Behaviour

The females often stay hidden Roer, 1970 - Untersuchungen zum Migrationsverhalten des Trauermantels (Nymphalis antiopa L.) (Lep., Nymphalidae) or fly around without being tied to a territory. It shows a male that it is ready to copulate by fluttering its wings briefly and intensively Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter I.

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The usual time of day for mating is midday or early afternoon Schricker, 1988 - Beobachtung der Balz und Paarung bei Nymphalis antiopa (LINNAEUS, 1758) im Freiland Roer, 1970 - Untersuchungen zum Migrationsverhalten des Trauermantels (Nymphalis antiopa L.) (Lep., Nymphalidae).

One interesting story tells of a secret string that the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is said to pluck during mating flight Schnack, 1947 - Das Leben der Schmetterlinge - Naturdichtung:

When the mourning cloak courts a female, it is said, as Marielle told me, to play soft love and wedding music during its courtship flight, as if it were the spirit of an enchanted musician. It is said to have a secret string on the rear edge of its front wing, a vein that is rubbed against one of the rear wings. A delicate sound is produced, which the chosen one must hear! And perhaps the wooer's way in butterfly language is: Mate of summer, how beautiful you are in your new black coat! Bask with me for a while in the sun!. And she, the beautiful black friend, does not deny herself to him. He has played too finely.

The Mating; sexual act in animalsCopulation Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter I, which takes place on the ground, lasts about 90 minutes Schricker, 1988 - Beobachtung der Balz und Paarung bei Nymphalis antiopa (LINNAEUS, 1758) im Freiland and soon afterwards the female begins to lay her eggs. Egg laying can take place in several batches. Not much later the female dies.

Do males live longer than females? Do they seek out several mating partners?

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Beatrice LaporteBeatrice Laporte discovered two Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties mating upside down under a dead tree trunk in June 2013.

Male and female adult butterflies find each other visually. It is assumed that a male only recognises a female when flying in circles Hesselbarth, 1995 - Die Tagfalter der Türkei unter Berücksichtigung der angrenzenden Länder.

At a daylight length of more than 16 hours, sufficient Juvenile hormones, important for the production of eggs, among other things.juvenile hormons are produced by the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty so that Mating; sexual act in animalsCopulation and Egg deposition; egg layingOviposition are possible Herman; Bennett, 1975 - Regulation, female specific protein production and male and female reproductive gland development by juvenile hormone in the butterfly Nymphalis antiopa.

If the females often remain hidden during the mating season, how do they find a mate? Presumption: As they have no territorial ties, they are immediately identified by males lying in wait when they enter male territories during their rare flights.

Males and females look very similar. How do they recognise the opposite sex? Guess: By the way they fly. Perhaps Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties are also able to see ultraviolet colours in a similar way to other animals? The wings of female or male adult butterflies could differ in ultraviolet light. See Spektrum der Wissenschaft 01/2007: Birds see the world in more colourful ways.

Is the process of finding a mate between male and female supported by pheromones? Presumption: Probably not, as this is common in moths but extremely rare in butterflies.

Are there typical mating sites?

What is the ratio of male and female adult butterflies?

The males of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty show pronounced territorial behaviour during the mating season. Territories of over 300 square metres are sometimes defended aggressively and fearlessly. I was also able to establish this in my few personal observations of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty, see MySightings.

If a male butterfly encounters a competitor, the two Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties fly upwards in vertical spirals, performing a 10 to 14 metre high circular flight. The intruder then usually leaves the area Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter IHesselbarth, 1995 - Die Tagfalter der Türkei unter Berücksichtigung der angrenzenden Länder.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty can also make clicking noises when attacking Fonseca, 2025 - Nymphalis antiopa.

Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties fly directly towards intruders, e.g. other adult butterflies, insects or birds Fonseca, 2025 - Nymphalis antiopa. Occasionally humans are also attacked Schricker, 1988 - Beobachtung der Balz und Paarung bei Nymphalis antiopa (LINNAEUS, 1758) im Freiland. There are reports of Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties that have also pursued, attacked and finally driven bats out of their own territory Nielsen, 1998 - Mourning Cloak Antics. One Big Brown BatEptesicus fuscusBig Brown Bat had to deal with two Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties Anich; White; Anich, 2020 - Interspecific Territorial Behavior of Two Mourning Cloaks (Nymphalis antiopa) in Response to a Diurnally Active Big Brown Bat (Eptesicus fuscus).

