Tagasaste is a valued forage for ruminants because of its good palatability and high protein content. It can be grazed directly, or cut and fed fresh or dried.
As a fodder crop tagasaste delivers between 23 and 27% crude protein (14–30% in Western Australia) and 18–24% crude indigestible fibre. With proper application of fertiliser it can maintain these levels even when grown on poor soils.
Phosphorus is particularly important both for the growth of tagasaste and for the growth of the animals grazing on it. Higher levels of phosphorus are required for the maximum growth of the animals than for maximum plant growth. Fertiliser will also reduce the level of phenolic compounds (similar to tannins) that increase when the plants are moisture stressed. These phenolic compounds make the tagasaste less palatable and reduce the animals' feed intake. The phenolics suppress the utilisation of protein in the rumen, and as a result feed intake. Despite the crude protein always being above 14%, supplementing with a high-protein feed like lupin seed stimulates animal feed intake when phenolics are high in tagasaste. Tagasaste typically has the same nutritional value as the best type of alfalfa when planted on good soil balanced in nutrients. Daily weight gains of 1 to 1.5 kg per steer per day are achieved during the growing season, with 6–10 rotations per year, typically one every 6–8 weeks.
Flowering also changes the palatability of tagasaste. In Western Australia it will commence flowering in winter (~ June). The seeds are mature and shed in early summer (first warm day in December). In the summer following flowering the palatability changes in different parts of the plant. In tagasaste that has not flowered the leaves will be grazed and bark on the stems ignored. After flowering the leaf becomes less palatable and the bark more palatable. This results in stock stripping bark off the trunk. Also the growth rate of the plant slows and leaves will be shed. The grazing management is designed to prevent flowering and keep tagasaste in the vegetative juvenile state. Trials have shown that if tagasaste is heavily grazed an  or mechanically cut in the first six months of the year, it will not flower in the second half of the year. Occasionally when mechanically cutting, one limb on a plant can be missed. This limb will go on to flower, and show all the other problems, while the rest of the plant is maintained in a vegetative state.
Initially tagasaste was developed to replace hand feeding sheep during autumn in Western Australia, when feed supplies were normally very low. This involved locking the tagasaste up for 11 months and then grazing it with 100 sheep per hectare for a month. As the tagasaste grows to about 3 metres height in 11 months, it must be mechanically cut while the sheep are in the paddock. Sheep can not be set stocked on tagasaste as this can lead to plant deaths.
A breakthrough came with the discovery that cattle can be set stocked on tagasaste. Cattle tongues are too large to pick off the new shoot buds, and some leaves always remain on the plant. Sheep can remove every leaf from tagasaste but this does not cause plant deaths. The removal of new buds by sheep, which appear about six weeks after grazing, can lead to plant deaths. Today the majority of tagasaste is used for cattle grazing with plantations able to be grazed at any time of year. Though it can grow up to 5 metres in height, when managed for grazing it is kept to less than 2 metres in height.
On the poor white sands in Western Australia tagasaste has increased the animal carrying capacity from 1 to 2 dry sheep equivalents, with annual pastures to 8-10 dry sheep equivalents (~ 1 cow) per hectare with mature tagasaste. This is roughly a tenfold increase in soil fertility-based carrying capacity. The yield of edible dry matter (leaves and fine stems) in the West Midlands is mostly in the range of 3–5 tonnes per hectare. It also prevents the wind erosion and excessive ground water recharge that were major environmental problems before. Recently it has been found that tagasaste can sequester carbon at the rate of about 6 tonnes CO₂ equivalent per hectare per year. About half the CO₂ being stored is as organic carbon in the soil and half is in the wood of the branches, trunk and roots. Tagasaste typically yields roughly 1 ton of edible material per 100mm of rain per hectare per year.


The potential of tagasaste as a Spanish fodder was identified by Dr Perez, a medical practitioner, based on La Palma island in the Canary Islands in the 1870s, and Spanish cattle farmers. He wrote to the Spanish authorities promoting tagasaste as a fodder shrub but could not get them interested. He then sent seed to Kew Gardens in England. Kew Gardens tested tagasaste and then sent seed to all its colonies around the world. In Australia, tagasaste's potential was promoted by a number of individuals over the next century (e.g. Dr Schomburge in South Australia and Dr Laurie Snook in Western Australia) but was not adopted on a large scale until the 1980s in Western Australia.
The first 2 ha of tagasaste in the West Midlands was planted by John Cook on his farm near Dandaragan in 1982. The success of this small paddock excited the interest of local farmers and researchers. Before this the farming systems in the region were totally based on annual pastures and crops. The region has strongly winter dominant rainfall, with up to eight months without rain over some summers. Until the development of tagasaste it was believed it was not possible to grow perennial pastures in this region.
In 1984 the Martindale Research Project was started by the University of Western Australia with a large grant from Sir James McCusker. This project encouraged the participation of local farmers and the Western Australia Department of Agriculture in research and development. Farmers such as John Cook and Bob Wilson at Lancelin developed seeding and mechanical cutting equipment that allowed tagasaste to be sown and managed cheaply by farmers on a broadacre scale. The Martindale Research Project resolved many issues relating to agronomy, animal production and economics that resulted in a reliable package that farmers could adopt with confidence. The Western Australia Department of Agriculture also conducted trials on their Badgingarra Research Station and at Bob Wilson's farm at Lancelin. Local farmers and researchers formed the West Midlands Fodder Shrub Improvement Group. This evolved into the Evergreen Group, which expanded its interests to include a range of other shrubs, grasses and perennial legumes.
There are now about 100,000 ha of tagasaste in Western Australia. In Western Australia it is mostly grown on deep, infertile sands in regions nearer the coast with 350 to 600 mm rainfall. Most of the tagasaste in Western Australia is in the West Midlands sand plain to the north of Perth.


In Australia a prostrate form of tagasaste has been developed by selection, called Weeping Tagasaste. It is hoped that this plant will not require mechanical cutting or pruning. It is too early to say whether Weeping Tagasaste will yield the same production levels as that of normal tagasaste. Initial indicators are that production is lower and that the plant may be more vulnerable to soil disease.
In South Africa three eco adapted genetic lines have been trademarked: "Green Kalahari" for arid areas (300–500 mm), "Cattle Candy" for temperate areas (600–800 mm) and "Kilimanjaro" cultivar for tropical highland areas (800–3500 mm) of rain. Commercial breeding of tree lucerne cultivars have led to improved varieties with increased hardiness, disease resistance and a wider range of growing conditions and applications. The use of tree lucerne for silvopasture agroforestry as a companion crop is one example of dual use of the same land for increased production. Sheep are then used to suppress weeds and keep the tree lucerne in check, preventing it from shadowing out the main forestry crop. The biological fertilisation is enhanced with green manure and trampling, improving the soil over time, and leading to production increases.


The laughing kookaburra (Dacelo novaeguineae) is a bird in the kingfisher subfamily Halcyoninae. It is a large robust kingfisher with a whitish head and a brown eye-stripe. The upperparts are mostly dark brown but there is a mottled light-blue patch on the wing coverts. The underparts are cream-white and the tail is barred with rufous and black. The plumage of the male and female birds is similar. The territorial call is a distinctive laugh that is often delivered by several birds at the same time, and is widely used as a stock sound effect in situations that involve a jungle setting.
The laughing kookaburra is native to eastern mainland Australia, but has also been introduced to parts of New Zealand, Tasmania, and Western Australia. It occupies dry eucalypt forest, woodland, city parks and gardens. This species is sedentary and occupies the same territory throughout the year. It is monogamous, retaining the same partner for life. A breeding pair can be accompanied by up to five fully grown non-breeding offspring from previous years that help the parents defend their territory and raise their young. The laughing kookaburra generally breeds in unlined tree holes or in excavated holes in arboreal termite nests. The usual clutch is three white eggs. The parents and the helpers incubate the eggs and feed the chicks. The youngest of the three nestlings or chicks is often killed by the older siblings. When the chicks fledge they continue to be fed by the group for six to ten weeks until they are able to forage independently.
A predator of a wide variety of small animals, the laughing kookaburra typically waits perched on a branch until it sees an animal on the ground and then flies down and pounces on its prey. Its diet includes lizards, insects, worms, snakes, mice and it is known to take goldfish out of garden ponds.
The International Union for Conservation of Nature (IUCN) has classed the laughing kookaburra as a species of least concern as it has a large range and population, with no widespread threats.


