Introduction

The Red Admiral is probably Britain’s most well known butterfly, and certainly one of the most beautiful, with it’s almost perfect pattern of red bands and white spots on a black background. It was first named as “the Admirable” by Moses Harris in 1766.

The butterfly is widely distributed and common throughout mainland Europe and north Africa. In northern Scandinavia it is a regular migrant. In Britain its status has changed over the years. Up to the end of the 19th century it was considered to be a resident that hibernated as an adult butterfly.

During the 20th century it occurred as a migrant, although there were occasional years when a few adults managed to survive the winter at sites along the south coast of England. Recently there has been a further change in status. The butterfly can now be seen flying regularly on sunny days even in mid winter in the southern counties of Hampshire, Dorset and Sussex. There is now evidence also that larvae resulting from eggs laid in late autumn can successfully overwinter and produce a new generation of adults in early spring.

The Red Admiral cannot be confused with any other European butterfly except the similar Indian Red Admiral Vanessa indica, which has a much wider, irregularly shaped red band on the forewings. It occurs together with atalanta on Madeira and the Canary Islands, and throughout much of the Oriental region, but is inexplicably absent from Europe and north Africa.

Elsewhere in the world there are several other Red Admiral species, including buana from Sulawesi, samani from Sumatra, dejeani from Bali, and the very beautiful gonerilla from New Zealand.

Habitats

This Red Admiral occurs in Britain as a migrant originating from north Africa and southern Europe, and being a highly mobile species, it can turn up in any habitat including woodlands, grasslands, meadows, heathlands and moors, coastal habitats, riverbanks, low montane habitats, gardens, parks, allotments and town centres. Males also gather at certain grassland hilltop sites, apparently to intercept migrating females.

The number of migrants varies according to climatic conditions in Europe, and in turn this greatly affects the number of UK bred butterflies seen later in the year.

Migrants arriving in the early spring oviposit on stinging nettles growing alongside hedgerows or in woodland glades, producing a summer brood in the UK which typically emerges from mid July to early August. In early autumn these butterflies migrate south, arriving in southern woodlands where they congregate to nectar at ivy blossom.

There is now mounting evidence that in mild winters a small percentage of Red Admirals are able to successfully over-winter in southern England. They awake on warm days, and sightings are frequent in southern woodlands on sunny days between October and late January.

In harsh winters Red Admirals are usually killed by the hard February frosts, but since the turn of the 21st century the species has successfully overwintered in significant numbers in many parts of southern England. In the winter of 2006/2007 for example there were almost unbroken sightings in West Sussex, Hampshire and Dorset from October until the following March. The surviving adults laid eggs as early as January, producing a new generation of adults which emerged in early May.

The chart above depicts the build up of Red Admirals following southward migrations into Stansted Forest, 4 miles from the English south coast on the Hampshire / West Sussex border; and the rapid reduction in the population following the gales, frosts and snow of late January & early February. In early March most of the surviving Red Admirals dispersed into the surrounding countryside. Their progeny emerged in May, coinciding with the arrival of a new wave of immigrants from Europe.

On 9th November 2007, I counted a minimum of 12 Red Admirals in a small section of Stansted Forest, but estimated that at least 150 must have been present in the whole wood.

Unfortunately, extremely wet and windy conditions prevailed over the following few weeks, decimating the population. On 23rd December however, I watched 2 Red Admirals basking on tree-trunks in the forest, despite having endured 6 consecutive nights of severe ( minus 3-C ) frost the previous week. At the end of January, in mild sunny conditions at least 3 Red Admirals were flying in the forest, but cold and wet conditions persisted throughout the spring, killing off the last survivors, and none were seen at Stansted after 9th February.

Compared to the previous winter, 2007-2008 was cooler, windier and considerably wetter, with fewer sunny days. Frosts were minimal prior to March, which produced many icy nights. April began very cold, with snow, sleet and hail. The harsh winter conditions had no negative impact on other hibernating butterflies, with Peacock, Comma, Brimstone and Small Tortoiseshell all being recorded in average numbers in mid-late April.

The low number of Red Admirals arriving in autumn 2007 was mainly attributable to lower breeding success and reduced immigration during the exceptionally cool and wet summer. The maximum count was on 21st October when 17 were recorded nectaring at ivy flowers.

Winter survival rates were very low, with only one sighting in March, and no more until 2 immaculate and extremely active individuals were seen on 25th May. Their condition and behaviour strongly suggested  they had emerged locally within the previous couple of days – providing fairly conclusive evidence that they were the progeny of post-hibernation adults that had successfully over-wintered at Stansted Forest.

Lifecycle

In late April and early May, female Red Admirals can be observed flying around nettle patches in woodland glades. They periodically alight on nettle leaves Urtica dioica, upon which they lay a single egg. Usually a dozen or so eggs will be laid on any sizeable nettle patch. Egg-laying is interspersed with periods of basking and nectaring – typically at blackthorn, crab apple, and wild cherry.

Second brood adults, which may be of either UK or European origin, are less fussy about oviposition sites. They will lay their eggs in any sheltered sunny lowland habitat where the foodplant grows e.g. in July 2008 I watched 2 Red Admirals laying eggs on the same small clump of unkempt nettles in my garden. Each laid about a dozen eggs between about 11.00am and 1.00pm.

The eggs were all laid singly on the upperside of terminal nettle leaves. The egg-laying bouts, which each lasted about 2-3 minutes were interspersed with periods of basking, and periodic flights to nectar at a Buddleia bush a few metres away. Interestingly a Comma was ovipositing simultaneously on the same nettle patch, and neither species seemed disturbed by the presence of the other.

The larva lives within a tent of folded nettle leaves, spun together with silk. Every few days, as the leaf-tent gets devoured, the larva moves house, and spins a new tent nearby. If the tent is opened, the spiky greenish-brown larva can be seen within, normally curled in a J shape, head-downwards.

The larvae of most butterfly species tend to wander away from their foodplants to pupate, but the Red Admiral is unusual, forming it’s greyish chrysalis within the final leaf-tent.

Adult behaviour

On sunny days the butterflies often bask on tree-trunks, always assuming a head-downwards or sideways facing position. In warm but overcast weather they like to bask on the foliage of bramble, hazel and other bushes.

In early spring Red Admirals nectar at sallow catkins and the blossom of blackthorn and hawthorn. 

Favoured summer nectar sources include dogwood, hemp agrimony, bramble, marjoram, devil’s bit scabious, ragwort, burdock, spear thistle and ivy blossom. They also visit dung, or imbibe mineral salts from damp ground, and attend sap runs on oak trunks. In gardens they will nectar at Buddleia, ice-plant and michaelmas daisies. In orchards they will feed at fallen apples or pears.

Males are highly territorial, chasing after Commas and Peacocks as well as other Red Admirals. When nectaring however they will accept the presence of other species, and I have often seen mixed groups of Red Admirals, Silver-washed Fritillaries, Commas, and Large Whites gathered on a single flower head of hemp agrimony or Buddleia.

At night, on cold overcast days, and during the winter months, Red Admirals roost head-downwards on the trunks or lower branches of oaks, larches and other trees, where the bark-like underside of the wings provides them with excellent camouflage.

Aglais urticae, Noar Hill, Hampshire – Adrian Hoskins

Introduction

The Small Tortoiseshell may have been present on Earth longer than any other butterfly species – a mid-Miocene fossil of Aglais karaganica estimated at 15 million years old is so similar to the modern day Aglais urticae as to be virtually indistinguishable!

Small Tortoiseshells vary considerably in appearance – some examples being a unicolorous dull orange, while others are much redder with contrasting yellow patches. In summers when weather conditions are cool, a high percentage of the butterflies tend to have the black markings enlarged, or merging together. In hot summers, and in warmer parts of the butterfly’s range these markings often become reduced in size.