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is active during the day, but in rare cases can also be on the move at night Williamson, 2002 - Nature Trails: A bashful beauty.

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Robert PetscharRobert Petschar lured an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty with a UV lamp at around 21:00 in early September.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty likes to suck wood sap from injured or felled trees, preferably oaks and birches, with its proboscis. This energy-rich food enables quick energy absorption, which is particularly advantageous in spring. It usually sucks sitting on the bark upside down, i.e. with its head facing the ground.

In the United StatesUnited States, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty has been observed sucking the sap oozing from maple trees where the bark has been gnawed by squirrels. It also sometimes sucks aphid honeydew from sticky leaves of infested plants. In April and May, the adult butterflies of United StatesUnited States also visit the flowers of MoosewoodAcer pensylvanicumMoose MapleMoosewoods and later those of Common MilkweedAsclepias syriacaCommon Milkweeds Weed, 1917 - Butterflies worth knowing.

After hibernation, flowering willows usually serve as the first food source, followed by sticky buds of stone fruit blossoms Brahm, 1791 - Handbuch der ökonomischen Insektengeschichte. But they also visit flowering plants (in residential areas in summer often on Orange Eye ButterflybushBuddleja davidiiButterfly BushOrange Eye Butterflybush flowers). Wet soil occasionally serves as a water source Weidemann, 1988 - Naturführer Tagfalter Band 2 - Biologie-Ökologie-Biotopschutz. In autumn, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty can be found sucking on overripe fruit.

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In the studies carried out by Allen M. YoungAllen M. Young in Wisonsin, United StatesWisonsin Young, 1980 - Some observations on the natural history and behavior of the Camberwell Beauty (Mourning Cloak) butterfly, Nymphalis antiopa (Linnaeus) (Lepidoptera; Nymphalidae) in the United States, an interesting observation was made in connection with the sap flow of injured trees:

In the morning hours, Yellow-Bellied SapsuckerSphyrapicus variusYellow-Bellied Sapsuckers regularly visit the same trees to work them with their beaks. The sap flow triggered by this is used by the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty as a primary carbohydrate-rich food source. As the sap dries quickly and hardens, the source dries up at midday.

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Like some other adult butterflies, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty sucks on sweat (see Behaviour (Confidingness)), and it also sucks on carrion Gu, 2014 - Animal biodiversity and food web restoration based on large vertebrate carcasses.

In the United Kingdom in particular, there are accounts that the Camberwell Beauty can be attracted and caught at night by strips of paper soaked in syrup or the sap of injured oak trees Williamson, 2002 - Nature Trails: A bashful beauty.

The adults can also be attracted by pieces of fermented pineapple in cheesecloth bags Young, 1980 - Some observations on the natural history and behavior of the Camberwell Beauty (Mourning Cloak) butterfly, Nymphalis antiopa (Linnaeus) (Lepidoptera; Nymphalidae) in the United States. Another good bait is a one-day fermented mixture of mashed overripe bananas, crushed tinned pineapple, sugar and beer Chermock, 1952 - The Use of Bait to Attract Butterflies.

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George Howard Parker (1864-1955)George Howard Parker discovered during his intensive observations at the beginning of the twentieth century in New England, United StatesNew England that the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty spends cold spring nights in places analogous to hibernation sites Parker, 1903 - The phototropism of the mourning-cloak butterfly, Vanessa antiopa Linn.. Driven by falling temperatures, the adult butterflies in an area visit these overnight roosts almost simultaneously (at the beginning of April this was between 04:00 PM and 04:15 PM).

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The population density of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty's is statistically between one adult butterfly per hectare and six adult butterflies per square kilometre Settele, 1999 - Die Tagfalter Deutschlands.

Population sizes are subject to more or less cyclical fluctuations. Population explosions can occur in some areas, while populations in other areas can decline dramatically or even disappear completely.

Is the distribution kumularaggregiert, uniform or random? Assumption: kumularaggregiert, i.e. local clusters of adult butterflies.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty likes to move at greater heights, especially in the canopy of trees or bushes. As a result, it often eludes observation.

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The adult butterfly is a fast and elegant flyer. Similar to the MonarchDanaus plexippus (Linnaeus, 1758)Monarch ButterflyMonarch, it holds its wings slightly above the horizontal when gliding. During leisurely flight, the wings beat at an angle of about 45º, during fast flight it becomes about 150º, i.e. they almost beat together Scott, 1986 - The Butterflies of North America - A Natural History and Field Guide.