The laughing kookaburra was first described and illustrated (in black and white) by the French naturalist and explorer Pierre Sonnerat in his Voyage à la nouvelle Guinée, which was published in 1776. He claimed to have seen the bird in New Guinea. In fact Sonnerat never visited New Guinea and the laughing kookaburra does not occur there. He probably obtained a preserved specimen from one of the naturalists who accompanied Captain James Cook to the east coast of Australia. Edme-Louis Daubenton and François-Nicolas Martinet included a coloured plate of the laughing kookaburra based on Sonnerat's specimen in their Planches enluminées d'histoire naturelle. The plate has the legend in French "Martin-pecheur, de la Nouvelle Guinée" (Kingfisher from New Guinea).
In 1783, the French naturalist Johann Hermann provided a formal description of the species based on the coloured plate by Daubenton and Martinet. He gave it the scientific name Alcedo novæ Guineæ. The current genus Dacelo was introduced in 1815 by the English zoologist William Elford Leach, and is an anagram of Alcedo, the Latin word for a kingfisher. The specific epithet novaeguineae combines the Latin novus for new with Guinea, based on the erroneous belief that the specimen had originated from New Guinea. For many years it was believed that the earliest description was by the Dutch naturalist Pieter Boddaert and his scientific name Dacelo gigas was used in the scientific literature, but in 1926 the Australian ornithologist Gregory Mathews showed that a description by Hermann had been published earlier in the same year, 1783, and thus had precedence.  The inaccurate impression of geographic distribution given by the name in current usage had not by 1977 been considered an important enough matter to force a change in favor of D. gigas.
In the 19th century this species was commonly called the "laughing jackass", a name first recorded (as Laughing Jack-Ass) in An Account of the English Colony in New South Wales by David Collins which was published in 1798. In 1858 the ornithologist John Gould used "great brown kingfisher", a name that had been coined by John Latham in 1782. Another popular name was "laughing kingfisher". The names in several Australian indigenous languages were listed by European authors including Go-gan-ne-gine by Collins in 1798, Cuck'anda by René Lesson in 1828 and Gogera or Gogobera by George Bennett in 1834. In the early years of the 20th century "kookaburra" was included as an alternative name in ornithological publications, but it was not until 1926 in the second edition of the Official Checklist of Birds of Australia that the Royal Australasian Ornithologists Union officially adopted the name "laughing kookaburra". The name comes from Wiradjuri, an endangered Aboriginal language.
The genus Dacelo contains four kookaburra species of which the rufous-bellied kookaburra and the spangled kookaburra are restricted to New Guinea and islands in the Torres Straits. The blue-winged kookaburra and the laughing kookaburra are both widespread in Australia.
Two subspecies are recognised:
D. n. novaeguineae (Hermann, 1783) – the nominate subspecies, east Australia, Tasmania and introduced into southwest Australia
D. n. minor Robinson, 1900 – Cape York Peninsula south to Cooktown


The laughing kookaburra is the largest species of kingfisher, outsizing even the giant kingfisher in body mass. It is a stout, stocky bird 41–47 cm (16–19 in) in length, with a large head, prominent brown eyes, and a long and robust bill. The sexes are very similar, although the female is usually larger and has less blue to the rump than the male. The male weighs 196–450 g (6.9–15.9 oz), mean 307 g (10.8 oz) and the female 190–465 g (6.7–16.4 oz), mean 352 g (12.4 oz). They have a white or cream-coloured body and head with a dark brown stripe across each eye and more faintly over the top of the head. The wings and back are brown with sky blue spots on the shoulders. The tail is rusty reddish-orange with dark brown bars and white tips on the feathers. The heavy bill is black on top and bone-coloured on the bottom. The subspecies D. n. minor has a similar plumage to the nominate but is smaller in size.
The laughing kookaburra can be distinguished from the similarly sized blue-winged kookaburra by its dark eye, dark eye-stripe, shorter bill and the smaller and duller blue areas on the wing and rump. Male blue-winged kookaburras also differ in having a barred blue and black tail.


The name "laughing kookaburra" refers to the bird's "laugh", which it uses to establish territory among family groups. It can be heard at any time of day, but most frequently at dawn and dusk.
This species possesses a tracheo-bronchial syrinx, which creates two sources of vibrations so it can produce two frequencies at the same time with multiple harmonics. The laughing kookaburras call is made through a complex sound production system, by forcing air from the lungs into the bronchial tubes. While the structure for producing calls is present from an early age, the kookaburra’s song is a learned behavior. The breeding pair within a riot of kookaburra teach the fledglings to produce the signature laughing call after the young have left the nest. The adult male will sing a short portion of the call while the offspring mimics this call, usually unsuccessfully. The singing lessons tend to last two weeks before the fledgling can properly sing and take part in crepuscular choral songs. Once mastered, the young can join in crepuscular chorus songs that aid in establishing territory.
One bird starts with a low, hiccuping chuckle, then throws its head back in raucous laughter: often several others join in. If a rival tribe is within earshot and replies, the whole family soon gathers to fill the bush with ringing laughter. The laughing chorus has 5 variable elements: 1. "Kooa"; 2. "Cackle"; 3. "Rolling", a rapidly repeated "oo-oo-oo"; 4. Loud "Ha-ha"; followed by 5. Male's call of "Go-go" or female's call of "Gurgle".  Hearing kookaburras in full voice is one of the more extraordinary experiences of the Australian bush, something even locals cannot ignore; some visitors, unless forewarned, may find their calls startling.
Those calls are produced to attract/guard mates, establish and maintain the social hierarchy, and declare and defend a territory, as their calls are more often correlated with aggressiveness. Calls are utilized as neighbor/kin recognition to exhibit that groups are still inhabiting a territory. These calls also demonstrate to receivers that highly coordinated groups are of better quality and health. Acoustic communication between laughing kookaburras increases 2–3 months before the breeding season, September to January, because male aggression also increases.
The duetting call requires higher levels of cooperation within the group. The coordination of calls amongst kookaburras has been hypothesized to strengthen the main long-term pair bond and may have evolved as a mechanism to solidify the group’s bonds since it is energetically costly to learn a new song. Neighboring groups exhibit degrees of cooperation as well since chorus songs between neighbors are delivered without any overlap, alternating between groups.


Squawking is another common form of acoustic communication in D. novaeguineae that is used in a slew of different contexts. Laughing kookaburras have been noted to squawk when nesting, exhibiting submissive behavior, and when fledglings are waiting to be fed. Laughing kookaburras have a greater repertoire of calls than other kookaburra species like the Blue-winged kookaburra (Dacelo leachii) that produces two simple types of calls: “barks” and “hiccups”. This large range of calls is highlighted through cadencing, intonation, and frequency modulations that allow more detailed information to be conveyed.


The laughing kookaburra is native to eastern Australia and has a range that extends from the Cape York Peninsula in the north to Cape Otway in the south. It is present on both the eastern and the western sides of the Great Dividing Range. In the south the range extends westwards from Victoria to the Yorke Peninsula and the Flinders Ranges in South Australia.
It has been introduced into many other areas probably because of its reputation for killing snakes. In December 1891, the Western Australian parliament included 'Laughing Jackass' in the schedule of strictly preserved Australian native birds in the Game Bill, moved by Horace Sholl, member for North District. He described it as native of the North West.  His nomination is, therefore, certainly a reference to the blue-winged kookaburra (Dacelo leachii), not the laughing kookaburra (Dacelo novaeguineae). The Game Act, 1892 (Western Australia), "An Act to provide for the preservation of imported birds and animals, and of native game," provided that proclaimed Australian native birds and animals listed in the First Schedule of the Act could be declared protected from taking. Laughing Jackass was one of 23 Australian native bird species named in the schedule. Laughing kookaburras from Eastern States were released near Mullewa in around 1896 and over the following decade hundreds of birds were imported from Victoria and released around Perth. By 1912 breeding populations had been established in a number of areas. The present range in Western Australia is southwest of a line joining Geraldton on the west coast and Hopetoun on the south coast. In Tasmania the laughing kookaburra was introduced at several locations beginning in 1906. It now mainly occurs northeast of a line joining Huonville, Lake Rowallan, Waratah and Marrawah. It was introduced on Flinders Island in around 1940, where it is now widespread, and on Kangaroo Island in 1926.
In the 1860s, during his second term as governor of New Zealand, George Grey arranged for the release of laughing kookaburras on Kawau Island. The island lies in the Hauraki Gulf, about 40 km (25 mi) north of Auckland on the North Island of New Zealand. It was thought that the introduction had been unsuccessful but in 1916 some birds were discovered on the adjacent mainland. It now breeds in a small region on the western side of the Hauraki Gulf between Leigh and Kumeu.
The usual habitat is open sclerophyll forest and woodland. It is more common where the understory is open and sparse or where the ground is covered with grass. Tree-holes are needed for nesting. It also occurs near wetlands and in partly cleared areas or farmland with trees along roads and fences. In urban areas it is found in parks and gardens. The range of the laughing kookaburra overlaps with that of the blue-winged kookaburra in an area of eastern Queensland that extends from the Cape York Peninsula south to near Brisbane. Around Cooktown the laughing kookaburra tends to favour areas near water while the blue-winged kookaburra keeps to drier habitats.