The race ichnusa, found in mountainous areas of Corsica and Sardinia has a fiery orange ground colour, and lacks the colon of black spots on the forewing. Some authors regard it as a separate taxon. Captive reared adults however closely resemble the normal form, so it seems likely that ichnusa is merely a habitat / climate generated form of urticae.

The Small Tortoiseshell occurs across the whole of Europe and temperate Asia. A closely related and very similar species Aglais caschmirensis is found in mountainous regions of Kashmir, Sikkim, Bhutan and Tibet. In the western Himalayas a third species occurs, Aglais ladakensis, which looks like a very heavily marked version of urticae. The only other member of the genus Aglais milberti is a north American species in which the basal half of the forewings is black, and in which the 2 small black spots in the median area are absent.

Aglais urticae 2nd brood, Hungerford, Berkshire – Adrian Hoskins

The butterfly has always been subject to dramatic fluctuations in abundance from year to year and in some seasons it can be very scarce. Historically it has always had the ability to bounce back, and was until recently regarded as one of Britain’s most familiar and common butterflies. However after about 1990 the species declined sharply in southern England, until the low point of 2007 when most recorders in Hampshire, Surrey and Sussex failed to see even a single specimen. There was some evidence of a minor come-back in 2008, when most observers saw at least one or two specimens of the late summer brood. Survival of hibernating adults in the cold, prolonged winters of 2008, 2011 and 2012 was exceptionally good, and by the summer of 2013 populations in many areas of Britain were fully recovered.

The cycles of abundance are the result of climatic variations which affect the population dynamics of the butterfly and its parasitoids, particularly the fly Sturmia bella ( Tachinidae ). The latter occurs mainly in warmer climates and first arrived in Britain in 1998. It lays its eggs on nettle leaves. When a Small Tortoiseshell caterpillar eats the leaves, the microscopic eggs are ingested undamaged and pass into its gut. There the eggs hatch, and the resulting grubs bore their way through the soft flesh, consuming non-vital body tissues. When they are almost full grown they eat the vital organs, and then break out through the skin of the dying caterpillar to pupate.

The fly is widespread throughout southern Europe, north Africa and Asia. In Japan it parasitises the caterpillars of a Danaine, Parantica sita. Research by Hirai & Ishii has revealed that P. sita can often survive attack by Sturmia bella – in experiments over 70 percent of infested sita larvae survived and went on to produce perfect adult butterflies. Sadly the Small Tortoiseshell lacks this immunity.

Although current research suggests that Sturmia bella is at least partially responsible for the decline of the Small Tortoiseshell, experiments have shown that only about 18% of Small Tortoiseshells are affected by parasitoids, and Sturmia bella is just one of several that attack the butterfly. It is also notable that other nettle-feeding Vanessids e.g. Peacock, Red Admiral and Comma are apparently less affected, even though their larvae also ingest the Sturmia eggs. The Comma and Peacock in fact increased their numbers substantially between 2005-2009 when urticae was at its nadir. All 3 species have similar biochemistry to that of urticae, so it seems unlikely that they possess a natural immunity. One cannot help wondering therefore why the Small Tortoiseshell seems to have been singled out, while Red Admiral, Comma and Peacock continue to thrive.

Habitats

The Small Tortoiseshell is a highly mobile species which can occur in almost any habitat, including woodlands, grasslands, heaths, gardens, country lanes, and even in city centres. It is a strong flyer, with high powers of dispersal, reaching remote islands and high mountain peaks. Adults have been recorded at altitudes as high as 1200m in Scotland, and larval nests have been found at 3350m. I have watched adults flying near the peak of Ben Lawers in Tayside, and high in the mountains at Langdale Pikes and various other sites in the Lake District.

The butterfly is perhaps commonest on dairy and cattle farmland, where the application of fertilisers and manure enriches the soil, encouraging the growth of stinging nettles Urtica dioica and U. urens.

Lifecycle

In Scotland there is usually only a single generation, emerging in early July, entering hibernation in August, and reappearing in March or April.  Elsewhere in Britain there are normally 2 broods per year, one emerging in June, and the other in August. Adults of the latter brood enter hibernation in September or October and awaken the following March. The resident UK population is supplemented by migrants from mainland Europe. The numbers arriving however are normally very low, and have little effect on population levels.

In spring female Small Tortoiseshells are often seen flying around young nettle patches, where they lay their greenish eggs in large untidy heaps of 80-100 on the underside of the upper leaves. Often more than one female will oviposit on the same leaf, sometimes simultaneously. Females typically select young plants growing near the edge of a nettle bed, and always growing in warm, sunny and sheltered conditions. Typical breeding sites include field margins at the bottom of south-facing hills, dykes, riverbanks, railway cuttings, and nettle patches growing on the south side of hedges or walls.

Aglais urticae, egg batch on stinging nettle ( Urtica ), Hampshire – Adrian Hoskins

The summer brood is less fussy about oviposition sites, and will lay on nettle patches in gardens, in woodland glades and rides, and on exposed hilltops.

In June 2009, I watched a female searching for oviposition sites at Martin Down in Hampshire. She spent about half an hour flitting back and forth, settling for a few seconds here and there on various terminal leaves, and took a great deal of time before she eventually found one which suited her. To my eyes it looked no different from any of the other leaves on which she landed, so I assume that she just found that particular leaf easier to grip. Once she had decided where to lay her eggs she clung tenaciously to the leaf, which was constantly battered by strong gusts of wind, until she had laid a batch of about 80 eggs.

Aglais urticae, 1st instar larvae on Urtica, Hungerford, Berks – Adrian Hoskins

The eggs hatch after about 12 days. Immediately after hatching, the larvae devour their empty egg shells, and then spin a communal silk web around the terminal leaves of the nettles. They shelter within the web at night, or in adverse weather conditions, and feed avidly whenever the sun shines. If disturbed the larvae react in unison, wriggling and jerking as a defence against parasitoid wasps or flies. Nevertheless a high proportion of larvae turn out to be parasitised, usually by the Tachinid fly Phryxe vulgaris. When young, they can easily be mistaken for the caterpillars of the Peacock, but Small Tortoiseshell larvae are paler, and even when quite small it is usually possible to discern pale lines running along their backs.

Aglais urticae, 2nd instar larvae on stinging nettle, Dorset – Adrian Hoskins

As they grow, they split up into progressively smaller groups, spinning a new web after each moult. The final instar sees a change in behaviour, with the larvae abandoning their webs to live solitarily. By this time they are a dull blackish colour, spiky, with broad yellow lines running along their backs and sides. These lines are usually prominent, but in some batches of larvae they can be pale and obscure. The fully grown larvae can often be seen curled in a J-shaped posture, resting on nettle leaves, and if disturbed will coil into a tight circle and drop to the ground.

The chrysalis is variable in colour, ranging from grey to olive or buff, often with a pinkish or golden metallic sheen. It can be found suspended by the cremaster, on woody stems, fence posts, walls, or beneath the stems or leaves of nettles. The adults emerge at dawn, about 12 days after pupation.

Adult behaviour

Over-wintered adults typically awaken in late March or early April, and remain on the wing until early May. They are usually seen as singletons, nectaring at dandelion, daisies or sallow catkins, often in the vicinity of nettle patches.

On cool days in spring they do not bask, but instead warm themselves up by shivering their wings prior to flight. The rapid shivering generates warmth by friction, and enables the butterfly to fly even in quite cool conditions. On warm days they frequently bask on bare soil or low foliage, but hot sunny conditions cause them to settle for long periods on bare soil with their wings closed, at which time the sombre ‘tortoiseshell’ pattern on the underside affords them excellent camouflage.