If an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is attacked in flight, it can drop lifelessly to the ground and play dead Cech; Tudor, 2005 - Butterflies of the East Coast: An Observer's Guide.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is a migratory butterfly.

In contrast to a typical migrant such as the Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady, which migrates to fixed destinations every year, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is considered a species that migrates primarily for the purpose of range expansion Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter I.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty therefore also exhibits ground-dwelling behaviour and in this way maintains local populations. Part of a population migrates and thus ensures the geographical spread of the species. This form of migration is advantageous, for example, in situations where the conditions in a habitat jeopardise the population, e.g. a winter that is too warm, in which overwintering adult butterflies die.

Some classifications of migratory behaviour from the literature:

• vagabonding Settele, 1999 - Die Tagfalter Deutschlands
• distinctive migratory butterfly Tolman; Lewington, 1998 - Die Tagfalter Europas und Nordwestafrikas
• suspected migrant, emigrant/internal migrant Eitschberger, 1991 - Wanderfalter in Europa (Lepidoptera)
• non-migratory species, but individual adult butterflies migrate (North America) Fonseca, 2025 - Nymphalis antiopa

I do not know how high the proportion of migrating adult butterflies is, but it could be that it depends dynamically, for example, on the existing population density (population too high or too low) or the environmental conditions such as the available food supply.

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Migrating adult butterflies usually migrate immediately after hatching and thus before summer diapausesummer rest Tolman; Lewington, 1998 - Die Tagfalter Europas und Nordwestafrikas Roer, 1970 - Untersuchungen zum Migrationsverhalten des Trauermantels (Nymphalis antiopa L.) (Lep., Nymphalidae).

The adult butterflies migrate individually or in small groups Tolman; Lewington, 1998 - Die Tagfalter Europas und Nordwestafrikas.

The direction and distance of migration depends to a large extent on the prevailing wind directions and strengths. This so-called co-wind migration can provide an explanation for the spread or dispersal of the butterfly Roer, 1970 - Untersuchungen zum Migrationsverhalten des Trauermantels (Nymphalis antiopa L.) (Lep., Nymphalidae).

Occasionally, mass migrations also occur. A large number of adult butterflies then move in a similar direction and over a similar distance within a narrow time corridor.

One example is the rare waves of immigration to the British Isles (see Distribution in United Kingdom). In North America, there were mass movements in South-Texas, United StatesTexas, South-Florida, United StatesFlorida and Bermuda, United StatesBermuda Scott, 1986 - The Butterflies of North America - A Natural History and Field Guide.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty can be quite trusting. It happens often that an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty settles on a person, even lingers there for a few minutes and often sucks sweat. Even movements by the person do not drive the adult butterfly away (an experience I have had myself with an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty).

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Just a few days after hatching, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty takes a summer diapausesummer rest lasting several weeks, which it spends in houses or sheltered bushes, for example. The adult butterfly find other hiding places under trunks or loose bark of dead trees Weed, 1917 - Butterflies worth knowing.

A remarkable characteristic of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is that it falls into a kind of dormant state. In this torpor, the wings are folded together, the antennae are tucked back between the wings and the legs are pulled up against the body. The adult butterfly usually sits with its wings pointing towards the ground Weed, 1917 - Butterflies worth knowing.

This rigidity is also adopted during winter resthibernation and can also be observed in newly hatched adult butterflies if they are disturbed Weed, 1917 - Butterflies worth knowing. The latter can certainly be seen as a form of playing dead.

The summer diapausesummer rest ends in early autumn when the days get colder Weed, 1917 - Butterflies worth knowing.

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In North America, the location of summer diapausesummer rest can be up to 70 km away from the hatching site Cech; Tudor, 2005 - Butterflies of the East Coast: An Observer's Guide.

Henry Gardiner Adams (1811-1881)Henry Gardiner Adams reports from Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties on the summit of Drachenfels, North Rhine-Westphalia, GermanyDrachenfels near Bonn, GermanyBonn Adams, 1854 - Beautiful Butterflies: The British Species described and illustrated with an introductory chapter containing the History of a Butterfly through all its changes and transformations. A strong wind was blowing and the adult butterflies were sitting on the bark on the side of the trees facing away from the wind and had fallen into a state of torpor. When they were thrown into the air, they barely opened their wings or fluttered weakly to the next place with protection from the wind (see full translation: Stories/Poems (Henry Gardiner Adams: Beautiful Butterflies: Camberwell Beauty (1854))).