During mating season, the laughing kookaburra reputedly indulges in behaviour similar to that of a wattlebird. The female adopts a begging posture and vocalises like a young bird. The male then offers her his current catch accompanied with an "oo oo oo" sound. However, some observers maintain that the opposite happens – the female approaches the male with her current catch and offers it to him. Nest-building may start in August with a peak of egg-laying from September to November. If the first clutch fails, they will continue breeding into the summer months.
The female generally lays a clutch of three semi-glossy, white, rounded eggs, measuring 36 mm × 45 mm (1.4 in × 1.8 in), at about two-day intervals. Both parents and auxiliaries incubate the eggs for 24–26 days. Hatchlings are altricial and nidicolous, fledging by day 32–40.  If the food supply is not adequate, the third egg will be smaller and the third chick will also be smaller and at a disadvantage relative to its larger siblings. Chicks have a hook on the upper mandible, which disappears by the time of fledging. If the food supply to the chicks is not adequate, the chicks will quarrel, with the hook being used as a weapon. The smallest chick may even be killed by its larger siblings. If food is plentiful, the parent birds spend more time brooding the chicks, so the chicks are not able to fight.


Kookaburras hunt much as other kingfishers (or indeed Australasian robins) do, by perching on a convenient branch or wire and waiting patiently for prey to pass by. Common prey include mice and similar-sized small mammals, a large variety of invertebrates (such as insects, earthworms and snails), yabbies, small fish, lizards, frogs, small birds and nestlings, and most famously, snakes.  Small prey are preferred, but kookaburras sometimes take large creatures, including venomous snakes, much longer than their bodies. When feeding their young, adult laughing kookaburras will make “Chuck calls”, which are deep, guttural calls that differ significantly from their daily chorus songs.


To further enhance territorial behavior, kookaburras will partake in two types of aerial displays: trapeze and circular flights. Trapeze flights are aptly named after the swooping motion that neighboring kookaburras will make towards one another in midair when defending territory. During trapeze flights, an individual from each riot will perch on branches bordering the others’ territory and fly back and forth between trees within their established home range and trees bordering neighboring kookaburras territory. These displays have been observed to last up to a half an hour and are usually accompanied by calls from the sender and members of that individual's riot. Circle flights are initiated when an individual circumnavigates a neighboring territory by flying over the area and then quickly invading neighboring territory. Once inside the neighboring territory, the individual will fly in circles around the other kookaburras inhabiting the area, which results in a typical laughing call or squawking depending on whether the neighbors’ dominance status. Flight displays are useful for communicating over long distances, but other forms of visual signals can be effective for short-range communication.
Close range visual signals can be used to convey aggression or indicate incoming threats to the flock. Aggressive posturing is used as a warning before attacking, a signal that is commonly received by foreign kookaburras encroaching on another groups’ territory. Laughing kookaburras will splay out their wings and propel their head forward while shaking their tail feathers to exhibit dominance and ward off intruders. The aggressive posturing is followed up by chasing off the unwanted individual before attacking. Visual displays are also used to communicate vigilance and the presence of threats via alert postures. D. novaeguineae will open its beak, ruffle up the feathers surrounding its cap, and angle their heads towards the direction of the threat. Depending on the urgency of the threat, alarm postures may be followed by loud laugh-like calls to warn other members of the flock.


Laughing kookaburras are a common sight in suburban gardens and urban settings, even in built-up areas, and are so tame that they will often eat out of a person's hands, and allow them to rub their bellies. It is not uncommon for kookaburras to snatch food out of people's hands without warning, by swooping in from a distance. People often feed them pieces of raw meat. Laughing kookaburras are often kept in zoos.
The kookaburra is also the subject of a popular Australian children's song, the "Kookaburra" which was written by Marion Sinclair in 1934.
Recordings of this bird have been edited into Hollywood movies for decades, usually in jungle settings, beginning with the Tarzan series in the 1930s, and more recently in the film The Lost World: Jurassic Park (1997).


The population density of the laughing kookaburra in Australia varies between 0.04 and 0.8 birds/ha depending on the habitat. Assuming an average of 0.3 birds/ha the total population may be as large as 65 million individuals. However, this may represent a severe over-estimate since the population of the laughing kookaburra seems to be undergoing a marked decline with Birdata showing a 50% drop in sightings from 2000 to 2019, and a drop in the reporting rate from 25% to 15% over the same period. The population in New Zealand is relatively small and is probably less than 500 individuals. Given the extended range and the large stable population, the species is evaluated as of "least concern" by the International Union for Conservation of Nature.


The blue dacnis or turquoise honeycreeper (Dacnis cayana) is a small passerine bird. This member of the tanager family is found from Nicaragua to  Panama, on Trinidad, and in South America south to Bolivia and northern Argentina. It is widespread and often common, especially in parts of its South American range.


In 1760 the French zoologist Mathurin Jacques Brisson included a description of the blue dacnis in his Ornithologie based on a specimen collected in Cayenne in French Guiana. He used the French name Le Pepit bleu de Cayenne and the Latin name Sylvia cayanensis caerulea. Although Brisson coined Latin names, these do not conform to the binomial system and are not recognised by the International Commission on Zoological Nomenclature. When in 1766 the Swedish naturalist Carl Linnaeus updated his Systema Naturae for the twelfth edition he added 240 species that had been previously described by Brisson. One of these was the blue dacnis. Linnaeus included a terse description, coined the binomial name Motacilla cayana and cited Brisson's work. The specific name cayana is from Cayenne. This species is now placed in the genus Dacnis that was introduced in 1816 by the French naturalist Georges Cuvier with the blue dacnis as the type species.
Eight subspecies are recognised:.        e
D. c. callaina Bangs, 1905 – west Costa Rica to southwest Panama
D. c. ultramarina Lawrence, 1864 – Honduras to northwest Colombia
D. c. napaea Bangs, 1898 – north, north-central Colombia
D. c. baudoana Meyer de Schauensee, 1946 – southwest Colombia and west Ecuador
D. c. caerebicolor Sclater, PL, 1851 – central Colombia
D. c. cayana (Linnaeus, 1766) – east Colombia to French Guiana, Trinidad and north, central Brazil
D. c. glaucogularis Berlepsch & Stolzmann, 1896 – south Colombia through east Ecuador and east Peru to north, east Bolivia
D. c. paraguayensis Chubb, C, 1910 – east, south Brazil, east Paraguay and northeast Argentina
The purplish honeycreeper (Chlorophanes purpurascens), a bird from Venezuela known only from the type specimen, is considered to be an intergeneric hybrid between the green honeycreeper and either the red-legged honeycreeper or the blue dacnis.


The blue dacnis is 12.7 cm long and weighs 13 g. Despite its alternative name, it is not a honeycreeper, which are longer-billed. The adult male is turquoise blue with a black around the eyes, and on the throat and back. The wings and tail are black, edged with turquoise. The female and immature are mainly green with a blue head, paler green underparts and green-edged brown wings. The blue dacnis's call is a thin tsip.


The blue dacnis occurs in forests and other woodlands, including gardens and parks.


Dactylorhiza fuchsii, the common spotted orchid, is a species of flowering plant in the orchid family Orchidaceae.
Dactylorhiza fuchsii is one of Europe's commonest wild orchids. It is widespread across much of Europe, with the range extending eastward into Siberia, Mongolia and Xinjiang. The species is also reportedly naturalised in the Canadian Province of Ontario.
Dactylorhiza fuchsii is a herbaceous perennial plant ranging from 10 to 60 centimetres (5 to 25 in) in height. The inflorescence is a dense-flowered spike, produced in June–August, that is at first conical then cylindrical. The flower colour can vary from white to pale purple with purple spots, a symmetrical pattern of dark purple loops or dots and dashes. The lip has three lobes. The bracts are usually shorter than the flower. The lip is smaller than that of the very similar Dactylorhiza maculata and has three deeper cuts. The middle lobe is more than half as large as a lateral lobe. Some colonies are highly perfumed, attractive to day-flying moths. The leaves are narrow lanceolate, keeled and often dark-spotted.


The orchids that are most similar belong to the Dactylorhiza maculata group. D. maculata ssp. maculata is distinguished by having the lip less deeply trilobed, while D. maculata subsp. saccifera has one spur large and saccular (sac-shaped) and the bracts of the inflorescence as long as or longer than the flowers. Outside of the "maculata group", D. majalis is very similar to D. fuchsii, but is distinguished by the following characters: the spots of the leaves are less elongated, the bracts of the inflorescence are longer and the lower transcend the inflorescence itself; it tends to be less cylindrical (a little more 'globular'), the stem is hollow (not solid) and the leaves are slightly larger. Other similar orchids are D. incarnata and D. lapponica but these species have hollow stems and different habitat (fens and bogs).


D. fuchsii is a Eurosiberian species occurring over Europe from Ireland in the west eastwards to Mongolia, the Altai Mountains and across northern Asia. It is sympatric with D. maculata.
Typical habitats are, variously across the range, conifer, beech and chestnut forests, moderately wet meadows, bogs and margins of streams. The preferred substrate is supposedly calcareous although it seems not to be particularly linked to this type of substrate. In mountain, subalpine and alpine ecosystems D. fuchsii is found from 900 to 2300 m above sea level. Elsewhere it is found from sea level.
The full list of areas (World Geographical Scheme for Recording Plant Distributions) for D. fuchsii is Finland, Great Britain, Iceland, Ireland, Norway, Sweden, Austria, Belgium, Czechia, Slovakia, Germany, Hungary, Netherlands, Poland, Switzerland, Corsica, Italy, Romania, former Yugoslavia, Belarus, Baltic States, Central European Russia, East European Russia, North European Russia, South European Russia, Northwest European Russia, Ukraine, Altay, Buryatiya, Chita, Irkutsk, Krasnoyarsk, Tuva, West Siberia, Yakutiya, Xinjiang, Mongolia.
In Italy it is found mainly in the Alps in the northern Apennines. In Britain it is widespread, the most common orchid, occurring from alkaline marshes to chalk downland. After the bee orchid, Ophrys apifera, it is the most successful orchid coloniser of waste land.