Aglais urticae 1st brood, Hungerford, Berkshire – Adrian Hoskins

Males establish vantage points in the early afternoon, from which they await passing females. When a female flies by she is intercepted and the courtship ritual begins. The male chases her until she settles on the ground. If she is receptive she opens her wings and the male approaches her from behind, with his wings also open. He then steps onto her hindwings, which he vigorously drums with with his antennae. The pair then fly a short distance and repeat the process. Any other males which attempt to interfere are briskly chased away by the resident male, who then returns to his female to continue wing-drumming.

This bonding process continues for several hours, until just before dusk, when the female accepts the male’s advances. A this point she leads him to a sheltered and shady spot, typically beneath a bush or hedge. Both sexes then hold their wings erect, and the male walks alongside the female, and curves his abdomen to copulate. After about 20 minutes the pair straighten out to face opposite directions. They remain copulated in this position until the following morning.

The summer brood butterflies, which emerge from mid June to early July, will nectar at almost any available wild or cultivated flowers. Their favourite however is undoubtedly creeping thistle. At Noar Hill in July 2009 I watched a female nectaring for over 2 hours at a clump of these flowers. When disturbed by a bee or hoverfly she would leave the thistles and briefly investigate other nearby flowers including marjoram, knapweeds, bramble, scabious and hemp agrimony, but after a quick taste always returned to the thistles.

The second generation is always more abundant than the first, and emerges at the end of August. In late summer Small Tortoiseshells often visit flowery areas in the countryside, where they congregate to nectar at thistles, ragwort and other Compositae. They also commonly visit gardens, where they are strongly attracted to Sedum, Buddleia and michaelmas daisies. Their mission at this time of year is to use every opportunity to gorge on nectar, building up sufficient protein and fat reserves in their bodies to enable them to survive hibernation.

In southern Britain the butterflies fly until late September, when they enter houses, sheds, churches and unheated farm buildings to hibernate. In the north they enter hibernation earlier, in late July or early August, after just a few days on the wing. They often hibernate communally – I’ve often found groups of 3 or 4 nestling together in an attic or outbuilding, and I once found 17 hibernating adults clustered tightly together on the ceiling in a hotel in Tayside, Scotland.

Aglais urticae 2nd brood, Lardon Chase, Berkshire – Adrian Hoskins

Painted Lady Vanessa cardui, Bentley Wood, Wiltshire – Adrian Hoskins

Painted Lady Vanessa cardui, Bentley Wood, Wiltshire – Adrian Hoskins

Introduction

The Painted Lady is the most widely distributed butterfly in the world, found in North America, and south to the Caribbean islands and Venezuela. In the Old World it occurs throughout Europe and temperate Asia, over most of Africa, Madagascar, the Azores, the Canary Islands, the Arab states, and throughout the Indian subcontinent including Sri Lanka. In the Far East it occurs in Thailand, Malaysia, Borneo and Sumatra – and extends it’s range south through the Indonesian islands to Western Australia.

The cosmopolitan distribution of the species results from a combination of its very strong migratory behaviour and its polyphagous nature – in Britain its caterpillars feed almost exclusively on thistles, but elsewhere in the butterfly’s range they use a vast range of foodplants amongst the Malvaceae, Labiatae, Compositae, Boraginaceae,  Ulmaceae, Rutaceae, Urticaceae, Rosaceae, Verbenaceae, Leguminosae, Chenopodiaceae, Hydrophyllaceea, Convolvulaceae, Plantaginaceae, Cucurbitaceae, Cruciferae, Umbelliferae, Rhamnaceae, and even one or two grass species !A feature common to most migratory species is that there tends to be very little variation in pattern between individuals. Apart from minor variations in colour a Painted Lady from N.America or Europe will be identical to one from Africa, Indonesia or Australia.

Habitats

In Britain the Painted Lady occurs as a migrant originating from North Africa, Arabia and southern Europe. C.B.

Williams in his book Insect Migration quotes Skertchley, who in March 1869 witnessed the commencement of a migration in desert behind Suakin by the Red Sea :”He noticed that the whole mass of grass, through which they were riding on camels, was in a state of violent agitation although there was no wind. When he dismounted he found that the cause was the emergence from the chrysalis of myriads of Painted Lady butterflies, which dried their wings and about half an hour later flew off together eastwards towards the sea”.

Being an extremely mobile species the Painted Lady can turn up in any almost any habitat – heaths, woodland, grassland, meadows, cliff-tops, urban gardens, city parks and town centres.

Migrants arriving in Britain in May often assemble at hilltop sites in southern England, but they soon disperse inland, and can reach northern Scotland by June. Eggs laid in May will produce a second brood of adults in August, which also migrate northwards and reach Scotland in early September. There does not appear to be any evidence of a southward return migration, but occasional records of Painted Ladies in January appear to indicate that the butterflies attempt to hibernate at certain sites on the south coast, although they do not seem able to survive the frosts of February.

The numbers arriving in Britain in the spring depends very much on weather conditions in Africa and continental Europe. A wet winter in north Africa can cause a huge population explosion and trigger a massive migration in which tens of thousands, or even tens of millions of Painted Ladies head north across the Mediterranean. They usually cross from Algeria and Tunisia to Italy, then east through Hungary, and then radiate out across northern Europe, laying eggs and producing successive generations along the route. Others follow a different route, heading north from Morocco to Spain and France, but numbers arriving in Britain via this route are usually lower. The butterflies also spread south from Morocco, heading around the coast of West Africa to Gambia and Nigeria. 

The chart above illustrates the massive immigration of Painted Ladies into Britain during May 2009. More migrants continued to arrive, albeit in much lower numbers, throughout the summer. The 1st brood of UK-bred adults emerged in vast numbers between mid-July and early August. The progeny of these butterflies emerged in much lower numbers during September and early October. Painted Ladies were not the only migrants to appear in 2009. There were also huge immigrations of Large White and Clouded Yellow; and above average numbers of Small White and Red Admiral. Other notable migrants included Long-tailed Blue, Short-tailed Blue & Queen of Spain Fritillary.

Painted Lady Vanessa cardui, Bentley Wood, Wiltshire – Adrian Hoskins

Lifecycle

In Britain the pale green eggs are nearly always laid on spear thistle Cirsium vulgare, creeping thistle C. arvense, or welted thistle Carduus acanthoides, but there are occasional records of nettle Urtica dioica or greater burdock Arctium lappa being used.

The eggs are laid singly on the upper surface of terminal leaves, and hatch after about a week.

The young larvae spin a silk web on the underside of a thistle leaf. They feed on the lower cuticle, leaving distinctive patches visible on the upper surface. When older they eat the entire leaf with the exception of the central vein and the spines. The fully grown larva is black, with a prominent yellow broken stripe along each side. It lives and feeds within a tent of leaves spun together with silk. These silk tents, in which the conspicuous droppings are enmeshed, make the larvae easy to find.

In early July the larva spins together a new tent of leaves, within which it pupates. The chrysalis is greyish pink, with an overall golden lustre.

Adult behaviour

On overcast days, the butterflies often spend long periods basking in rabbit scrapes or other small depressions on the ground, and are frequently seen basking on patches of bare chalk.

Males establish territories at ride intersections within woodland, or in sheltered dry gullies, chalk pits, or around field edges where they are sheltered by hedgerows. I have also commonly found them aggregating at hilltop sites e.g. at Old Winchester Hill NNR, and Noar Hill, both in Hampshire. They generally have about 4 or 5 spots within their territories where they regularly bask.

The butterflies are powerful flyers and habitually fly from flower to flower, stopping for just a few seconds at each to take nectar. They visit a wide range of species including marjoram, knapweeds, thistles, burdock, hemp agrimony, fleabane, devil’s bit scabious, and bramble blossom; and in gardens will nectar at buddleia, michaelmas daisies, sedum and many other cultivated flowers.

Painted Lady Vanessa cardui, Hampshire – Adrian Hoskins

Introduction

The Comma was regarded as a common species in the 18th century but declined to became a great rarity between about 1850 and 1910. By 1930 it was on the verge of extinction but then it’s fortunes began to reverse. By the 1980’s its numbers had completely recovered and it is now once again a common species in southern Britain.