In addition to summer diapausesummer rest, the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty as a adult butterfly also keeps a winter resthibernation.

Hibernation begins in autumn and ends in GermanyGermany around the end of March, but leaving the hibernation site also depends on the altitude: At higher altitudes, hibernation ends later than at lower altitudes.

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The adult butterfly uses these as hibernation sites:

• Tree hollows or hollow bark Rösel, 1746 - Insecten-Belustigung erster Theil. Der Tagvögel erste Classe. N. 1. Die grosse gesellige Dornen-Raupe mit gelb-rothen Flecken, und ihre Verwandlung bis zum Papilion, especially near the roots Scudder, 1899 - Every-day Butterflies: A Group of Biographies
• Hwood piles Scudder, 1899 - Every-day Butterflies: A Group of Biographies
• Piles of brushwood
• Loose bark Weed, 1917 - Butterflies worth knowing
• Sheds Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter I oder Scheunen Scudder, 1899 - Every-day Butterflies: A Group of Biographies
• Buildings Roer, 1970 - Untersuchungen zum Migrationsverhalten des Trauermantels (Nymphalis antiopa L.) (Lep., Nymphalidae) und deren Keller Scudder, 1899 - Every-day Butterflies: A Group of Biographies
• Bridges Weed, 1917 - Butterflies worth knowing
• Open canalisations Weed, 1917 - Butterflies worth knowing, Scudder, 1899 - Every-day Butterflies: A Group of Biographies
• Rock crevices Weed, 1917 - Butterflies worth knowing
• Stone piles Scudder, 1899 - Every-day Butterflies: A Group of Biographies

Although the adult butterfly is comparatively insensitive to low temperatures, ice crystals in its body pose a deadly danger to it. It protects itself by ingesting natural antifreeze such as Sorbitol (also known as glucitol or hexanhexol) belongs to the alditols (sugar alcohols) and is the reduced polyol form of glucose.Sorbitol, which it only produces when the weather gets colder. In summer, the butterflies lacks these means, so that butterflies die immediately in very cold summer temperatures. As frost protection is only effective down to -38° C (source: Joseph BelicekJoseph Belicek, pers. comm., 2009), winters that are too cold can lead to the death of the adult butterfly Taron, 2002 - Mourning Cloak: Meet Your Neighbours.

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The two previous photos by Libor DvořákLibor Dvořák show the hibernation site of an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty in the Czech Šumava National Park, Bohemia, CzechiaŠumava National Park. The hibernating Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty was under a piece of bark lying on a piece of meadow.

Samuel Hubbard ScudderSamuel Hubbard Scudder noticed that an Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty hibernating in the cellar of his house had moved within a few centimetres a few times during hibernation Scudder, 1899 - Every-day Butterflies: A Group of Biographies. I personally made the same observation when winterising a [#io|case=Genitiv#] 2024/2025 in the basement of my house.

Kenelm W. PhilipKenelm W. Philip placed individual Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties for overwintering in Fairbanks, Alaska, United StatesFairbanks in openings that he had drilled into a piece of wood. He closed the two open sides of the opening with fabric so that the butterflies were protected. He placed the piece of wood on the ground where, covered in snow, the adult butterfly successfully overwintered (Source: Joseph Belicek, pers. comm., 2009).

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Generally speaking, cold winters without major temperature fluctuations are ideal overwintering conditions for the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty. If the winter is too warm, individual butterflies may leave the hibernation site early, which can be fatal for the butterflies.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty leaves its hibernation site as early as the first warm days of spring (around the flowering time of ColtsfootTussilago farfaraColtsfoot) Weidemann, 1988 - Naturführer Tagfalter Band 2 - Biologie-Ökologie-Biotopschutz. In some areas with long periods of snow well into spring, it is not unusual to see Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties flying around despite the snow still lying on the ground.

