and choose which prey to attack. Whatever comes its way - bees, ants, moths, butterflies, will be fair game for it. Even size does not matter! From the tiny Lesser Grass Blue to large butterflies like a Common Mormon or even a Cruiser, the Crab Spider will find a juicy meal.


Danger lurks below! A Crab Spider waiting for the right moment to strike as a Club Silverline perches unaware of the killer spider just millimetres away

The momentary distraction that a flower's nectar offers to a hungry butterfly is enough time for the Crab Spider to sneak up on the unsuspecting butterfly to grab it. Hence whether it is a fast-flying Skipper or a large and robust Cruiser that is many times the size of the spider, the distracted butterfly is vulnerable. There is probably enough venom in the spider's bite that can easily immobilize a prey of any size - provided the spider can hang on to it long enough, whilst the venom paralyses the prey.


Distasteful species like the Crows and Tigers are fair game as far as the Crab Spider is concerned.  The toxins in these species do not seem to affect spiders.

Another observation is that the aposematic colouration and distastefulness in butterflies that feed on toxic plants that endow them with "immunity" from predators like birds, do not apply to spiders. Hence there are equally as many "distasteful" Crab Spider victims as there are other species of butterflies, like the Plain Tiger shown above and other Danainaes.


And the one that got away.... almost.  A dead but relatively pristine Lesser Jay found near where a number of Crab Spiders lay in wait at a flowering Syzygium tree.  This one must have been bitten, but manage to get away before the venom paralysed it. But it did not fly far before it succumbed to the Crab Spider's venom, and died. 

In some cases, the butterfly victim still manages to get away from the clutches of the Crab Spider, despite being bitten and poisoned. However, it is not likely that the prey will be able to fly far, as the spider's venom starts to work. This could explain why some dead butterflies are found with their wings intact and not damaged at all.

Text by Khew SK : Photos by Janice Ang, Foo JL, Heng ZX, David Ho, Khew SK, Michael Khor, Koh CH, Lim CA and Richard Ong

Further Reading :