The butterfly gets its common name from the C or comma-shaped silver mark on the underside of the hindwings. The genus name Polygonia is a reference to the beautiful ragged wing shape. This is unique among British butterflies but there are other similarly shaped species found elsewhere in the Holarctic region, including the Southern Comma P. egea which inhabits the Mediterranean area, and the Question Mark and Angle-wing butterflies of North America.

Polygonia c-album, male at roost on birch sapling, Stansted Forest, West Sussex – Adrian Hoskins

The Comma is distributed across most of Europe, but absent from Scotland and sub-arctic regions. Beyond Europe it occurs across temperate Asia to northern China, Korea and Japan. It also occurs in Morocco, Algeria and Tunisia.

Comma Polygonia c-album, male, March, Stansted Forest, West Sussex – Adrian Hoskins

Comma Polygonia c-album, f. hutchinsoni, June, Stansted Forest, West Sussex – Adrian Hoskins

Habitats

In spring Commas breed primarily in woodland edge habitats where stinging nettles grow in damp but sunny situations, typically in glades or at the side of grassy tracks. The summer adults are more mobile however, and can breed in gardens, old quarries, along country lanes and railway cuttings, on sheltered areas of scrubby grassland, and at coastal habitats. They freely roam the countryside and can be found anywhere where stinging nettles or elms grow but favour sunny sheltered sites with bramble bushes nearby, and a profusion of wild flowers.

Polygonia c-album, f. hutchinsoni, July, Titchmarsh Wood, Northamptonshire – Adrian Hoskins

Lifecycle

There are 2 generations each year. Larvae hatching from eggs laid in early March by overwintered females develop slowly, pupating in mid-June to produce adults of the brightly marked hutchinsoni form. These emerge in early July and remain on the wing until mid-August. The overwintered adults however remain alive until late spring and continue laying eggs until mid-May. Larvae hatching from these later eggs develop more slowly to produce normal adults which emerge in August.

Meanwhile the offspring of the hutchinsoni brood produce another generation of normal adults which emerge in September. In late summer it is possible to see both forms flying together. The hutchinsoni adults die by early September. The normal form adults enter hibernation and awaken the following spring.The green, ribbed eggs are laid singly, close to the edge on the upperside of stinging nettle leaves. Commas normally oviposit on nettles growing in sheltered situations along hedgerows or in sunny woodland glades, close to nectar sources such as blackthorn ( spring ) or bramble (  summer ).

Two hundred or so years ago the primary foodplant used in Britain by Commas was hop Humulus lupulus, which was grown all over southern England as an ingredient for ale. The dramatic decline of the butterfly in the late 19th and early 20th centuries is probably linked to changes of practice which greatly reduced this formerly widespread and abundant plant.

Fortunately the Comma was able to adapt to making widespread use of another foodplant, stinging nettle Urtica dioica and this has now  undoubtedly become the primary foodplant in Britain. Larvae are also found occasionally on sucker growth of English elm Ulmus procera, and more rarely on wych elm U. glabra, sallow Salix caprea, hazel Corylus avellana, or cultivated blackcurrant Ribes nigrum.

The fully grown caterpillar is unmistakable, being brownish black, with orange spikes on the front segments, and a long splash of white along the back. It often rests on the upper surface of a leaf, adopting a semi-curled posture, and at a glance can be mistaken for a bird dropping. The chrysalis is marbled in shades of brown, and decorated with spangles of silver and gold. I have occasionally found pupae attached to fence posts, but they are more often found suspended from woody stems or shaded tree trunks.

Comma Polygonia c-album, Steyning, West Sussex – Adrian Hoskins

Adult behaviour

In spring Commas occupy ride intersections and glades, where they nectar at blackthorn blossom, sallow catkins and dandelions. They spend long periods basking on leaf litter, bare ground, on logs, or on bramble leaves or dead bracken. It is also common to see them basking head-downward on fence posts, or on the trunks of birch and ash trees.

In the spring males establish territorial perches on twigs, logs or favoured leaves which they use as bases from which to launch flights to ambush other passing butterflies. Intruding male Commas and butterflies of other species including Peacocks and Orange tips are always ousted by the ‘owner’ of the territory. Summer brood Commas often perch on hazel bushes and intercept Gatekeepers and Speckled Woods. Passing female Commas are intercepted, but I have never witnessed a courtship ritual, or found any mated pairs. Copulation reportedly takes place high in the tree tops.

Comma Polygonia c-album, Stansted Forest, West Sussex – Adrian Hoskins

On warm sunny days in spring, butterflies of both sexes frequently settle on paths, tree stumps or wood shavings to imbibe moisture. Summer nectar sources include bramble, hemp agrimony, wild carrot, marjoram, thistles and traveller’s joy.

In cool cloudy weather, Commas roost openly on foliage, and if disturbed will feign death, falling to the ground, with their wings closed and their white legs tucked tightly against their bodies. Towards dusk they seek overnight roosting sites. One evening in early July 2009, I watched a female settling down to roost – she spent a couple of minutes fluttering around on the shady side of an ivy covered fence, and eventually settled for the night under an ivy leaf, resting on the mid-vein, with her head pointing towards the stem.

On various occasions I have also found Commas roosting at ground level among grasses – in these cases the butterflies always roost head-downward. Their disguise is quite remarkable, giving the impression of a dead oak leaf that has fallen onto the grass.

Comma Polygonia c-album, July, Alice Holt forest, Hampshire – Adrian Hoskins

In early autumn the butterflies nectar avidly at devilsbit scabious, buddleia, ragwort and fleabane in preparation for hibernation. The feeding frenzy continues into September and October when they nectar at ivy, and gorge themselves on sugary juices exuding from fermenting blackberries.

In late October they seek out hibernation sites, usually choosing to spend the winter months hiding in wood stacks, hollow tree trunks, or sometimes out in the open, hanging beneath branches. The butterfly depicted below was found hibernating under a branch of young coppiced sweet chestnut, about 0.5m above ground level in Stansted Forest.

I found it on 26th January 2008 but suspect that it had been hibernating there since the previous October. It was still there in the same position on 9th February, but by 24th February had disappeared, presumably having awoken from it’s diapause with the arrival of sunny and slightly warmer weather.

Comma Polygonia c-album, hibernating openly under a low branch – Adrian Hoskins

Nymphalis polychloros  – Peter Bruce-Jones

Introduction

The Large Tortoiseshell is widely distributed in Europe, and common in the Mediterranean region where it breeds in open woodland areas. It also occurs in Morocco, Algeria and Tunisia; and across temperate Asia from Turkey to Kazakhstan and the foothills of the Himalayas. It can confused with the similar Yellow-legged Tortoiseshell Nymphalis xanthomelas which has much brighter and redder colouring, and enlarged blue lunules on the hindwings. The latter species is only found in eastern Europe and temperate Asia, where it breeds in wooded river valleys. Old records of it’s occurrence in Britain are dubious.

Nymphalis polychloros – Lars Anderson

Habitats

A century ago the Large Tortoiseshell was a common butterfly in southern and eastern Britain. It had always been subject to periodic fluctuations in abundance, but declined very suddenly in the early part of the 20th century, and apart from a short period of abundance in the 1940’s never recovered. By the 1970’s it had become a great rarity.

The butterfly’s decline was originally attributed to increased parasitism but I believe this is unlikely. Parasitoid / host abundance is normally cyclic, and linked to the differing climatic requirements of host and parasitoid. After a period of parasitoid-induced scarcity, butterflies generally recover and return to abundance until climatic conditions again favour the parasitoid.