Pollination may be by bumblebees, or the longhorn beetle Alosterna tabacicolor.
Dactylorhiza fuchsii forms mycorrhizal associations with fungi in the Tulasnellaceae and Ceratobasidium groups.


Dactylorhiza is Ancient Greek for finger root, referring to the shape of the plant's roots. The species name fuchsii honours the 16th-century German botanist Leonhart Fuchs. The English name 'common spotted' refers to the species' abundance and the spots on its leaves. The French and German common names also honour Leonhart Fuchs.
This plant belongs to a problematic group of orchids. D. fuchsii is very variable in flower colour and flower morphology, plant height and the scent of flowers. This is due to the ease of introgression (the transfer of genetic material from one sympatric species to another, only partially isolated from the first, through interspecific hybridization and repeated backcrossing to a parental species), the ability of these plants to adapt quickly and easily to habitat and different substrates and possibly other causes. As a result, a multitude of forms have been defined for this plant. The World Checklist of Kew Gardens lists over 25 varieties, of which 7 are recognized as valid.


Dactylorhiza fuchsii subsp. carpatica (Batoušek & Kreutz) Kreutz – Slovakia
Dactylorhiza fuchsii subsp. fuchsii (widespread from Spain and Ireland to Siberia and Mongolia)
Dactylorhiza fuchsii subsp. hebridensis (Wilmott) Soó – British Isles
Dactylorhiza fuchsii subsp. okellyi (Druce) Soó – British Isles
Dactylorhiza fuchsii subsp. psychrophila (Schltr.) Holub – northern and central Europe to Western Siberia
Dactylorhiza fuchsii subsp. sooiana (Borsos) Borsos – Slovakia, Hungary


Hybrids with other species of the same genus are frequent. Species include:
Dactylorhiza majalis (Rchb.) PFHunt & Summerh.
Dactylorhiza traunsteineri (Saut. ex Rchb.)
Dactylorhiza incarnata (L.) Soó subsp. ochroleuca
This plant hybridizes easily with species of different genera (intergeneric hybridization). The list below shows some of these intergeneric hybrids (these hybrids are not always recognized by all botanists):
×Dactylodenia lawalreei P.Delforge & D.Tyteca – hybrid with Gymnadenia odoratissima
×Dactylodenia st-quintinii (Godfery) J. Duvigneaud in De Langhe et al. – hybrid with Gymnadenia conopsea
×Dactyloglossum mixtum (Ascherson & Graebner) Rauschert – hybrid with Coeloglossum viride
×Rhizanthera martysiensis Balayer – hybrid with Platanthera chlorantha


Dactylorhiza incarnata, the early marsh-orchid, is a perennial, temperate-climate species of orchid generally found growing in wet meadows, and generally on base-rich soils, up to about 2100m asl. The species occurs widely in Europe and Asia from Portugal and Ireland east to Siberia and Xinjiang.
There are several subspecies and also hybrids, rendering the identification of this species more difficult, but typically, the flowering spike is robust with a hollow stem, 25–60 cm tall, and bearing up to 50 flowers. Plants grow to a height of from 15 to 70 cm. The 4–7 erect yellowish-green leaves are hooded at the tip. The inflorescence is 4–12 cm long, with up to 50 blooms. The labellum appears long and narrow, since its sides are strongly reflexed (folded back). The tip is shallowly three-lobed. The flower is often flesh-coloured (the meaning of incarnata) and the labellum normally has loop-shaped markings.
The flowering period is from May to mid-July, dependent on latitude and subspecies.


Many names have been proposed for subspecies, varieties and forms within the species. Subspecies recognized as of June 2014:
Dactylorhiza incarnata subsp. coccinea – British Isles
Dactylorhiza incarnata subsp. cruenta – from France and Ireland east to Siberia and Xinjiang
Dactylorhiza incarnata subsp. gemmana  – Great Britain, Ireland, Germany, Netherlands
Dactylorhiza incarnata subsp. incarnata – from Spain and Ireland east to Siberia and Kazakhstan
Dactylorhiza incarnata subsp. jugicrucis – Transcaucasus
Dactylorhiza incarnata nothosubsp. krylovii ined. – France and Western Siberia
Dactylorhiza incarnata subsp. lobelii – Denmark, Norway, Netherlands
Dactylorhiza incarnata subsp. ochroleuca – British Isles, Sweden, France, Germany, Switzerland, Austria, Baltic Republics
Dactylorhiza incarnata subsp. pulchella – British Isles, Sweden, France, Austria, Czech Republic
Dactylorhiza incarnata nothosubsp. versicolor – Germany and Austria  (D. incarnata subsp. incarnata × D. incarnata subsp. ochroleuca)
Hybrids have been reported between D. incarnata and D. maculata, D. praetermissa, D. purpurella and D. kerryensis.


Dactylorhiza majalis, the broad-leaved marsh orchid,  is a terrestrial Eurasian orchid.
Subspecies include: western marsh orchid (Dactylorhiza majalis subsp. occidentalis), southern marsh orchid (Dactylorhiza majalis subsp. praetermissa), Hebridean marsh orchid (Dactylorhiza majalis subsp. ebudensis), and narrow-leaved marsh orchid (Dactylorhiza majalis subsp. traunsteinerioides).
The broad-leaved marsh orchid grows mainly in nitrogen-poor marsh areas that consist of several plant communities. More rarely, it is found in fens. Its flowering period begins at lower elevations as early as the beginning of May and ends in higher elevations at the end of July. The lowest blossoms usually open even before the stem has reached its full height.


The broad-leaved marsh orchid is usually 15 to 40 cm (6 to 15.5 in) tall, though some specimens may reach 60 cm (2.0 ft). Three to eight dark spotted leaves are distributed on the stem, which is hollow. The lower leaves are ovate to lanceolate and 6 to 18 cm (2.5 to 7 in) long and 1.5 to 3.5 cm (⅝ to 1⅜ in) wide. The upper leaves are increasingly smaller and more lanceolate. The bracts are about as long as the blossom and cover it before it blooms. The densely flowered inflorescence, which is 4 to 15 cm (1.5 to 6 in) long, is at first conical, but distinctly cylindrical when in full blossom. The seven to forty blossoms are colored purplish red, rarely light pink or white. The lateral tepals of the external circle of the perianth stand obliquely or vertically upright. They are 7 to 12 mm (¼ to ½ in) long and 2.5 to 5 mm (⅛ to ³⁄₁₆ in) wide. The middle tepal is smaller and forms a "helmet" together with the two lateral tepals of the internal circle. These are 6 to 11 mm (¼ to ⁷⁄₁₆ in) long. The trilobate lip is 5 to 10 mm (³⁄₁₆ to ⅜ in) long and 7 to 14 mm (¼ to ⁹⁄₁₆ in) wide. The shape and pattern of the lips are variable. In the lighter central area of the lip the markings are made up of lines, streaks, or dots. The spur is bent slightly downwards and is not quite as long as the ovary. The tuber has a palmate division and an irregular shape.
The broad-leaved marsh orchid has a karyotype of two sets of forty chromosomes. The seed of this orchid contains no endosperm for the embryo. Therefore, germination can take place only by means of infection with a root fungus (mycorrhiza).


In 1828 Ludwig Reichenbach described the broad-leaved marsh orchid as Orchis majalis. The name became the basionym after Peter Francis Hunt and Victor Samuel Summerhayes transferred the species to the genus Dactylorhiza in 1965. Sometimes the name Dactylorhiza fistulosa is used, but since this description is not valid, the name cannot be used despite its earlier publication in 1794 as Orchis fistulosa.
Many synonyms have been published:
Orchis majalis Rchb.
Dactylorchis majalis (Rchb.) Verm
Orchis baltica (Klinge) A.Fuchs
Dactylorchis baltica (Klinge) Verm.
Dactylorhiza baltica (Klinge) N.I.Orlova
Orchis longifolia Neuman
Dactylorhiza ebudensis (Wief. ex R.M.Bateman & Denholm) P.Delforge
Orchis francis-drucei Wilmott
Dactylorhiza francis-drucei (Wilmott) Aver.
Orchis kerryensis Wilmott
Dactylorchis kerryensis (Wilmott) Verm.
Dactylorhiza kerryensis (Wilmott) P.F.Hunt & Summerh.
Dactylorhiza parvimajalis D.Tyteca & Gathoye
Orchis occidentalis (Pugsley) Wilmott
Dactylorchis occidentalis (Pugsley) Verm.
Dactylorhiza occidentalis (Pugsley) P.Delforge
Orchis sphagnicola Höppner
Dactylorchis sphagnicola (Höppner) Verm.
Dactylorhiza sphagnicola (Höppner) Aver.
Dactylorchis hoeppneri (A.Fuchs) Verm.
Orchis hoeppneri (A.Fuchs) Höppner ex Verm.
Dactylorchis deweveri Verm.
Dactylorhiza deweveri (Verm.) Soó
Dactylorhiza hoeppneri (A.Fuchs) Soó
Dactylorhiza sennia Vollmar
Orchis traunsteinerioides (Pugsley) Pugsley
Dactylorchis traunsteinerioides (Pugsley) Verm.
Dactylorhiza traunsteinerioides (Pugsley) Landwehr