Kind of hibernation/overwintering of the 143 butterflies distributed in GermanyGermany (source: Settele, 1999 - Die Tagfalter Deutschlands):

Hibernation	Species		migration
as egg	24	(16,8%)	0	(0%)
as caterpillar	92	(64,3%)	10	(15,6%)
as chrysalis	18	(12,6%)	7	(38,9%)
as adult butterfly/imago	6	(4,2%)	6	(100%)
moveing to warmer areas	3	(2,1%)	3	(100%)

• None of the butterflies that overwinter as eggs migrate.
• One sixth of the butterflies that overwinter as caterpillars migrate.
• Approximately one third of the butterflies overwintering as chrysalis (38.9 %).
• All species overwintering as butterfly in GermanyGermany migrate.

Jürgen HensleJürgen Hensle pointed out to me that for Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow, Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady and Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral the type and manner of overwintering cannot be answered across the board and simply with as a caterpillar or as a imago. In addition, the behaviour of the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral in particular has changed in recent years or decades, which means that the behaviour patterns given in the literature do not always describe the current status (Source: Jürgen Hensle, pers. comm., 2006).

His notices are interesting and sometimes a little surprising:

Hibernation

Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow, Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady and Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral generally overwinter without Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause. The Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow and the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral form an uninterrupted succession of generations in the warmest areas. The Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady always has an interrupted succession of generations.

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With late oviposition (November) and low temperatures (Central Europe), the eggs of Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow and Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral can at least partially overwinter during the first half of winter. However, the egg does not overwinter in the classic sense (inactive), but the caterpillar develops in the egg during the winter and hatches in January. If temperatures are higher after egg-laying, development is faster and the caterpillar hatches earlier, which can lead to the egg phase being completed before hibernation. Once the caterpillar has hatched, it remains active and feeds. Frost phases are survived inactive but without Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause (The caterpillar of Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow can very occasionally lapse into an approx. six-week short-term Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause during the coldest weeks of the year). If temperatures rise above 0° Celsius again, the caterpillar continues to feed.

The Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow overwinters, usually as a caterpillar, in the warmest regions (e.g. Canary Islands or CyprusCyprus) in an uninterrupted succession of generations. The caterpillar of Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow tolerates frost without any problems, but the host plant does not: it freezes back in longer periods of frost. If it is cold for too long, if there is snow on the leaves of the host plant or if the leaves freeze, the caterpillar starves to death (exception: isolated caterpillars in short-term Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause).

If the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral overwinters, then as an egg/caterpillar (see above) or as a adult butterfly. The caterpillar of the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral is sensitive to cold to varying degrees depending on its origin. Southern European Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral caterpillars tolerate almost no frost. Individual populations in central and south-eastern Europe are frost-hardy. As the food plant of the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral's freezes back during longer periods of frost, the caterpillar of the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral's can also starve to death - as with the Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow.

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If the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral hibernates as a imago, it rests - but without Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause. If it gets warm for a short time in winter, it wakes up again immediately and becomes active, even in December and January. The adult butterfly currently tolerates between seven and fourteen days of permafrost in Central Europe and can withstand temperatures down to at least -20° Celsius. As it does not Hormone-controlled dormancy of insects and other invertebrates, especially during prolonged cold or dry periodsdiapause, long winters are problematic for it.

The Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady does not have a stage suitable for overwintering.

Hibernation areas

The Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow hibernates or attempts to hibernate in warmer areas of western Europe (south coast of EnglandEngland, West-FranceFrance), central Europe (catchment area of the Rhine and middle Danube) and southern Europe in general.

The Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral has now become native to GermanyGermany. The majority of Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admirals in Central Europe also overwinters in Central Europe - mainly as adult butterflies. Some of the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admirals migrate south to the Po Valley and Burgundy in autumn. Today, hardly any Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral migrate from the Mediterranean region to Central Europe. The Mediterranean region is a completely independent migratory area that is closed off to Central Europe.

Adult butterflies overwintering in Central Europe are mainly found in the catchment area of the Rhine Which I can confirm from my own observations in my home town of Bonn, GermanyBonn and in northern GermanyGermany, east to Rügen. But also in inland eastern Germany and Bavaria. In winter, it is only absent in the middle and higher altitudes of the low mountain ranges, above approx. 500 metres and perhaps in Lusatia.

As the Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady does not have a stage suitable for hibernation, it leads a vagabond's life: it is constantly on the move. The western Palaearctic populations of the Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady spend most of the winter in Africa, sometimes also in southernmost southern Europe.

Generations

The Clouded YellowColias croceus (Geoffroy, 1785)Colias croceaColias edusaClouded Yellow and the Red AdmiralVanessa atalanta (Linnaeus, 1758)Pyrameis atalantaRed AdmirableRed Admiral form three to four generations in GermanyGermany and Central Europe respectively. In warmer areas, such as CzechiaCzechia, there can be five to six generations.