Whatever the cause of the decline, it was certainly exacerbated by the arrival in Britain in 1976 of a particularly virulent strain of Dutch elm disease, which rapidly proceeded to almost eradicate the main larval foodplants wych elm Ulmus glabra, and English elm U. procera. The disease arrived in Britain on logs imported from North America, and quickly spread across the country, transported by Scolytus elm bark beetles. These live under the bark, and propagate an ascomycete microfungus Ophiostoma novo-ulmi which is responsible for the disease.

By 1990 the Large Tortoiseshell was considered to be either extinct or on the verge of extinction in the UK. It has been suggested by some that it still maintains its presence as a breeding resident in southern England, but that populations have become reduced to the point where the butterfly is now unobservable. It seems very unlikely though, as the very conspicuous larval nests would surely still be found occasionally. None have been reported in Britain for several decades.

Since 1998, sporadic sightings of adults have been reliably reported from sites along the south coast of Hampshire and Dorset. Most of these records have referred to very worn post-hibernation insects seen in early spring. It is unknown whether these represent the progeny of migrants or of captive bred stock. In June / July 2007 fresh specimens were seen at several locations on the Isle of Wight and scattered along several miles of the Hampshire coastline.

Records of overwintered specimens from other coastal sites every spring from 2007 to 2012 appears to indicate that the butterfly is now breeding regularly in low numbers on the Isle of Wight and possibly elsewhere long the south coast although no larvae have been recorded. It is however possible that one or more amateur breeders may be releasing captive-bred European livestock of this butterfly in an attempt to re-establish the species in Britain.

In Europe the Large Tortoiseshell is usually encountered as singletons. It is encountered widely but never in numbers. It is a highly mobile species and breeds in forests and along hedgerows where elms Ulmus glabra & U. procera, sallow Salix caprea, poplars Populus alba & P. nigra, wild cherry Prunus avium, or aspen Populus tremula grow.

Lifecycle

The adults emerge in late June or early July, but are only active for about a fortnight, as they enter hibernation very early – typically in late July, and are not seen again until the following spring. The eggs are laid in April, and hatch after about 3 weeks. They are laid in large batches in a neatly arranged ring around twigs, about 2-4 metres up, on the sunny side of various trees, but particularly on wych elm Ulmus glabra. The larvae are black, covered with sharp orange spikes and peppered with tiny white dots, giving them a greyish appearance.

They live communally in conspicuous silk webs spun on the twigs of the foodplants. If they are disturbed by a bird the whole group jerks in unison. As they grow older they split into smaller groups, and become solitary just prior to pupation. When fully grown in early June, the larvae descend from the treetops, and wander a short distance to pupate.

The pupa is brown, marked with gold spots, slightly spiky in appearance, and resembles a withered dead leaf. It is formed hanging by the cremaster from twigs or branches on the lower part of various bushes and trees. In Turkey I have found them commonly suspended from the walls and eaves of buildings. In my experience at least 60 percent of pupae are found to be parasitised.

Adult behaviour

Males are seen as solitary individuals, usually either in flight, or when basking on bare ground in lightly wooded habitats. In Britain they have been reported nectaring at sallow catkins in spring, and bramble blossom and various herbaceous garden plants in summer.

Females habitually bask on tree trunks, often in a head-downwards posture. The males patrol along wood edges until they locate a female, and then settle beside it. The female then flies off, with the male in hot pursuit, until after an hour or more the pair settle on a tree trunk and copulate. They remain paired all afternoon, and possibly overnight.

In Europe they are sometimes seen flying around the tops of sallows, elms and poplars, but are normally encountered in flight, when dispersing in search of breeding sites. In western France I’ve observed them flying in inhospitable terrain including motorway car parks, petrol stations and town centres, indicating that the species is highly mobile and probably migratory in behaviour.

Near Dijon in central France I once found 3 Large Tortoiseshells clustering together in the company of several bees and a dozen Woodland Graylings, all feeding at a sap run on a hedgerow hawthorn. While feeding the Large Tortoiseshells nervously fanned their wings, but made no attempt to fly away even when I approached very closely. However on other occasions when I’ve encountered this species, they have been extremely wary.

In Turkey I have found recently emerged butterflies hanging from twigs along dry river beds. They remained aestivating on the twigs for several days in late May.

Introduction

There are about 25-30 species in the genus Limenitis. Some such as populi and camilla are widely distributed across Europe and temperate Asia. The majority however are largely restricted to China and the countries of the former Soviet Union.

The White Admiral is distributed across much of southern Britain and throughout central Europe but is absent from northern Scandinavia, most of the Iberian peninsula, and the Mediterranean region with the exception of western Italy. Beyond Europe it is found in Turkey, and across temperate Asia to north China, Korea and Japan.

White Admiral Limenitis camilla, Alice Holt forest, Hampshire – Adrian Hoskins

The White Admiral can’t really be confused with any other British butterfly although novice butterfly-watchers sometimes mistake its silhouette in flight for that of the Purple Emperor Apatura iris which breeds in similar habitats and flies at the same time of year. Purple Emperors however are larger, with more pointed wings and a much more powerful flight.

In southern Europe and western Asia camilla can be confused with its close relative reducta but the latter has a prominent white spot in the discal cell of the upperside forewing, and only a single row of black spots on the underside hindwing.

White Admiral Limenitis camilla, male, Hampshire, England – Adrian Hoskins

Habitats

In Britain camilla can be found in most broadleaved woodlands in the southern and central counties of England, but is commonest in Wiltshire, Hampshire, Berkshire, West Sussex, Surrey, Oxfordshire and Buckinghamshire.

Within these woodlands the butterflies inhabit small sunny glades, narrow grassy rides, and lightly wooded areas. It breeds in places where the larval foodplant honeysuckle grows in dappled sunlight, and hangs as thin wisps from the boughs of oak, ash and various other trees. White Admirals rarely venture away from these areas, and are not normally encountered in clearings, wide sunny avenues or dry open woodland.

White Admiral Limenitis camilla, male, Hampshire – Adrian Hoskins

Lifecycle

The butterflies are normally univoltine, emerging in mid-late June. They have a lifespan of about 2 weeks. Late emerging females can still be seen until the end of July, and sometimes as late as early August. In exceptionally warm summers there may be a partial second brood in early autumn.

Females usually spend a minute or two carefully inspecting the vegetation at ground level, before entering the undergrowth to oviposit on leaves of Lonicera periclymenum. They meticulously select leaves growing semi-concealed in bushes or on the boughs of trees, always in dappled light or semi-shade. Luxuriant growth, or plants growing in sunlight are ignored.

The egg is globular, greyish-olive, and looks like a miniature sea urchin. The surface is covered with tiny hexagonal cells, and bears many short spines.

The caterpillars hatch after about 10 days. They give away their presence by leaving characteristic feeding marks – eating the tips of the leaves either side of the midrib but leaving the uneaten midrib protruding. The young larva disguises itself by covering its body with pellets of frass ( droppings ), and often rests at the tip of the protruding midvein. This appears to be a defensive action which has evolved to protect it from marauding ants. Many tropical relatives in the Limenitidinae take matters a stage further by constructing ‘frass chains’ by silking together pellets of frass along the protruding midvein. The tiny larvae habitually rest at the extremity, where ants are unable or unwilling to venture, possibly deterred by toxins within the grass.

In September, when in its second instar, the larva constructs a shelter by folding a half eaten leaf, which it fastens together with strands of silk. It spends the winter months hibernating within the shelter, and awakens in the following March to resume feeding, becoming fully grown in late May.

The mature larva is a beautiful creature, bright green with short dark orange spikes along the back and longer spikes on the front segments. It rests in a characteristic posture, with the tail raised and the front segments arched, and is extremely difficult to find in the wild.

The chrysalis is equally exquisite, having a purplish abdomen, bright green wing cases, and metallic silver spots on the back. There is a flattened knob projecting from the back, and a pair of long black labial knobs projecting from the head. It hangs suspended by the tail from a honeysuckle stem, and strongly resembles a withered leaf. The pupal stage lasts for about 2 weeks.