Many names have been proposed at the subspecies, variety and form levels, but as of June 2014 only the following are recognized:
Dactylorhiza majalis subsp. baltica (Klinge) H.Sund. – Finland, Germany, the Baltic Republics, Russia, Siberia, Kazakhstan
Dactylorhiza majalis subsp. ebudensis (Wief. ex R.M.Bateman & Denholm) M.R.Lowe – Outer Hebrides of Scotland
Dactylorhiza majalis var. francis-drucei (Wilmott) R.M.Bateman & Denholm – Scotland
Dactylorhiza majalis var. kerryensis (Wilmott) R.M.Bateman & Denholm – western Ireland
Dactylorhiza majalis subsp. majalis – widespread across much of Europe from Spain to Russia
Dactylorhiza majalis subsp. occidentalis (Pugsley) P.D.Sell – Britain and Ireland
Dactylorhiza majalis subsp. sphagnicola (Höppner) H.A.Pedersen & Hedrén – Scandinavia, Germany, France, Belgium, Netherlands
Dactylorhiza majalis subsp. traunsteinerioides (Pugsley) R.M.Bateman & Denholm – Britain and Ireland


The broad-leaved marsh orchid hybridizes quite commonly with other species of its genus.
Dactylorhiza × aschersoniana (Dactylorhiza incarnata × D. majalis)
Dactylorhiza × braunii (Dactylorhiza fuchsii × D. majalis)
Dactylorhiza × dufftiana (Dactylorhiza traunsteineri × D. majalis)
Dactylorhiza × godferyana (Dactylorhiza praetermissa × D. majalis)
Dactylorhiza × kuehnensis (Dactylorhiza ruthei × D. majalis)
Dactylorhiza × townsendiana (Dactylorhiza maculata × D. majalis)
Dactylorhiza × rupertii (Dactylorhiza sambucina × D. majalis)
More rarely, hybrids with other genera (intergeneric hybrids) occur.
×Dactyloglossum drucei (Coeloglossum viride × Dactylorhiza majalis)
×Dactylodenia lebrunii (Dactylorhiza majalis × Gymnadenia conopsea)


Dactylorhiza majalis is widespread across much of Europe and north-central Asia, from Spain and Ireland to Siberia and Kazakhstan.
In Germany the broad-leaved marsh orchid is widespread but with several gaps. In many places, especially from western to northern Germany, it is extinct.
In Switzerland it is also quite widespread. A significant gap is found south of the Aar between Aarau and Lake Neuchâtel.


Although the broad-leaved marsh orchid is commonly found in some regions, it is nevertheless protected as an orchid.
As with many marsh plants, the numbers of this species have been dwindling for quite some time. The main causes are the entry of nitrogen via fertilizer, drying out of the habitat, and intensive conversion to pasture. The broad-leaved marsh orchid does not react so sensitively to changes in its habitat as for example the early marsh orchid, Dactylorhiza incarnata. It is usually the last of the native orchids to disappear. This tolerance makes it a still relatively common species.


Dactylorhiza sambucina, the elder-flowered orchid, is an herbaceous plant belonging to the family Orchidaceae. It is quite common and widespread throughout much of Europe from Portugal east to Finland and Ukraine. The flowers appear in spring and summer, in various colors from yellow to purple.


The name of the genus Dactylorhiza is formed from Greek words δάκτυλος "daktylos" meaning "finger" and ρίζα "rhiza" meaning "root" and refers to the tubers of this plant, that are split into several tubercles. The specific Latin name "sambucina", refers to the smell of Elder (Sambucus nigra) emanating by some plants of this species.
The scientific binomial name of this plant was initially Orchis sambucina, proposed by the Swedish naturalist and botanist Carl Linnaeus in 1755. The name was subsequently amended to the one currently accepted (Dactylorhiza sambucina), by the Hungarian botanist Károly Rezső Soó in 1962.


Dactylorhiza sambucina reaches on average 10–40 centimetres (3.9–15.7 in) in height. These plants are bulbous geophytes with underground tubers or bulbs that annually produce new stems, leaves and flowers.
The leaves are amplexicaul and vary from 4 to 7 per plant. The lower leaves are oblong-obovate with obtuse apex, while the upper leaves are lanceolate with acute apex. Size of leaves: width 1 to 2.5 cm, length 6 – 12 cm.
The underground part of the stem has two tubers each one more or less deeply divided into several lobes (a characteristic of the genus Dactylorhiza), the first one plays the important functions of supplying the stem, while the secondary ones collect nutrient materials for the development of the plant that will form in the coming year.
The inflorescence is 5–10 centimetres (2.0–3.9 in) long and it is composed of flowers gathered in a dense spike. The flowers grow in the axils of bracts that are membranous and lanceolate-shaped. The flowers appear from mid-April to early July. Their colors vary from yellow with light reddish stains or purple speckled with darker spots on the labellum.
They are hermaphrodite and pollinated by insects (entomophily), especially bumblebees (Bombus species). These orchids are almost without nectar, but they are visited by various pollinator insects as they are confused with other plants with nectar. The seed germination is conditioned by the presence of specific fungi.


Dactylorhiza sambucina is quite common and widespread throughout much of Europe from Portugal east to Finland and Ukraine. It is absent from the British Isles, the Low Countries, the Dinaric Alps and less frequent on the north side of the Alps.


The elder-flowered orchid prefers fresh or dry meadows (subalpine and alpine grasslands), light woods and clearings or scrubland. They grow on siliceous and calcareous substrate, at an altitude of 300–2,000 metres (980–6,560 ft) above sea level, although locally, for instance in Uppland, Sweden, they can be found at sea level as well. The germination behavior, plant longevity, and soil requirements have recently been reviewed for the "Biological flora of Europe" .


Dactylotum bicolor, also known as the rainbow grasshopper, painted grasshopper, or the barber pole grasshopper, is a species of grasshopper in the family Acrididae. It is native to the United States, Canada and northern Mexico and exhibits aposematism (warning coloration). It was first described by the German entomologist Toussaint de Charpentier in 1843.


Dactylotum bicolor grows to an average length of about 20 mm (0.8 in) for males and 35 mm (1.4 in) for females. It is mainly black with distinctive reddish and yellowish markings, a pale green prothorax and pale green wingpads. The tibia of the hind leg bears six to eight spines. This species does not develop wings and is unable to fly.
Three subspecies are recognised:
D. b. bicolor – Northern Texas, New Mexico and Mexico
D. b. pictum – Northern and eastern part of range
D. b. variegatum – Southern Arizona and western part of range
The coloring varies across the insect's range, with D. b. pictum being black with little red, D. b. variegatum having distinct red markings, and D. b. bicolor having a purplish or violet sheen to the background black color.
Before the discovery of Dichroplus silveiraguidoi in Uruguay in 1956, Dactylotum bicolor had the lowest known number of chromosomes among grasshopper species, with seventeen acrocentric chromosomes.


It is found in shortgrass prairie, desert grasslands, thinly vegetated areas and alfalfa fields throughout the western Great Plains of the United States (and southern Canada), southward to Arizona, New Mexico, Texas, and into northern Mexico.


Its eggs are laid in soft soil in several batches of about one hundred. These overwinter and hatch in late spring or early summer, with adults being present until September or October. There is a single generation each year.
Although adult rainbow grasshoppers are polyphagous and feed on many species of plant, the nymphs feed entirely on Wright's false willow (Baccharis wrightii) in Arizona and New Mexico. It has been found that the nymphs orient themselves relative to the sun, positioning themselves around the bush so as to thermoregulate (keep their body temperature within an acceptable range). In the morning and evening they feed near the ground in full sun, but at midday they move to the shady center. At night they roost in upper branches of the bush, but this may primarily be to avoid ground-based predators.
The insect's coloration was shown to be aposematic in an experiment of predation by little striped whiptail lizards. These brightly-colored grasshoppers were less attractive to the lizards as food than cryptically-colored Trimerotropis sp of similar size, even though the lizards were unfamiliar with the rainbow grasshopper, which was not present in their natural habitat. The lizards may previously have encountered other prey with warning colors such as stink bugs, seed bugs, or velvet ants. The rainbow grasshopper is also distasteful to birds.


Dahlia coccinea is a species of flowering plant in the daisy family Asteraceae. Its common name is red dahlia, although the flowers can be orange or occasionally yellow, as well as the more common red. The species is native to Mexico but has long been cultivated in other countries for its showy flowers.


Tubers of the plant were used as a food source by the Aztecs, though this use largely died out after the Spanish conquest. The tubers are highly nutritious.