The Painted LadyVanessa cardui (Linnaeus, 1758)Cynthia carduiCosmopolitanPainted Lady forms one to two generations in GermanyGermany or Central Europe. Throughout the year, including its winter flight areas and the spring and autumn inter-settlement areas, there are six generations.

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is a r-strategists rely on a high reproduction rate (=r) when colonising a habitatr-strategy, i.e. it pursues a propagation strategy.

In contrast, a K-strategists reproduce according to the capacity (=K) of the ecological framework conditionsK-strategy pursues an adaptation strategy.

A typical r-strategists rely on a high reproduction rate (=r) when colonising a habitatr-strategy is, for example, the Large WhitePieris brassicae (Linnaeus, 1758)Cabbage ButterflyCabbage WhiteLarge Cabbage WhiteLarge White Settele, 1999 - Die Tagfalter Deutschlands.

One of the much more numerous K-strategists reproduce according to the capacity (=K) of the ecological framework conditionsK-strategy in GermanyGermany is, for example, the ApolloParnassius apollo (Linnaeus, 1758)Mountain ApolloApollo Settele, 1999 - Die Tagfalter Deutschlands.

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Egg

Clarence Moores Weed (1864-1947)Clarence Moores Weed observed in the early twentieth century how a Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty female laid a group of eggs on a branch of a Willow SpeciesSalix spec.Willow Species Weed, 1917 - Butterflies worth knowing. Clarence Moores Weed (1864-1947)Weed noticed that immediately after the adult butterflies had completed its work and flown away, a tiny fly ran excitedly over the newly laid eggs (the fly was later identified by Leland Ossian Howard (1857-1950)Leland Ossian Howard as the Telenomus graptaeTelenomus graptae). In most eggs, the tiny fly laid its own, much smaller egg with its ovipositor. Each egg took between one and two minutes to lay. It walked around for a few seconds between laying two eggs.

It is noteworthy in this context that the small fly was immediately on the spot to lay its eggs. This phenomenon has also been observed by other people Weed, 1917 - Butterflies worth knowing.

A small larva soon hatched from the tiny egg, which developed inside the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty egg and eventually pupated in the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty egg. The hatching fly finally left the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty egg through a hole eaten into the egg shell.

Caterpillar

The caterpillars of Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty, like the caterpillars of many other butterflies, are infested by a large number of Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids. The Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids can decimate their populations, in some cases severely.

The following families of Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids infest the caterpillars:

1. Chalcid waspChalcidoideaChalcid wasps (Chalcid waspChalcidoideaChalcidoidea)
2. Ichneumon waspIchneumonidaeIchneumon wasps (Ichneumon waspIchneumonidaeIchneumonidae)
3. Tachinide FlyTachinidaeTachinide Flies (Tachinide FlyTachinidaeTachinidae)
4. ScelionidaeScelionidae (ScelionidaeScelionidae)

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The tiny four-winged Chalcid waspChalcidoideaChalcid wasps lay their eggs in the caterpillars of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty. There, the larvae continue to develop and eventually kill the caterpillar. After pupation and hatching, it then goes in search of a host. Up to 145 Chalcid waspChalcidoideaChalcid wasps can leave an infested dead caterpillar Weed, 1917 - Butterflies worth knowing.

In contrast, the much larger Ichneumon waspIchneumonidaeIchneumon wasps only develop in one or two specimens in a caterpillar of Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty Weed, 1917 - Butterflies worth knowing.

Tachinide FlyTachinidaeTachinide Flies lay their eggs on the skin of the caterpillars. When the larva hatches, it eats through the skin of the egg and the skin of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty caterpillar into the body of the caterpillar, where it feeds on the inside of the caterpillar. The Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoid continues to weaken the caterpillar and it becomes increasingly sluggish. At this point, infestation with a Tachinide FlyTachinidaeTachinide Fly can also be recognised by the fact that infested caterpillars do not crawl away like the other caterpillars when disturbed. Once the larva of the Tachinide FlyTachinidaeTachinide Fly is fully grown, it breaks through the skin of the dying caterpillar to the outside, where it drops to the ground and pupates. In this case, pupation means that the skin of the larva becomes brown and harder. Finally, after one to two weeks, the Tachinide FlyTachinidaeTachinide Fly Weed, 1917 - Butterflies worth knowing hatches.