If the weather between late May and early-mid June is warm, the larvae and pupae develop rapidly, but in cool summers the rate of development is much slower. Consequently the time during which they are vulnerable to predation and parasitism is extended, and adult butterflies become scarce.

White Admiral Limenitis camilla, Fermyn Wood, Northamptonshire – Adrian Hoskins

Adult behaviour

The White Admiral is amongst the most graceful of butterflies in flight. Males glide high in the tree tops, then descend, flitting and gliding delicately and with great precision, in and out amongst the foliage in search of females.

On 16th June 2007, I watched 7 males flying around a small oak in a Hampshire wood, and noted that they frequently settled to feed at acorn buds. They also regularly visited the flowers of bramble and dog rose – White Admirals rarely if ever visit any other species of flower.

Both sexes, but particularly the males, also imbibe honey-dew from the upper surface of oak leaves high in the canopy. This sugary fluid is a by-product expelled by the oak aphid Phylloxera quercus, as it sucks protein-rich fluids from oak leaves. Vast quantities of this substance coat the uppersides of oak leaves in mid-summer. Rain washes the honey-dew off of the leaves, so after rainfall White Admirals and other honey-dew feeding butterflies tend to seek alternative forms of sustenance.

White Admiral Limenitis camilla, male, Hampshire – Adrian Hoskins

In hot weather males often descend to imbibe dissolved mineral salts from stony forest tracks, and I have seen them feeding on sticky fluids coating the stems of dock and other plants. I’ve also seen freshly emerged White Admirals drinking at frothy ‘cuckoo-spit’ ( froghopper secretions ) on twigs.

On overcast but warm days, the butterflies often spend long periods basking with wings outspread on the foliage of bramble bushes, bracken or hazel, or on leaf litter on the forest floor. They will fly even during light rain if the weather is warm, but when it gets really hot they seek shade, resting on the upper surface of heavily shaded foliage.

White Admiral Limenitis camilla, male, Hampshire – Adrian Hoskins

Purple Emperor Apatura iris male, Hampshire – Adrian Hoskins

Introduction

The magnificent Purple Emperor is undoubtedly the British butterfly most admired and most sought after by butterfly watchers, breeders, photographers and general naturalists alike. There are more stories told, and more myths about this species than any other.

It is the second-largest species in the UK ( the largest is the Swallowtail ), measuring up to 85mm across the wings. The deep purple-blue sheen on the wings of the male is produced structurally by light refracted by ridges on the wing scales. The colour is only visible from certain angles and under bright lighting conditions.

Less experienced butterfly watchers often mistake the White Admiral for this species. Both species are dark with white banding, both fly around oaks, and both fly at the same time of year. The White Admiral however is smaller, far more graceful and delicate in flight, has much more rounded wings, and lacks the purple sheen. In Europe it can be confused with its slightly smaller relative ilia, but that species has an orange-ringed black spot near the outer margin of the upperside forewing.

The Purple Emperor is distributed throughout much of central Europe but is localised and scarce in southern France, Spain and Portugal. It does not occur in Scandinavia, peninsular Italy or on any of the Mediterranean islands. Beyond Europe its range extends across temperate Asia from the Baltic states to north-east China and Korea.

Purple Emperor Apatura iris male, Hampshire – Adrian Hoskins

Habitats

In Britain the butterfly occurs as a breeding species in most of the larger woodlands in southern and central England, where the larval foodplant Salix grows along the edges of tracks. It once occurred over a wider area but it is now more local. In Wiltshire, Hampshire, Sussex and Surrey it occurs as a breeding species in almost all of the larger woodland complexes. The largest colonies are found in Rockingham Forest in Northamptonshire. A handful of isolated colonies also occur in Warwickshire, Herts, Oxfordshire, Buckinghamshire, Lincolnshire, Somerset and Devon.

Most colonies comprise of less than 20 adults per day at peak season, although in the East Midlands up to 30 can be seen in a single day under ideal conditions in certain forests. Individual butterflies have a range that probably encompasses several hectares, and is usually based on a single area of contiguous forest, with satellite colonies in adjacent smaller woodlands.

Both sexes can be found at least a kilometre away from the egg-laying localities, assembling at so-called master trees, of which there may be several in a large woodland complex. The master trees are usually mature oaks, but beeches, poplars and conifers are also used. In all cases the trees are tall specimens, often located on high ground, typically on the brow of a hill. At these trees the males indulge in spectacular ‘sorties’, competing for the best vantage points. Strings of 4 or 5 males can often be seen chasing in circles around the tree tops.

Females are seen less often. They visit the master trees to find mates, and thereafter disperse to the egg-laying sites, which are often on north facing woodland edges, or in semi-shaded and low-lying areas of the wood where there are high densities of sallow.

Purple Emperor Apatura iris male, Hampshire – Adrian Hoskins

Lifecycle

The conspicuous domed eggs are laid singly on the upperside of sallow leaves. The usual foodplant is broad-leaved sallow Salix caprea but the narrow-leaved S. cinerea is also sometimes used.

In July 2005 I watched a female which in the course of 3 or 4 minutes laid about 10 eggs at various heights between 2 and 5 metres on broad-leaved and narrow-leaved sallows growing on both sides of a track in a Wiltshire wood. The eggs were laid on semi-shaded leaves in the interior of the trees. At other sites in Hampshire and Surrey I have watched females ovipositing on low, sunlit leaves on woodland edge sallows; and at a thicket in north-east Hampshire I have seen a female oviposit at eye-level on a totally shaded broad-leaved sallow.

When first laid the eggs are pale green, but after about 5 days they develop a dark purplish band near the base. They hatch after about 14 days and the newly hatched larva makes it’s first meal of the eggshell.

The young larva is greyish-brown and has 2 prominent horns on its head. It feeds until October and then enters hibernation while in the 2nd or 3rd instar. It spends the winter resting on a silk pad spun on the upperside of a withered sallow leaf, or in the fork of a twig on the damper and shadier north or east facing side of the tree. It resumes feeding in early April.

When fully grown in mid-June the larva is plump and green. It is marked along the sides with dark-edged diagonal cream stripes that perfectly simulate the veins of a leaf. Two horns project forward from the head, and the body is strongly tapered at the tail. It is quite unlike that of any other British butterfly or moth.

The larva rests by daytime along the midrib on the upper surface of a leaf. It periodically nibbles at the tissue either side, leaving the midrib and leaf tip intact. At dusk it vacates it’s resting place and wanders all over the tree to feed. It lays a near-invisible trail of fine silk along its route, and uses this as a map to return to its ‘home’ leaf before dawn.

The chrysalis, which wriggles frantically like a wet fish if touched, is a beautiful shade of translucent silvery green, marked on the abdomen with short whitish diagonal dashes. It is slightly flattened in shape. The camouflage is so perfect that it is virtually impossible to locate, as it hangs suspended by the cremaster from a sallow leaf. The pupal stage lasts for about 14 days.

Adult behaviour

The butterflies begin emerging in late June and early July, although they tend to emerge a week or two later if the early summer has been cool. They remain on the wing until early August.

Purple Emperors spend most of their time resting high in trees, the males favouring oaks while the females more often rest in tall sallows. Females in particular may spend an hour or more on their tree-top perches, especially if the weather is cloudy or breezy. When the sun appears, even if only for a brief period, the males take flight and circle around the vicinity in search of food sources.

Both sexes will feed at aphid secretions which often coat the upper surface of sallow or oak leaves. They are also strongly attracted to sap runs. Males additionally feed at carrion, but are most often encountered when imbibing fluids from carnivore dung or urine-soaked ground. These provide them with essential alkaloids which are passed to females during copulation. Females are sometimes seen imbibing moisture from forest tracks between bouts of egg-laying, but do not visit carrion or dung.