Dalea aurea, commonly called golden prairie clover, is a species of flowering plant in the legume family (Fabaceae). It is native to North America, where it is found in the Great Plains and southwestern United States, and in the state of Coahuila, Mexico. Its natural habitat is in silty or gravelly prairies, often over limestone.
It is an herbaceous perennial, with pinnate-compound leaves. It produces yellow flowers in spring and early summer.


Dalea candida is a species of flowering plant in the legume family known by the common name white prairie clover. It is native to North America, where it can be found throughout central Canada, the central United States, and northern Mexico. It can sometimes be found outside its range as an introduced species. It grows in many types of habitat, including several types of prairie, foothills, woods, forests, and disturbed areas.
It is a perennial herb growing erect to a maximum height around 1 meter (3 ft 3 in), its taproot growing up to 5 or 6 feet (1.5 or 1.8 m) deep. The alternately arranged leaves are each composed of several narrow, gland-dotted, light green leaflets. The inflorescence is a dense cylindrical spike of flowers at the tip of each stem or stem branch. The spike is packed with the pointed green calyces of sepals, the lower ones bearing corollas of white petals and the higher ones blooming later. The fruit is a green oval legume pod containing one seed.
A specimen of this species was collected by Meriwether Lewis in Nebraska in 1804.


Among the Ramah Navajo, the candida variety is used for stomachache and  as a "life medicine", especially for fever. A compound decoction is used to treat "snake infection" in sheep.


Dalea purpurea is a species of flowering plant in the legume family known as purple prairie clover. Native to central north America, purple prairie clover is a relatively common member of the Great Plains and prairie ecosystems. It blooms in the summer with dense spikes of bright purple flowers that attract many species of insects.


It is native to central North America, where it occurs from central Canada to the southeastern and southwestern United States, except for the east and west coasts. It is a common and widespread plant within its range, especially on the Great Plains. In 1804, Meriwether Lewis collected a specimen in Nebraska.


Dalea purpurea is a perennial herb growing 20 to 90 cm (8 to 35 in) tall. The mature plant has a large taproot 5.5 to 6.5 feet (1.7 to 2.0 m) deep. The stem is woody with several branches. The leaves are a few centimeters long and are divided into 3 to 7 narrow leaflets. The inflorescence atop each stem branch is a spike up to 7 cm (2+3⁄4 in) long containing many purple flowers. The fruit is a legume pod containing 1 or 2 seeds. The Latin specific epithet purpurea means purple.


This plant is adapted to a habitat with periodic wildfires. In some areas, it depends on fire to clear encroaching woody vegetation, as it cannot tolerate shade.
Dalea purpurea is a common member of the flora on the plains of central North America, occurring in a variety of habitat types, including several types of grassland. It occurs in glades, riverbanks and floodplains, oak woodlands, pinyon-juniper woodlands, shrubsteppe, many types of forests, and the Sand Hills of Nebraska. It occurs in a variety of prairie ecosystems. On tallgrass prairie it is associated with plants such as little bluestem (Schizachyrium scoparium), big bluestem (Andropogon gerardi), prairie Junegrass (Koeleria macrantha), prairie dropseed (Sporobolus heterolepis), lead plant (Amorpha canescens), and silky aster (Symphyotrichum sericeum). On midgrass prairie it grows alongside several grasses such as silver bluestem (Bothriochloa saccharoides), purple threeawn (Aristida purpurea), sideoats grama (Bouteloua curtipendula), and sand dropseed (Sporobolus cryptandrus). On shortgrass prairie it is associated with grasses such as blue grama (Bouteloua gracilis), hairy grama (B. hirsuta), and buffalo grass (B. dactyloides). This species may be considered an indicator of pristine prairie.
The nectar and pollen of Dalea purpurea attract many bees, wasps, flies, butterflies, and skippers. Several plasterer bees (genus Colletes) are specialist pollinators of Dalea species, and other insects eat the seeds and leaves.  It is a larval host to the southern dogface (Zerene cesonia).


This species is used for revegetation efforts on reclaimed land, such as land that has been strip mined. It is good for preventing erosion and for fixing nitrogen in soil. Though it is often found in mid- to late-successional stages of ecological succession, it may also be a pioneer species, taking hold in bare and disturbed habitat, such as roadsides.
Purple prairie clover provides food for a number of animals, such as pronghorn. It also grows in cultivated fields and becomes included in hay for livestock. It is nutritious and is "considered one of the most important legumes in native grasslands on the Great Plains." It also had a number of uses for Native Americans. The leaves are edible and good for making tea and medicines, and the roots are palatable when chewed. The stems were used as brooms by the Pawnee people.


Dalea purpurea has been found to contain several active constituents, including pawhuskin A, pawhuskin B, pawhuskin C, and petalostemumol. The pawhuskins possess affinity for the opioid receptors, and pawhuskin A, by far the most potent of the group, acts as a non-selective antagonist of all three opioid receptors, with preference for the κ- and μ-opioid receptors over the δ-opioid receptor.


The queen is a member of the genus Danaus, which includes D. plexippus (monarch) and D. eresimus (soldier). It is of the family Nymphalidae of the order Lepidoptera. There are as many as eleven subspecies recognized. It is a native of the Nearctic and Neotropical realms. The conservation status of this species is secure, with no reported management needs.


Females lay small white eggs one at a time on larval host plants, usually members of the milkweed subfamily Asclepiadoideae. The egg hatches into a black caterpillar with transverse white stripes and yellow spots, and three pairs of long, black filaments. The caterpillar feeds on the host plant and sequesters chemicals that make it distasteful to some predators. It then goes through six instars, after which the larva finds a suitable spot to pupate. The adult emerges 7 to 10 days afterwards. The queen butterfly has multiple generations per year.


The queen butterfly oviposits one egg at a time. Each individual egg can be found on leaves, stems, and flower buds of the host. The eggs are usually pale green, ovate to conical in shape, with a flattened base and slightly truncated top, and is longitudinally ribbed with raised cross-lines between the ridges. Compared to that of the monarch butterfly, the egg of the queen butterfly is taller relative to its width.


The mature queen caterpillar is darker and not as brightly colored as the monarch caterpillar. It is nearly identical to the caterpillar of Danaus chrysippus.

In the larval stage, the queen is bluish-white dorsally, with a reddish-brown underside. It has three pairs of black, fleshy tentacles—one is on the head, one is on the second thoracic segment and one is on the eighth abdominal segment. When mature, the caterpillar is brown with purplish prolegs. It has been observed with the following color variants of its transverse stripes: blue, green, yellow, white, and blackish brown. The head is black with white rings. The caterpillar lacks spines, and there is no hair on its body.


The pupa is relatively short and thick, tapering rapidly at the end of the abdomen. It is pale green, rarely pale pink, and is frequently ornamented with golden spots. A black transverse band edged with gold is on the abdomen. Below this black abdominal band lies another one in blue. The pupa has very few projections; most notably, it is suspended by a long cremaster from a button of silk. As such, the pupa resembles a pendant.
In general, the pupa of the queen is smaller and more slender than that of the monarch.


The queen butterfly  is related to two species to which it bears a striking resemblance: the monarch (Danaus plexippus) and the soldier (Danaus eresimus).
The queen is a moderately large butterfly, with an average wingspan of 3.1 inches (7.9 cm) to 3.3 inches (8.4 cm). It is easily distinguishable from its relative the monarch by its darker brown ground colour, and lack of stripes decorating the wings. The queen bears a closer resemblance to, the soldier (Danaus eresimus). It possesses a very tough and flexible chitinous exoskeleton, unlike most other butterflies.
Its wing coloration varies from bright, reddish brown to rich chocolate-brown, with black marginal bands that are dotted with white or yellow. The underside of the wing resembles the upper wing, but is paler. The queen has less-prominent veins on its wings and lacks the darker, apical shading found in the monarch.
Both sexes are morphologically similar. The male's and the female's forewing lengths range from 3.7 centimetres (1.5 in) to 4.6 centimetres (1.8 in), with a mean length of 4.2 centimetres (1.7 in). The antennae lack scales. Although all danaids have two pairs of walking legs, the forelegs, the first pair located on the prothoracic segment of the abdomen, are reduced and not used for locomotion. The forelegs are relatively smaller in the male than in the female. The female uses its short forelegs to scratch the surface of a leaf to determine which ones are suitable hosts for its eggs. In both sexes, the reduced forelegs lack claws.
The male queen has an androconium, a specialized, scale-covered scent-pouch, on each of  its dorsal hindwings. The positions and structures of androconia are used to identify different genera. The male also has an extensible hair-pencil on each side of its abdomen, which exudes sex pheromones. The abdominal hair-pencils, when in contact with the androconium scales, are able to disseminate pheromones near the female at integral stages of courtship.


