Hiding in plain sight - Butterfly Gear

The Lepidoptera Log - by Steve Woodhall

Hiding in plain sight

How Not to Be Seen

(with apologies to Monty Python’s Flying Circus)

As insects, butterflies and moths occupy a low position in the food chain, positioned just above the plants that serve as their primary food source. Many animals prey upon them. Over hundreds of millions of years, these species have developed various adaptations to reduce the likelihood of predation by sight hunters.

Some strategies include developing toxicity (and displaying warning colouration), mimicking toxic species, or using camouflage to avoid detection. Predators such as birds and mammals often hunt visually and may overlook individuals that resemble dead leaves, twigs, or rocks. The wings of adult butterflies and moths facilitate these forms of concealment. It’s not just the colours and patterns on the wings that can hide them. There are all sorts of strategies like patterns that break up their outline. Many moths, in particular, have resting postures that add to the camouflage effect.

Not only do adults employ these tactics; caterpillars and chrysalides are also targeted by predators, especially birds that feed them to their offspring. While they lack wings, their coloration and shapes often allow them to blend into their surroundings.

This adaptive strategy is known as camouflage, and it is common among butterflies and moths. Here we’ll show you a few examples from South Africa.

Because many moths are night-flying (nocturnal) they need to take refuge when at rest during the day. Having cryptically marked forewings and resting with them facing outwards allows many moths to ‘hide in plain sight‘. The image many people have in mind when they hear the word ‘camouflage’ is of a soldier wearing ‘camo kit’. The neutral earth and leat tones combine on this Accented Hawkmoth Nephele accentifera (a member of the family Sphingidae in the superfamily Bombycoidea) to break up its outline and disappear into the background. Its caterpillar uses a different strategy. Its tail horn (to the right) is the same colour as the leaf petioles on the Ficus (fig) host plant and it has pale lateral stripes that mimic the leaf’s veins. Its body is countershaded, being paler on the dorsal (back) surface and darker ventrally (on the belly). This allows it to blend into the foliage and avoid being seen.

There is great variety among the Hawkmoths. Verdant Hawkmoth Euchloron megaera appears to be extremely conspicuous when placed on bark but among green leaves it vanishes, as shown in the slide show in the header. Grey Temnora Temnora murina is plainer in appearance and relies on its dull colour to protect it when at rest.

Superfamily Noctuoidea comprises two large families, Noctuidae and Erebidae. The latter includes many small species that use camouflage. The Decorous Widebar, Metachrostis decora, is difficult to see when resting on mossy tree bark with lichen patches. The Eyed Snout, Hypena erastrialis, is found in woodland areas among dead bark and leaves; its straight median line resembles a leaf midrib.

Walker’s Owl Erebus walkeri is the largest member of the Erebidae with a wingspan of up to 65mm. It spends most of its time hidden in shady places or flying at night but if it is caught out in the open (as this one was) its wavy patterning hides it well against tree bark. Narrow Jigsaw Dysgonia properans is another small species whose geometric forewing patterns help to break up its outline when resting amongst dead leaves.

The family Noctuidae is difficult for the uninitiated to separate from Erebidae because the critical difference lies in the pattern of wing veins in the forewings. This is not easily visible in naturally posed specimens. It has many well-camouflaged small species like White Quaker Mentaxya albifrons which like the Narrow Jigsaw has patterns that break up its outline. White-vein Meliana, Meliana tenebra, is a grassland species whose streaked forewings hide it well when resting among grasses.

The Geometridae family of moths within the superfamily Geometroidea is renowned for its advanced camouflage abilities. Adult moths in this family typically rest with their wings open, exhibiting appearances that mimic leaves (both living and dead), bark, lichen, bird droppings, or twigs. The term ‘Geometridae’, meaning ‘earth measurers’, refers to the locomotion method of their caterpillars. These larvae possess three pairs of articulated ‘true’ legs on the thorax near the head, with only two pairs of ‘prolegs’ at the posterior end of the abdomen. When stationary, they often adopt a rigid posture on their prolegs, closely resembling dead twigs, leaf petioles, or dried leaves. During movement, they extend their bodies forward to grip the substrate with their true legs, then release the prolegs and bring the rear of their bodies towards the head, forming a loop. This sequential motion—alternately grasping and releasing with each set of legs—creates a distinctive looping gait, giving rise to the impression of ‘measuring’ the surface beneath them, which is reflected in their name. The Common Bark moth, Ascotis reciprocaria, is a classic example of this, as shown in the video of its caterpillar walking.