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Ichneumon waspIchneumonidaeIchneumon wasps Friedrich, 1986 - Breeding Butterflies And Moths and Tachinide FlyTachinidaeTachinide Flies Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae) are the largest plague in Europe. Clarence Moores Weed (1864-1947)Clarence Moores Weed reports that in the early twentieth century in New Hampshire (United StatesUnited States) Tachinide FlyTachinidaeTachinide Flies were the most common Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty-Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoid Weed, 1917 - Butterflies worth knowing.

The following table shows some known Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty:

Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoid	Family [Ordnung]	Region	Source
Ichneumon fossoriusIchneumon fossorius	Ichneumon waspIchneumonidaeIchneumon wasp [HymenopteraHymenoptera]	Europe	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Hoplismenus terrificusHoplismenus terrificus	Ichneumon waspIchneumonidaeIchneumon wasp [HymenopteraHymenoptera]	Europe	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Hoplismenus morulusHoplismenus morulus	Ichneumon waspIchneumonidaeIchneumon wasp [HymenopteraHymenoptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Thyrateles procaxThyrateles procax	Ichneumon waspIchneumonidaeIchneumon wasp [HymenopteraHymenoptera]	North-America	Andrew M. R. BennettAndrew M. R. Bennett, 2013 with reference to (Bradley, 1978)
Pteromalus puparumPteromalus puparum	Chalcid waspChalcidoideaChalcid wasp [HymenopteraHymenoptera]	Europe	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Pteromalus vanessaePteromalus vanessae	Chalcid waspChalcidoideaChalcid wasp [HymenopteraHymenoptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Pteromalus puparumPteromalus puparum	Chalcid waspChalcidoideaChalcid wasp [HymenopteraHymenoptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Derostenus antiopaeDerostenus antiopae	Chalcid waspChalcidoideaChalcid wasp [HymenopteraHymenoptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Telenomus graptaeTelenomus graptae	ScelionidaeScelionidae [HymenopteraHymenoptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Telenomus kolbeiTelenomus kolbei	ScelionidaeScelionidae [HymenopteraHymenoptera]	FranceFrance	Kan; Shaw; Kan - van Limburg Stirum; Warot; Groussie, 2021 - First record of Telenomus kolbei (Hymenoptera, Scelionidae) in France, parasitizing the eggs of Nymphalis antiopa and Aglais io (Lepidoptera, Nymphalidae)
Phorocera edwardsiiPhorocera edwardsii	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	New England, United StatesNew England (United StatesUnited States)	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Phorocera concinnataPhorocera concinnata	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	Europe	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Phorocera antiopaePhorocera antiopae	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	Europe	Scudder, 1889 - The Butterflies of the Eastern United States and Canada with special Reference to New England, Vol. III
Exorista larvarumExorista larvarum L.Exorista larvarum	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	North-GreeceGreece	Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae)
Phorocera concinnataPhorocera concinnata	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	North-GreeceGreece	Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae)
Sturmia bellaSturmia bella Meig.Sturmia bella	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	North-GreeceGreece	Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae)
Masicera cuculliaeMasicera cuculliae R.D.Masicera cuculliae	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]	North-GreeceGreece	Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae)
Pelatachina tibialisPelatachina tibialis	Tachinide FlyTachinidaeTachinide Fly [DipteraDiptera]		Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae) with reference to Herting, 1960 - Biologie der westpaläarktischen Raupenfliegen Dipt., Tachinidae

In general, populations in Montains (Alps: 800-1850m altitude, Low mountain range 450-1500m altitude)montane areas seem to be less infested by Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids Friedrich, 1986 - Breeding Butterflies And Moths.

The degree of parasitisation of the caterpillars may also be responsible for population fluctuations of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty Ebert; Rennwald, 1991 - Die Schmetterlinge Baden-Württembergs. Band 1, Tagfalter I.

In North America, caterpillars of Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty are eaten on the way to pupation (but also the chrysalis) by squirrels and chipmunks (Allen M. YoungAllen M. Young Young, 1980 - Some observations on the natural history and behavior of the Camberwell Beauty (Mourning Cloak) butterfly, Nymphalis antiopa (Linnaeus) (Lepidoptera; Nymphalidae) in the United States with reference to L. TevisL. Tevis stomach content studies of the two rodents conducted in Northeast-CaliforniaCalifornia in 1953).