The queen belongs to a family (Danaidae) that is common to both New and Old Worlds, specifically found throughout the tropics and into the temperate regions of the Americas, Asia, and Africa. Stray specimens are found in Europe.
The queen is chiefly a tropical species. In the US, it is usually confined to the southern portion of the country. It can be found regularly in peninsular Florida and southern Georgia, as well as in the southern portions of Texas, New Mexico, Arizona, and California. Occasionally, the subspecies of the queen can be found somewhat north, in Kansas, Colorado, and Utah. Periodically, a stray may be found in the Midwest, such as in Missouri. The berenice subspecies is found largely in the Southeast and the strigosus in the Southwest. The queen is also found in Cuba.
It is more common in southern Central America, with numbers beginning to rise in Mexico. The queen can be found as far south as Argentina.
Although the queen does not undertake dramatic migrations like the monarch, most undertake short-distance travel at tropical latitudes in areas that have a distinct dry season. During those periods, the queen will fly from lowlands to high elevations.
Throughout its distribution, the queen can be found on open land, in meadows, fields, and marshes. It displays a more xeric preference in Hispaniola and will fly to the edge of, but seldom penetrate, hammocks and forests. In the southern US, the queen prefers open woodland, fields, and desert.


The queen larvae feed on Apocynaceae (milkweeds and dogbanes). It can survive on a number of hosts. Common plants include butterflyweed (Asclepias tuberosa) and bloodflower (Asclepias curassavica). In the West Indies, blunt-leaved milkweed (Asclepias amplexicaulis) and honey vine (Cynanchum laeve) are favored. The caterpillar has also been observed on Asclepias nivea, Calotropis procera, and Apocynaceae nerium. Other reported host genera include the Apocynum, Gonolobus, Sarcostemma, and Stapelia.


As an adult, its feeding habits are less specific. The butterfly feeds predominantly on nectar from flowers and dead foliage, but can also feed on rotting fruit, sweat, and dry or wet dung, among other substances.
Even as an adult, the queen is drawn to milkweeds (Apocynaceae). However, the butterfly is also attracted to the Nerium, Funastrum, Vincetoxicum, Philabertia, Stapelia.
In addition to the above food sources, males are attracted to Heliotropium, Eupatorium, Senecio, and Crotalaria, plants known to contain the alkaloid lycopsamine. The alkaloid and other precursor compounds from these plants are used to create pheromones used to attract mates. Pheromone precursors are predominantly obtained from Boraginaceae, Asteraceae, and Fabaceae.


The queen is one of many insects that derives chemical defenses against its predators from its food plant. Most of the toxic cardenolides that make queens so unpalatable to its predators are sequestered from larval host plants.


For quite some time, the queen had been regarded as highly unpalatable to its vertebrate (mainly avian) predators. This is due to the fact that the queen, like its cousin the monarch, feeds largely on Asclepiads. As the queen and the monarch are closely related, it was assumed that the queen also possesses the ability to effectively sequester and store cardenolides present in milkweeds. As such, the queen and the Florida viceroy was long regarded a classic model-mimic example of Batesian mimicry, similar to the relationship exhibited by the monarch and the viceroy.
However, the unexpected failure of birds to reject successive queens in an experimental setting called into question the legitimacy of this relationship. In fact, experimental evidence suggested that Florida viceroys could be significantly more unpalatable than representative queens. Because experimental evidence showed sampled queens were significantly less distasteful than viceroys, it was purported that Florida viceroys and queens were Müllerian co-mimics. Furthermore, evidence from this study led to the hypothesis that the queen actually enjoys an asymmetric mimicry relationship, gaining an advantage from flying in the company of the relatively more unpalatable viceroy.


Further experimentation suggested that chemical defense of queens is highly labile. It was shown that queens reared on the high-cardenolide A. curassavica sequester and store levels of cardenolides similar to those found in monarchs. These butterflies were regarded as very distasteful and were largely rejected by avian predators. Furthermore, those that were eaten elicited high rates of distress behavior. However, queens reared on S. clausum, a larval host plant known to be a very poor cardenolide source, contain no detectable cardenolide and are essentially palatable to predators. These highly variable responses of avian predators to queens reared on different plants suggest the existence of a food-plant-related palatability spectrum in Florida queen butterflies.
Micro-geographic differences in the environment lead to variation in the dynamics of mimetic relationships even at a local level. Spatiotemporal variation throughout different areas lead to large differences in unpalatability of queens separated by only a few kilometers. This extensive variation supports the idea that automimicy occurs at the intrapopulation level – palatable queens mimic individuals that have higher cardenolide content. By extension, interspecific mimicry is also highly variable. At hydric inland sites, which contain large numbers of A. curassavica, queens and viceroys are distasteful Müllerian mimics of one another, while at coastal sites queens probably serve as the palatable Batesian mimics of viceroys.


During courtship, which occurs while both butterflies are in flight, the male everts his hair-pencils and brushes them against the female's antennae. This act is called "hair-pencilling." The secretion associated with these hair-pencils plays an important role in seducing the female. When the female comes to rest, the male hovers closely above her and subjects her to further “hair-pencilling” before alighting next to the female and copulating with her. Afterwards, the two engage in a postnuptial flight - the male flies with the female dangling beneath him.


The chemicals that comprise the pheromone are secreted by trichogen cells, which are located at the base of each hair-pencil. This liquid secretion moves from these cells, through the cuticle of the hairs, to coat the numerous free, cuticular dust particles that adhere to the hair-pencil surface. Two of the chemicals that comprise this secretion have been identified – a crystalline pyrrolizidinone (ketone) and a viscous terpenoid alcohol (diol). The diol imparts a stickiness that allows the secretion to stay on the dust, and the dust on antennae. The ketone is a releaser pheromone, inducing females to mate. Although insufficient levels of ketone present in the dust particle correlates to lower seductive capacity in the male, some males with low levels of ketone – and even some without hair-pencils – have been known to mate successfully with females. This suggests that although hair-pencil pheromones are of major importance, they are not absolutely essential to mating.


Many butterflies possess extrusible brushlike structures, called hair-pencils. In the queen, the hair-pencils, which are present in the posterior abdomen in the male, are tucked away when the male is not interacting with the female. As such, these organs are thought to serve as important tools for pheromone dissemination during courtship.
Hair-pencils play an important role in courtship success. Although the lack of hair-pencils does not affect the rate at and enthusiasm with which males pursue females, males without hair-pencils experience significantly lower success in achieving copulation. Male queen butterflies with physically normal but chemically deficient hair-pencils also suffer from lower mating success. In addition, adult female queens whose antennae have been blocked are not receptive to advances from competent male queens. However, physical contact between the male's hair-pencil and the female's antennae does not affect a male's mating success. Males without hair-pencils are no less fertile than males with hair-pencils.
That actively hair-pencilling males emit a very definite odor that can even be perceived by humans also supports the idea that it is not the hair-pencil itself that is important in courtship, but rather, the pheromone which the hair-pencil transports.


Danaus petilia, the lesser wanderer,  is a species of butterfly in the nymphalid Danainae subfamily. It is a migratory species which is found in Australia and in tropical countries. Previously considered a subspecies of Danaus chrysippus, this species came about through allopatric speciation. The deep sea barrier called Lydekker's Line, located by the Molluccas and the Sahul Shelf, was what separated Danaus petilia from Danaus chrysippus cratippus. Its caterpillars feed on native and introduced cotton bush species.
A common butterfly with nomadic populations, particularly in central Australia where many native host plants die of during dry periods, and in south eastern Australia where it is too cold for them in winter. Butterflies prefer an open country or farmland habitat. Flight is generally slow and close to the ground.


Host plants include of a range of native and introduced plants that have a milky sap like Gomphocarpus, Cynanchum, Secamone, Marsdenia, Ascelpias, Calotropis, Stapelia and many more.


The monarch butterfly or simply monarch (Danaus plexippus) is a milkweed butterfly (subfamily Danainae) in the family Nymphalidae. Other common names, depending on region, include milkweed, common tiger, wanderer, and black-veined brown. It is often considered to be amongst the most familiar of North American butterflies. It is often considered an iconic pollinator species, although it is not an especially effective pollinator of milkweeds. Its wings feature an easily recognizable black, orange, and white pattern, with a wingspan of 8.9–10.2 cm (3+1⁄2–4 in) A Müllerian mimic, the viceroy butterfly, is similar in color and pattern, but is markedly smaller and has an extra black stripe across each hindwing.
The eastern North American monarch population is notable for its annual southward late-summer/autumn instinctive migration from the northern and central United States and southern Canada to Florida and Mexico. During the fall migration, monarchs cover thousands of miles, with a corresponding multigenerational return north. The western North American population of monarchs west of the Rocky Mountains often migrates to sites in southern California, but has been found in overwintering Mexican sites, as well.  In 2009, monarchs were reared on the International Space Station, successfully emerging from pupae located in the station's Commercial Generic Bioprocessing Apparatus.