Two notable examples highlighting the diversity within the Geometridae family are the Storm Wave, Chrysocraspeda leighata, and Neptune’s Vestal, Traminda neptunaria. While the adult Storm Wave exhibits striking coloration when displayed on foliage—as demonstrated by this reared specimen—it becomes nearly imperceptible against tree bark. In contrast, Neptune’s Vestal is commonly recognized as the quintessential ‘little green moth on a wall,‘ often serving as an introduction to these moths for many individuals. Its presence among green leaves renders it difficult to detect.

The caterpillars of these species further illustrate distinct camouflage strategies employed by Geometrids. The Storm Wave caterpillar, when adopting a rigid posture, closely resembles the midribs of its host plant, the Hiccup-nut Combretum bracteosum, making it virtually indistinguishable from its surroundings. Similarly, the larva of Neptune’s Vestal mimics the appearance of a dried leaf from its host, a Thorn-tree Vachellia species, enhancing its concealment.

The Confused Omphax Omphax bacoti is a member of the Geometridae family that does not rest with its wings fully open. This species is diurnal, and when disturbed, it exhibits erratic flight patterns within its grassland habitat before rapidly descending and concealing itself. Upon closer examination, individuals can be found hiding beneath green leaves, where they rest with their wings folded downward. The Scribbled Duster Pingasa distensaria, another notable Geometridae species, is relatively large and at rest closely resembles a patch of lichen. This camouflage is effective on tree bark and equally so in environments where lichen is present, such as rocks or walls.

The superfamily Bombycoidea encompasses several moth families aside from the Sphingidae. Notably, the Notodontidae are recognised for their conspicuous and weirdly shaped larvae, although the adult moths—such as the Olive Prominent, Desmeocraera latex, typically exhibit effective camouflage. Their grey-green colouration provides concealment on tree bark or among foliage. The Saturniidae family includes some of the largest moth species, including the Saturnine Emperor, Lobobunaea angasana, which is the largest moth in South Africa with a wingspan reaching up to 210mm. The forewings of this species are a subdued grey-brown colour that mimics dead leaves, complete with a simulated ‘midrib’. When at rest with forewings covering hindwings it is almost invisible among fallen leaves on the forest floor. This species employs not only camouflage but also a secondary defence mechanism: when threatened, it exposes its hindwings marked with prominent red eyespots. This startle display induces a shock reflex in potential predators, affording the moth an opportunity to evade capture.

Another representative, the Related Lappet, Leipoxais acharis of the family Lasiocampidae, demonstrates defensive adaptations at both life stages. The caterpillars use chemical deterrents such as irritating bristles or stinging hairs, while adults employ morphological mimicry, resembling dead leaves.

The Pyraloidea constitutes another substantial moth superfamily, primarily consisting of small to minute species. Within it is the Crambidae family, many of which exhibit vibrant coloration; however, the subfamily Crambinae, commonly referred to as ‘Grass Moths’, predominantly rely on cryptic colouring for concealment among dry grass stems and leaves. The Shining Grass Moth, Ancylolomia chrysographellus, exemplifies these adaptive strategies.

The superfamily Papilionoidea in South Africa encompasses five butterfly families. While the majority of these species are diurnal, there are exceptions. Many exhibit vividly coloured dorsal wing surfaces, which often serve a role in sexual display. Conversely, when resting with closed wings, these butterflies become susceptible to predation, making camouflage an important defensive mechanism.

Among the largest families within Papilionoidea is the Nymphalidae, commonly referred to as the Brush-footed Butterflies. Many species in this family are noted for their vividly coloured dorsal wing surfaces, or upper sides, which they often display prominently. For concealment, however, these butterflies typically rely on effective camouflage on the ventral surface (underside). A notable example is the Dead-leaf Commodore Precis tugela, whose bright orange upper side contrasts sharply with its ventral side, an exceptional mimic of a dead leaf. The royal blue male Blue-spotted Charaxes Charaxes cithaeron utilizes his striking upper side for display purposes within the forest canopy, particularly to assert dominance over rivals or attract potential mates. Conversely, when feeding at ground level—often on animal dung—the butterfly’s grey-green underside aids in blending into its surroundings. Occasional flashes of the upper wing colours serve as threat displays to deter competitors.

The Satyrinae subfamily of the Nymphalidae, commonly called ‘Browns’ and ‘Ringlets,’ is noted for its use of camouflage. Species such as the Black-haired Bush Brown Bicyclus safitza safitza and the Common Evening Brown Melanitis leda are crepuscular, typically active during dawn and dusk. These butterflies often inhabit shaded wooded areas, resting near the ground among fallen leaves. Their undersides are camouflaged, helping them avoid predators. The Common Evening Brown possesses large forewing eyespots that may deter potential threats, similarly to Emperor Moths. The grass-feeding caterpillars are camouflaged by their colours, although being crepuscular they are seldom seen anyway.