A large American beetle, the Caterpillar HunterCalosoma scrutatorCaterpillar Hunter - also known as the Caterpillar Hunter (German: Raupenjäger) - eats the caterpillars of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty and can destroy a large number of caterpillars Weed, 1917 - Butterflies worth knowing.

In den südlichen Staaten der United StatesUnited States wurden Polistin waspPolistesPolistin wasp beim Angriff auf Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty-Raupen beobachtet und es gibt wahrscheinlich viele weitere Insekten, deren Nahrung auch Raupen des Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty sind Weed, 1917 - Butterflies worth knowing.

As the caterpillars of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty are very prickly, there are comparatively few bird species that prey on the caterpillars. For North GreeceGreece the Black-billed MagpiePica pica L.Black-billed Magpie and for Bonn, GermanyBonn the House SparrowPasser domesticus L.House Sparrow are named Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae), for North America the: Yellow-billed CuckooCoccyzus americanusYellow-billed Cuckoo, the Black-billed CuckooCoccyzus erythropthalmusBlack-billed Cuckoo and the Baltimore OrioleIcterus galbulaBaltimore Oriole Weed, 1917 - Butterflies worth knowing.

After all, they also seem to be on the menu of toads in North America Weed, 1917 - Butterflies worth knowing.

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With its red spots on a black background, the caterpillar of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty's signals to potential predators that it will be bad for them. This effect is reinforced by the caterpillar's sociability. In addition, the strong spines carried in later stages of development offer protection.

If a caterpillar is disturbed, it flinches in defence Fonseca, 2025 - Nymphalis antiopa, or all the caterpillars in a community move their front bodies back and forth in the same rhythm Hesselbarth, 1995 - Die Tagfalter der Türkei unter Berücksichtigung der angrenzenden Länder.

Chrysalis

If the chrysalis is disturbed/touched, it twitches in defence Fonseca, 2025 - Nymphalis antiopaWeed, 1917 - Butterflies worth knowing.

In North America, the pupae of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty's (but also caterpillars on the way to pupation) are eaten by squirrels and chipmunks (Allen M. YoungAllen M. Young Young, 1980 - Some observations on the natural history and behavior of the Camberwell Beauty (Mourning Cloak) butterfly, Nymphalis antiopa (Linnaeus) (Lepidoptera; Nymphalidae) in the United States with reference to L. TevisL. Tevis stomach content studies of the two rodents carried out in north-east California in 1953).

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In North GreeceGreece, the Forest Caterpillar HunterCalosoma sycophantaForest Caterpillar Hunter - a predatory beetle - is a predator of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty chrysalis, although this beetle probably plays an insignificant role due to different biotope preferences Roer, 1981 - Massenwechsel-Untersuchungen an einer Südosteuropäischen Population von Nymphalis antiopa (Lep.: Nymphalidae).

Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids that infest the caterpillars of the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty can also be Organism that lives in close association with its host at the host's expense, eventually resulting in the death of the hostParasitoids of the chryasalis, such as Tachinide FlyTachinidaeTachinide Flies.

Adult Butterfly

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The typical enemies of adult Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauties are birds (predators).

The Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty is also on the menu of other animals, such as the HornetVespa crabroHornet, which Uwe LorenzenUwe Lorenzen reported on impressively in summer 2021:

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For a few days now, I have been observing a large butterfly in my garden in Schleswig, Schleswig-Holstein, GermanySchleswig, which only ever stayed on my summer lilac for a short time. Besides the MapAraschnia levana (Linnaeus, 1758)Maps and PeacockAglais io (Linnaeus, 1758)Inachis ioNymphalis ioEuropean PeacockPeacock ButterflyPeacocks, I noticed it because of its size.

On 13 August I got him in front of the camera and took one photo after the other.

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Suddenly there was a loud humming noise and the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty fell to the ground, fluttering wildly, and disappeared under the ground cover. I quickly plucked some leaves from the plants and then I saw the misery. A HornetVespa crabroHornet had bitten into the trunk. My attempt to separate the two was in vain and only put me in danger because the hornet pounced on me, buzzing loudly, but then flew away. And now he's dead the Camberwell BeautyNymphalis antiopa (Linnaeus, 1758)Mourning CloakCamberwell Beauty. I wonder how he got here?

Are adults infested by parasites or parasitoids?