Purple Emperor Apatura iris male, Hampshire – Adrian Hoskins

In July 1984 I found no less than 6 males feeding at the carcass of a roe deer that was floating in an open cesspit in a Hampshire thicket. The butterflies were so stupefied by their unsavoury meal that 2 of them remained on the carcass as I lassoed a rope around the antlers and hauled it to the edge of the cesspit to take photographs !

Mud-puddling and dung-feeding activity typically begins at about 8.30am on warm sunny mornings, or later if it is cooler or slightly overcast, and can continue until 12.30pm or later although there is usually a lull in activity between noon and 2pm. Sometimes a male will spend an hour or more on the ground without moving. Further puddling sometimes occurs in the late afternoon at about 6.00-7.00pm if it remains sunny.

In the early afternoon both sexes fly to the highest point in the forest – the ‘master tree’, typically a tall oak. As many as 6 or 7 may gather there on any particular day. Males usually arrive first, and soon begin to chase each other around in circles, competing to obtain the best ‘throne’ – usually a prominent clump of leaves on which they sit to await the arrival of the females.

When a female appears the dominant male charges after her, followed by any other males at the tree, and the butterflies fly in a string, one behind the other, following the female as she flits and glides around the upper branches. The strongest and most aggressive male eventually chases off his lesser rivals, and then follows the female until she settles on a sunlit clump of leaves, often half a kilometre or more distant from the master tree.

Copulation takes place there in early afternoon. The mating pair spend most of the remaining afternoon sitting on their love nest, but sometimes fly in tandem from one branch to another, or more rarely descend to ground level.

In July 1986 for example, while watching a mud-puddling male in a Surrey wood, I saw a copulated pair float down from an oak, and settle on gravel. When I approached they flew up into another tall oak, settling near the top of the tree. On another occasion I spotted a copulating pair late in the afternoon, at a height of about 6 meters in a spruce. The pair were still copulating at 7.00pm, when I had to leave, and I suspect that they remained joined until dusk, or possibly until the following morning.

In some years prolonged periods of dull or rainy weather occur during the flight season. Then when the sun does briefly appear, a frenzy of courtship, mating and egg-laying takes place. They are able to make the most of very limited opportunities, e.g. weather conditions during throughout the 2007 flight season in the UK were cool, overcast and often wet, with precious little time for mate location or egg laying.

Nevertheless 2008 and 2009 produced the highest Purple Emperor counts on record. Similarly in 2012 Britain experienced atrocious weather, with cool wet and windy conditions lasting from early May until late July. The weather then suddenly changed and became very hot and sunny for about a week, during which time the butterflies emerged en masse, and were able to lay a large number of eggs.

Introduction

The most recent revision by Walberg includes the 6 Aglais species ( Small Tortoiseshells ), plus the Peacock Inachis io, the Blue Admiral Kaniska canace, the Camberwell Beauty N. antiopa and the 5 Polygonia species within the genus Nymphalis which consequently now comprises of 29 species. The genus is very widespread with representatives across the entire Holarctic region. Only one species canace reaches into the Oriental region.

The genus can be recognised by their angular wing-shape and dark undersides which are marked with numerous fine blackish striations. Several of the species carry a white or silvery mark on the underside hindwing, which often appears in the shape of a symbol or letter, hence the Latin species names interrogationalis ( question mark ), comma, g-argenteum, c-album, c-aureum and l-album. The uppersides of the true Nymphalis species ( sensu stricta ) are a rich orange-brown colour, marked with a distinctive pattern of black spots and blotches.

Nymphalis antiopa is distributed across most of Europe but only occurs in Britain as a very scarce migrant. Beyond Europe it is found across temperate Asia, and over most of North America where it is known as the Mourning Cloak. The English name refers to the fact that the first recorded example was captured in Camberwell in London. It was originally known as the Grand Surprise, then went through a succession of other popular names including Willow Beauty and White Petticoat before the current name became widely adopted.

Habitats

Nymphalis antiopa breeds primarily in deciduous woodlands but is migratory in behaviour so can be found in a wide variety of habitats ranging from coastal dunes to dry rocky gullies, alpine meadows and city gardens. It has been recorded at elevations as high as 2300m but is most commonly seen between about 50-1500m.

Lifecycle

The eggs are laid is large batches forming a ring around a twig of the foodplant, usually quite high in the tree. The foodplants include various Salix and Populus species ( Salicaceae ). The larvae feed gregariously and live within a silken web during the early instars, but gradually split into smaller groups as they mature, and live solitarily in the final instar. The chrysalis is formed on the twigs of bushes or on tree trunks.

Adult behaviour

The adults emerge in late summer and spend 2 or 3 weeks feeding at sap runs or fermenting fruit to build up protein reserves prior to hibernating overwinter in hollow trees or log piles. Summer and autumn specimens are a beautiful deep maroon colour, with rich creamy yellow wing margins, but this quickly fades. Post-hibernation examples are duller and invariably have whitish or very pale margins. In spring the adults awaken and nectar at Salix catkins.

Silver-washed Fritillary Argynnis paphia male, Wiltshire, England – Adrian Hoskins

Introduction

The genus Argynnis comprises of about 25 species, found variously in Europe, temperate Asia and North America. Certain workers include an additional 18 Speyeria species within Argynnis.

The Silver-washed Fritillary is one of Europe’s largest and most magnificent butterflies. The common name refers to the suffused silvery markings on the underside hindwings. The male, shown above, is easily distinguished from the female by the 4 prominent horizontal dark streaks on its forewings. These contain androconial scales, from which pheromones are released during the courtship flight.

Silver-washed Fritillary Argynnis paphia female, Wiltshire, England – Adrian Hoskins

Females have a similar pattern of black spots, but lack the horizontal streaks. They are normally a dull golden colour, often with a slight bronzy tinge. The greyish female depicted below is the scarcer and very beautiful form valesina. It only occurs at a few sites, usually in long established colonies in oak forests. Captive breeding experiments have proven that about 10% of females in Hampshire, Wiltshire and Dorset are of the valesina variety. The numbers actually observed in the wild however are much lower as they are more secretive in behaviour than normal females. The common name refers to the suffused silvery markings on the greenish underside hindwings.

Silver-washed Fritillary Argynnis paphia female form valesina – Adrian Hoskins

The butterfly is common and widely distributed across much of Europe, but absent from northern Scandinavia, northern Britain, and the southern parts of the Iberian peninsula. Beyond Europe it occurs in northern Algeria, and is widespread across temperate Asia to China, Korea and Japan.

Argynnis paphia males basking on bracken in early morning – Adrian Hoskins

Habitats

The Silver-washed Fritillary prefers slightly shadier conditions than most other woodland Fritillaries and is better able to survive in high canopy forests and woods that have become neglected and overgrown. The highest populations occur in mature oak and beech plantations where a program of regular thinning encourages a profusion of dog violets to germinate beneath the trees. The best sites are also typified by having glades and grassy tracks where thistles, brambles and other nectar sources are abundant.

The butterfly is a powerful flyer and quite mobile, being often found on scrubby downland or along hedgerows or railway cuttings where these habitats are close to its woodland homelands. Thus it is able to recolonise neglected woods which have been thinned or returned to coppice management and once again become suitable as breeding sites.

In the early 19th century this species was found as far north as Dumfries, but has always been very scarce in northern Britain, and has been extinct in Scotland for about 150 years. There was however a freak migration in 1910, when the species found its way as north as Aberdeen. The butterfly is still very rare north of a line running from Liverpool to Peterborough, but has recently shown signs of expansion, probably as a result of climate change. In 2009 for example I found a fresh male Silver-washed Fritillary visiting bramble blossom in a wood near Witherslack in Cumbria.