Species within the genus Pseudonympha, also part of the Browns group, are found in grassland habitats. Like other Satyrinae, they display eyespots on their wings which can serve to distract predators. Many such species like the Gaika Brown Pseudonympha gaika have pale streaks on brown backgrounds on their hindwing undersides, aiding concealment in dry grasses.

The Lycaenidae, or Gossamer-winged butterflies, represent another major butterfly family in South Africa, comprising seven subfamilies within the region. These butterflies are generally small, with the largest species attaining a wingspan of approximately 50mm. Despite their diminutive size, they are notable for their distinctive behaviours and striking appearance.

The Poritiinae subfamily is particularly noteworthy for its larval feeding habits, which include non-plant sources such as lichens and cyanobacteria. This adaptation enables them to occupy challenging environments that are inhospitable to many other butterfly species. Within this subfamily, the genus Durbania (commonly known as Rocksitter butterflies) is endemic to South Africa; no comparable taxa exist elsewhere. As the common name implies, adult Durbania typically inhabit rocky areas, predominantly in grasslands, with certain species also present in the Karoo veld.

Durbania amakosa flavida, found near Durban, is classified as a red-listed species. Despite sharing a name, it is not named after the city but rather after Sir Benjamin D’Urban, whose name also inspired the city’s designation. The larvae consume the cyanobacterial component of rock lichens and exhibit effective camouflage, often gathering in groups on the underside of rocks within their environment. The butterflies’ hindwing undersides feature patterns and colours that provide optimal camouflage against lichen-covered rocks. When threatened, the adults rapidly open their wings to display vivid red-and-black upper surfaces. Preliminary evidence suggests these butterflies may be unpalatable to predators; however, further research is required to substantiate this claim.

Within the Lycaenidae family, the subfamily Miletinae is distinguished by its predominantly aphytophagous (non-plant-eating) characteristics. Most species feature caterpillars that are parasitic or predaceous, and certain genera comprise adults that lack functional mouthparts. Consequently, these adults do not consume nectar and must rely on fat reserves accumulated during their larval stage for sustenance. The genus Thestor, commonly referred to as ‘Skollies‘, exemplifies this adaptation: adult specimens inhabit harsh rocky or desert environments and possess camouflage suited to these habitats, such as the Rock Skolly Thestor petra found among limestone formations, or the Roggeberg Skolly Thestor pringlei native to the arid semi-desert regions of the high Karoo. Their upper sides are also dull brown or grey in colour.

The behavioural patterns of the caterpillars are not well documented. Females frequently deposit eggs on dead plant material devoid of greenery, as observed in the Roggeberg Skolly. Other species select egg-laying sites based on the presence of scale insects utilized by their larvae, such as the Basuto Skolly Thestor basutus. This deposition typically occurs in proximity to nests of Pugnacious Ants Anaplolepis custodiens, upon which the caterpillars are wholly dependent, spending their lifecycle within the ant nests where camouflage becomes unnecessary. While some feed on scale insects during early development, all eventually migrate into the ant nests, resembling small white grubs. To date, only a limited number of species have been photographed and documented.

Several notable Lycaenidae species belong to the subfamily Theclinae. These butterflies often feature prominent colouration on their upper wing surfaces and cryptic patterns underneath, such as the Common Fig-tree Blue Myrina silenus ficedula and the Russet Protea Capys disjunctus. The Fig-tree Blue frequently perches on Combretum trees, with its ventral side closely resembling the appearance of the tree’s seed pods. Similarly, the grey underside of the Protea Scarlet provides effective camouflage among the dead flower buds of its host plant.

Russet Protea caterpillars are not reliant on camouflage, as they reside within flower heads, consuming seeds. In contrast, Fig-tree Blues inhabit the shoots and young leaves of fig trees, where their colouration affords them effective concealment.

Some Theclinae are extremely conspicuous as adults like this Southern Sapphire Iolaus silas male. By contrast the caterpillars of this genus are masters of camouflage. Most of them feed on mistletoes in the Loranthaceae family, some of which have leaves covered in short fuzz. Not only does this caterpillar closely resemble a leaf of its host plant, it has covered itself with the fuzz to render itself almost invisible!

The Aphnaeinae subfamily of Lycaenid butterflies includes several visually distinctive species. Many feature vividly coloured ventral surfaces with ‘dazzle‘ patterns designed to disrupt predator recognition—distinct from traditional camouflage. Certain terrestrial Aphnaeinae, such as the Veined Russet Aloeides pierus and Common/Star Opal Chrysoritis thysbe, exhibit earth-toned or stone-coloured undersides that provide effective camouflage while at rest, although some display conspicuous dorsal markings.

The caterpillars of these butterflies are typically associated with various ant species and generally remain concealed during daylight hours.

The Polyommatinae subfamily of Lycaenidae is commonly known as the ‘Blues’ due to the colouration of the upper sides of many species. These butterflies are not typically known for strong camouflage. Most have whitish or grey undersides with dark spots outlined in lighter tones, which blend moderately with various backgrounds and may function as a form of crypsis. For example, the Brenton Blue Orachrysops niobe, now critically endangered, is difficult to distinguish when resting on dead fern leaves. The Sabi Smoky Blue Euchrysops dolorosa displays a bright blue upper side, while its underside provides concealment when at rest on soil.

When in flight, the alternating visibility of their blue and grey colouring can make visual tracking challenging, and their rapid, unpredictable directional changes followed by immediate settling may reduce detection by predators.

Many caterpillars of Polyommatinae exhibit cryptic characteristics. Species of Zebra Blues Leptotes feed on seed pods from several plants, including Plumbago auriculata; these caterpillars resemble the seed pods. The Black-striped Ciliate Blue Anthene amarah caterpillar feeds on Vachellia or Senegalia thorn-trees, and the diagonal lines along its body visually align with the compound leaves of these host plants, aiding in concealment.

The butterfly family Pieridae consists mainly of white and yellow species that may also have darker or red-orange patterns. These characteristics can make them more noticeable, but they also employ various camouflage techniques. The Autumn-leaf Vagrant Afrodryas leda is normally very conspicuous, but here it’s observed resting on yellowed leaves of the Castor Oil plant Ricinus communis. This butterfly is found throughout much of the year in warmer climates and tends to choose yellow leaves as resting spots. It has been seen flying along forest edges and landing specifically on yellow foliage. Research shows that butterflies possess advanced colour vision. Humans are trichromats with cone cells responsive to blue, green, and red light, while pierid butterflies are hexachromats, detecting six wavelengths: ultraviolet, violet, blue, green, red, and infrared. Each lens in their compound eyes contains light receptors specialized for different wavelengths. This capability likely enables butterflies to select leaf colours that closely match their wing colouration for camouflage.

The Pointed Caper White Pseudanaphaeis gidica abyssinica displays a winter or dry season form known as abyssinica, which is commonly seen on dead grass foliage during this period. This form features brown and cream colouring for effective concealment, unlike the summer or wet season forms, which do not exhibit these colours. Camouflage in butterflies frequently relates to seasonal changes in appearance. Although butterflies do not moult like birds, individuals that emerge during the dry season often display different colours or patterns compared to those emerging in the wet season.

Swallowtail butterflies belonging to the family Papilionidae typically represent the largest local species. Few members of this family display camouflage adaptations; most bear undersides that either mirror the vibrant patterns of their upper wings or imitate unpalatable butterflies from other families.

These insects can often avoid predation by simply closing their wings and seeking shelter within dense foliage. Swallowtails commonly emerge from these hiding places at sunrise or after overcast conditions, basking with open wings to warm their flight muscles—a period during which they are vulnerable to predators. If threatened, they react by swiftly folding their wings.

The male Mocker Swallowtail Papilio dardanus cenea is frequently observed sunning itself in this way. Butterfly photographers recognize that early morning walks along forest edges offer prime observation opportunities. With wings closed, this species exhibits an underside pattern resembling dead leaves, and it is known to select roosting spots among dead foliage that matches its wing coloration, similar to members of the Pieridae family.

The male Bush Kite Swallowtail Papilio euphranor also engages in sunbathing. It’s our only endemic Swallowtail and quite rare – it’s only found in high, cool, Afrotemperate forests. It uses a different family of larval host plant to other local Papilio species. Larvae feed on Wild Laurel Cryptocarya woodii instead of the Rutaceae or Apiaceae used by the others. Typically found soaring high over the forest canopy—hence its common name—the cryptically patterned underside of this species is rarely visible due to its active behaviour. However, during mating, these butterflies often close their wings, revealing these concealed markings.

Papilionid caterpillars are large and fleshy and represent an attractive meal to predators. They are usually well camouflaged to protect them. Most are green in colour and counter shaded to assist in hiding them among leaves, like this final instar Narrow Green-banded Swallowtail, Papilio nireus lyaeus.