Silver-washed Fritillary Argynnis paphia males at bramble, Hampshire – Adrian Hoskins

Lifecycle

The butterflies emerge in June and July and have a life expectancy of about 3-4 weeks, although many are killed by birds before fulfilling their potential life spans.

The greenish-white eggs are laid singly in crevices in the bark of oaks, and more rarely on other trees including beech, ash and larch; normally at a height of about 2 metres, on the moss-covered east or north-facing side of the trees. The trees chosen tend to be about 50-100 year old specimens growing along the edges of rides, and illuminated by dappled sunlight. Often a female will revisit the same favoured tree trunk, laying a dozen or more eggs in the space of an afternoon.

Immediately after hatching in August, the tiny caterpillars eat their egg-shells, which contain vital nutrients. They then attach themselves to a little pad of silk which they spin on the tree trunk, and enter a state of diapause, remaining hidden in a chink in the bark until the arrival of sunny weather in March of the following year. They then descend the trees and wander in search of dog violets Viola riviniana. They feed diurnally, eating only the lobes of the violet leaves, and then move on to another plant. The fully grown larvae can sometimes be found basking on dead oak leaves in May or early June. They are black, marked with a double yellow line along the back, and have a series of dull orange branched spikes on the back and sides. The 2 spikes on the first segment are black, and inclined forward over the head.

The slightly spiky pupa is mottled in shades of brown, and decorated with golden spots. It is almost impossible to find in the wild, but is reportedly suspended by the cremaster from the woody stems of bushes, or from twigs on oak trees. The pupal stage lasts for about 3 weeks.

Silver-washed Fritillary Argynnis paphia, Hampshire – Colin Baker

Adult behaviour

The first males emerge in late June, appearing about a week before the first females. Individuals can live for up to a month, with worn specimens seen as late as early September.

Overnight, and during dull weather, the butterflies roost amongst oak leaves high in the tree tops. In the early morning they bask high up until the rays of the sun reach the forest floor, and then drift down to continue basking on bracken, hazel leaves, and other low vegetation.As the day warms up they become more active, nectaring avidly at thistles, bramble blossom, hemp agrimony, hogweed and ragwort. Both sexes also imbibe honey-dew from the surface of leaves, and males often settle on stony tracks to imbibe mineral-rich moisture.

Silver-washed Fritillary Argynnis paphia males, Wiltshire ï¿½ Adrian Hoskins

The courtship ritual of the Silver-washed Fritillary is one of the most endearing and familiar sights of the English summer. The female flies in a straight line along woodland tracks at a height of about 2 metres, and as she does so she emits an aphrodisiac scent from the tip of her abdomen. The male responds by following her closely, repeatedly looping under and over her, and showering her with pheromones released from the 4 black bars of androconial scales which run along the veins of his forewings. In many cases this tantalising display fails to entice the female into mating, but if she is receptive she leads the male to a clump of leaves high in an oak tree where copulation takes place. Periodically the pair fly down to settle on bracken or hazel, or to nectar at bramble, but return to the tree tops if disturbed. Copulation lasts about 2 hours and usually takes place in late morning.

Silver-washed Fritillary Argynnis paphia female, Straits Inclosure, Hampshire – Adrian Hoskins

Silver-washed Fritillary Argynnis paphia male, Bentley Wood, Wiltshire – Adrian Hoskins

Introduction

The genus Argynnis comprises of about 25 species, found variously in Europe, temperate Asia and North America. Certain workers include an additional 18 Speyeria species within Argynnis.

The High Brown Fritillary Argynnis adippe is distributed across much of Europe, but is absent from northern Scandinavia, and the Mediterranean islands ( except Sicily ).

It can easily be mistaken for the Dark Green Fritillary, which often shares its habitats, but the male of aglaia lacks the conspicuous black sex brands which run along veins 2 and 3 of the forewing of adippe. The ground colour of female High Brown Fritillaries is golden, whereas in females of aglaia it is more reddish brown with noticeably pale outer margins. In both sexes the outer edge of the fore-wings is almost straight in aglaia, but is slightly concave in adippe. The size and positioning of the black spots is variable in both species. On the undersides the differences are more pronounced – the High Brown Fritillary has a yellowish ground colour with no trace of green, and has an additional row of white-centred reddish spots in the submarginal area of the hindwings.

In continental Europe this species can be confused with the Niobe Fritillary Argynnis niobe, but the latter is smaller and has more chevron-like lunules on the underside.

Argynnis adippe male, Arnside Knott, Cumbria – Adrian Hoskins

Habitats

The High Brown Fritillary was formerly very widespread in Britain, and occurred commonly in most of the larger woodlands in southern England and Wales, although it became scarcer in the northern counties. During the mid 20th century the butterfly declined very rapidly as a result of habitat loss. The coppiced woodlands where it once bred became neglected and overgrown, or were converted to plantations of oak, beech or conifers. Consequently the larval foodplants were shaded out and the species was lost.

This is one of the most rapidly declining species in Britain. It has proven incapable of surviving at the small number of isolated sites which are still actively coppiced. The butterfly’s long term viability depends on the existence of extensive tracts of well drained and lightly wooded habitat where the larval foodplant, dog violet Viola riviniana grows profusely and in continual supply. These habitats are now very scarce, so most of the woodland colonies have long disappeared, and the butterfly is now largely restricted to a few small and isolated areas of scrubby grassland / woodland mosaics on limestone hills, where periodic cattle grazing, bracken control, and retention of scrub are vital elements in ensuring it’s continued survival.

In Europe it is much more catholic in it’s choice of habitats, occuring on scrubby grassland, sparsely vegetated limestone plateaux, and in sub-alpine meadows and woodland / grassland mosaics. It’s choice of larval foodplants is also wider and includes Viola hirta, V. riviniana and V. odorata.

Argynnis adippe male, Arnside Knott, Cumbria – Adrian Hoskins

As well as the normal form, adippe produces a variety cleodoxa in which the silvery spots on the underside are replaced with yellow.

Argynnis adippe, male form cleodoxa, Aggtelek, Hungary – Peter Bruce-Jones

Lifecycle

The butterflies emerge in late June and early July. They lay their pinkish conical eggs singly on dead bracken fronds, dead leaves, dry stems or bits of moss, very close to the larval foodplants. They are normally deposited in sheltered nooks, in the dappled sunlight beneath bracken, gorse or straggling patches of low bramble growth. The tiny caterpillars quickly develop within the eggs, but don’t hatch until March of the following year.

The fully grown larva is dark brown, with a white stripe along the back, and is covered with rows of brown or pinkish spikes along the back and sides. It lives solitarily, wandering from one violet plant to another, and feeds diurnally, periodically retiring to rest beneath dead leaves.

The dark brown, shiny chrysalis resembles a withered leaf, and is virtually impossible to locate in the wild. In captivity it is formed attached by the cremaster to a silk pad spun on twigs or stems, and in the wild it is probably suspended from twigs or woody stems beneath bushes. The pupal stage lasts for about 3 weeks.

Argynnis adippe male, Arnside Knott, Cumbria – Adrian Hoskins

Adult behaviour

On cool or slightly overcast days the butterflies bask for long periods on the ground, choosing small depressions with dead vegetation, where they are sheltered from the wind. If disturbed they tend to only fly short distances and quickly settle in another nearby depression.

On warm sunny days males patrol rapidly across the habitat in search of females, pausing every now and then to nectar at thistle, bramble, heather, knapweed, burdock, hemp agrimony and other flowers. Females use the same nectar sources, but alternate between nectaring and ovipositing, typically dropping down to lay an egg at each of about half a dozen spots within a small area, and then spending 2 or 3 minutes nectaring at nearby flowers before repeating the process.

In late afternoon the butterflies tend to bask on bracken. Both sexes roost overnight in trees, on leaf clumps at the tips of the highest branches. They also fly up to settle in similar positions during overcast or wet weather.