Some species, like this Emperor Swallowtail caterpillar, Papilio ophidicephalus, have a combination of green and brown bark-like markings to break up their outline. This species is found in forests where many trees have foliose lichens growing on their branches and twigs – this helps in the deception. This individual is exhibiting another protective mechanism of Papilio larvae. It has extended its osmeterium, an organ that flickers to and fro like a snake’s tongue and gives off a strong smell of plant oils. The sudden appearance of this appendage is often enough to startle a small predator into retreating.

The Hesperiidae family, commonly referred to as Skipper butterflies, includes numerous species that exhibit cryptic or subdued colouration. Many species within this family rest with their wings open, similar to moths, displaying upper wing surfaces adapted for camouflage. The Forest Elfin, Sarangesa motozi, exemplifies this trait. These butterflies are challenging to observe in flight due to their speed, agility, and preference for shaded habitats with dappled light. When settled among leaf litter on the forest floor, they become virtually indistinguishable from their surroundings.

A distinct subgroup of Skippers, known as the Sandmen, is characterized by their diminutive size and exceptionally rapid, low-altitude flight. Their erratic movement through the air aptly reflects their common name, as they seem to ‘skip’ and are difficult to track visually. Although not overtly camouflaged when at rest with wings open, their muted coloration and small stature contribute to their inconspicuousness. For instance, the Dwarf Sandman, Ernsta nanus, possesses a disruptively patterned underside that effectively breaks up its outline when resting with closed wings.

Caterpillars of the Hesperiidae are infrequently encountered since they reside within shelters constructed from leaves or debris bound together with silk threads. Both the Orange Elfin caterpillar, Sarangesa phidyle, and the Forest Sandman, Ernsta dromus, were temporarily removed from their shelters for photographic documentation. A distinctive morphological characteristic of all Skipper larvae is the pronounced constriction resembling a ‘neck’ between the thoracic segments and the head. While these species possess green larvae that blend seamlessly into their environment should they venture outside their shelters, others display more vivid coloration, potentially conferring protection through mimicry of unpalatable butterfly species.

Accented Hawkmoth ♂

Accented Hawkmoth caterpillar

Verdant Hawkmoth

Grey Temnora

Decorous Widebar

Eyed Snout

Walker's Owl

Narrow Jigsaw

White Quaker

White-vein Meliana

Common Bark caterpillar 'looping'

Common Bark caterpillar

Common Bark

Storm Wave

Neptune's Vestal

Storm Wave caterpillar

Neptune's Vestal caterpillar

Confused Omphax

Scribbled Duster

Olive Prominent

Saturnine Emperor

Related Lappet

Shining Grass Moth

Dead-leaf Commodore upper side

Dead-leaf Commodore underside

Blue-spotted Charaxes ♂ upper side

Blue-spotted ♂ Charaxes underside

Black-haired Bush Brown ♂ upper side

Black-haired Bush Brown underside

Black-haired Bush Brown caterpillar

Common Evening Brown caterpillar

Common Evening Brown upper side

Common Evening Brown underside

Gaika Brown ♀ upper side

Gaika Brown ♀ underside

Yellowish Amakosa Rocksitter ♂ showing upper side

Yellowish Amakosa Rocksitter underside

Natal Amakosa Rocksitter larva

Natal Amakosa Rocksitter larvae

Rock Skolly

Roggeberg Skolly

Peninsula Skolly caterpillar fed by ants © Alan Heath

Basuto Skolly eating a scale insect © John Joannou

Common Fig-tree Blue ♂

Common Fig-tree Blue

Russet Protea ♂

Russet Protea

Russet Protea caterpillar inside Protea head

Common Fig-tree Blue caterpillar

Southern Sapphire ♂

Southern Sapphire caterpillar

Veined Russet

Veined Russet

Common or Star Opal ♂

Common or Star Opal

Brenton Blue ♀

Brenton Blue ♂

Sabi Smoky Blue ♂

Sabi Smoky Blue

Zebra Blue caterpillar

Black-striped Ciliate Blue caterpillar

Autumn-leaf Vagrant ♂

Autumn-leaf Vagrant ♀ hiding on yellow Ricinus leaf

Pointed Caper White

Pointed Caper White

Mocker Swallowtail ♂

Mocker Swallowtail ♂

Bush Kite Swallowtail ♂

Bush Kite Swallowtail mating pair

Narrow Green-banded Swallowtail caterpillar

Emperor Swallowtail

Forest Elfin

Forest Elfin

Dwarf Sandman ♂

Dwarf Sandman ♂

Orange Elfin caterpillar

Forest Sandman caterpillar