The name "monarch" is believed to have been given in honor of King William III of England, as the butterfly's main color is that of the king's secondary title, Prince of Orange. The monarch was originally described by Carl Linnaeus in his Systema Naturae of 1758 and placed in the genus Papilio. In 1780, Jan Krzysztof Kluk used the monarch as the type species for a new genus, Danaus.
Danaus (Ancient Greek Δαναός), a great-grandson of Zeus, was a mythical king in Egypt or Libya, who founded Argos; Plexippus (Πλήξιππος) was one of the 50 sons of Aegyptus, the twin brother of Danaus. In Homeric Greek, his name means "one who urges on horses", i.e., "rider" or "charioteer". In the 10th edition of Systema Naturae, at the bottom of page 467, Linnaeus wrote that the names of the Danai festivi, the division of the genus to which Papilio plexippus belonged, were derived from the sons of Aegyptus. Linnaeus divided his large genus Papilio, containing all known butterfly species, into what we would now call subgenera. The Danai festivi formed one of the "subgenera", containing colorful species, as opposed to the Danai candidi, containing species with bright white wings. Linnaeus wrote: "Danaorum Candidorum nomina a filiabus Danai Aegypti, Festivorum a filiis mutuatus sunt." (English: "The names of the Danai candidi have been derived from the daughters of Danaus, those of the Danai festivi from the sons of Aegyptus.")
Robert Michael Pyle suggested Danaus is a masculinized version of Danaë (Greek Δανάη), Danaus's great-great-granddaughter, to whom Zeus came as a shower of gold, which seemed to him a more appropriate source for the name of this butterfly.
The three species of monarch butterflies are:
D. plexippus, described by Linnaeus in 1758, is the species known most commonly as the monarch butterfly of North America. Its range actually extends worldwide and can be found in Hawaii, Australia, New Zealand, Spain, and the Pacific Islands.
D. erippus, the southern monarch, was described by Pieter Cramer in 1775. This species is found in tropical and subtropical latitudes of South America, mainly in Brazil, Uruguay, Paraguay, Argentina, Bolivia, Chile, and southern Peru. The South American monarch and the North American monarch may have been one species at one time. Some researchers believe the southern monarch separated from the monarch's population some 2 million years ago, at the end of the Pliocene. Sea levels were higher, and the entire Amazonas lowland was a vast expanse of brackish swamp that offered limited butterfly habitat.
D. cleophile, the Jamaican monarch, described by Jean-Baptiste Godart in 1819, ranges from Jamaica to Hispaniola.
Six subspecies and two color morphs of D. plexippus have been identified:
D. p. plexippus – nominate subspecies, described by Linnaeus in 1758, is the migratory subspecies known from most of North America.
D. p. p. "form nivosus", the white monarch commonly found on Oahu, Hawaii, and rarely in other locations.
D. p. p. (as yet unnamed) – a color morph lacking some wing vein markings.
D. p. nigrippus (Richard Haensch, 1909) – South America - as forma: Danais  [sic] archippus f. nigrippus. Hay-Roe et al. in 2007 identified this taxon as a subspecies
D. p. megalippe (Jacob Hübner, [1826]) – nonmigratory subspecies, and is found from Florida and Georgia southwards, throughout the Caribbean and Central America to the Amazon River.
D. p. leucogyne (Arthur G. Butler, 1884) − St. Thomas
D. p. portoricensis Austin Hobart Clark, 1941 − Puerto Rico
D. p. tobagi Austin Hobart Clark, 1941 − Tobago
The population level of the white morph in Oahu is nearing 10%. On other Hawaiian islands, the white morph occurs at a relatively low frequency. White monarchs (D. p. p. "form nivosus") have been found throughout the world, including Australia, New Zealand, Indonesia, and the United States. However, some taxonomists disagree on these classifications.
Monarchs belong in the subfamily Danainae of the family Nymphalidae. Danainae was formerly considered a separately family Danaidae.


The monarch was the first butterfly to have its genome sequenced. The 273-million-base pair draft sequence includes a set of 16,866 protein-coding genes. The genome provides researchers insights into migratory behavior, the circadian clock, juvenile hormone pathways, and microRNAs that are differentially expressed between summer and migratory monarchs. More recently, the genetic basis of monarch migration and warning coloration has been described.
No genetic differentiation exists between the migratory populations of eastern and western North America. Recent research has identified the specific areas in the genome of the monarch that regulate migration. No genetic difference is seen between a migrating and nonmigrating monarch, but the gene is expressed in migrating monarchs, but not expressed in nonmigrating monarchs.
A 2015 publication identified genes from wasp bracoviruses in the genome of the North American monarch leading to articles about monarch butterflies being genetically modified organisms.


Like all Lepidoptera, monarchs undergo complete metamorphosis; their life cycle has four phases: egg, larva, pupa, and adult. Monarchs transition from eggs to adults during warm summer temperatures in as little as 25 days, extending to as many as seven weeks during cool spring conditions. During their development, both larvae and their milkweed hosts are vulnerable to weather extremes, predators, parasites, and diseases; commonly fewer than 10% of monarch eggs and caterpillars survive.


The egg is derived from materials ingested as a larva and from the spermatophores received from males during mating. Female monarchs lay eggs singly, most often on the underside of a young leaf of a milkweed plant during the spring and summer. Females secrete a small amount of glue to attach their eggs directly to the plant. They typically lay 300 to 500 eggs over a two- to five-week period.
Eggs are cream colored or light green, ovate to conical in shape, and about 1.2×0.9 mm in size. The eggs weigh less than 0.5 mg each and have raised ridges that form longitudinally from the point to apex to the base. Although each egg is 1⁄1000 the mass of the female, she may lay up to her own mass in eggs. Females lay smaller eggs as they age. Larger females lay larger eggs. The number of eggs laid by a female, which may mate several times, can reach 1,180.
Eggs take three to eight days to develop and hatch into larvae or caterpillars. The offspring's consumption of milkweed benefits health and helps defend them against predators.  Monarchs lay eggs along the southern migration route.


To prepare for the pupal or chrysalis stage, the caterpillar chooses a safe place for pupation, where it spins a silk pad on a downward-facing horizontal surface. At this point, it turns around and securely latches on with its last pair of hind legs and hangs upside down, in the form of the letter J. After "J-hanging" for about 12–16 hours, it soon straightens out its body and goes into peristalsis some seconds before its skin splits behind its head. It then sheds its skin over a period of a few minutes, revealing a green chrysalis. At first, the chrysalis is long, soft, and somewhat amorphous, but over a few hours, it compacts into its distinct shape – an opaque, pale-green chrysalis with small golden dots near the bottom, and a gold-and-black rim around the dorsal side near the top. At first, its exoskeleton is soft and fragile, but it hardens and becomes more durable within about a day. At this point, it is about 2.5 cm (1 in) long and 10–12 mm (3⁄8–7⁄16  in) wide, weighing about 1.2 g. At normal summer temperatures, it matures in 8–15 days (usually 11–12 days). During this pupal stage, the adult butterfly forms inside. A day or so before emerging, the exoskeleton first becomes translucent and the chrysalis more bluish. Finally, within 12 hours or so, it becomes transparent, revealing the black and orange colors of the butterfly inside before it ecloses (emerges).


The adult emerges from its chrysalis after about two weeks of pupation. The emergent adult hangs upside down for several hours while it pumps fluids and air into its wings, which expand, dry, and stiffen. The butterfly then extends and retracts its wings. Once conditions allow, it flies and feeds on a variety of nectar plants. During the breeding season, adults reach sexual maturity in 4–5 days. However, the migrating generation does not reach maturity until overwintering is complete.
The adult's wingspan ranges from 8.9 to 10.2 centimetres (3.5 to 4.0 in). The upper sides of the wings are tawny orange, the veins and margins are black, and two series of small white spots occur in the margins. Monarch forewings also have a few orange spots near their tips. Wing undersides are similar, but the tips of forewings and hindwings are yellow brown instead of tawny orange and the white spots are larger. The shape and color of the wings change at the beginning of the migration and appear redder and more elongated than later migrants. Wings size and shape differ between migratory and nonmigratory monarchs. Monarchs from eastern North America have larger and more angular forewings than those in the western population.
In eastern North American populations, overall wing size in the physical dimensions of wings varies. Males tend to have larger wings than females, and are typically heavier than females. Both males and females have similar thoracic dimensions. Female monarchs tended to have thicker wings, which is thought to convey greater tensile strength and reduce the likelihood of being damaged during migration. Additionally, females had lower wing loading than males, which would mean females require less energy to fly.
Adults are sexually dimorphic. Males are slightly larger than females and have a black spot on a vein on each hindwing. The spots contain scales that produce pheromones that many Lepidoptera use during courtship. Females are often darker than males and have wider veins on their wings. The ends of the abdomens of males and females differ in shape.
The adult's thorax has six legs, but as in all of the Nymphalidae, the forelegs are small and held against the body. The butterfly uses only its middle and hindlegs when walking and clinging.
Adults typically live for 2–5 weeks during their breeding season. Larvae growing in high densities are smaller, have lower survival, and weigh less as adults compared with those growing in lower densities.


Physiological experiments suggest that monarch butterflies view the world through a tetrachromatic system. Like humans, their retina contain three types of opsin proteins, expressed in distinct photoreceptor cells, each of which absorbs light at a different wavelength. Unlike humans, one of those types of photoreceptor cells corresponds to a wavelength in the ultraviolet range; the other two correspond to blue and green.
In addition to these three photoreceptors cells in the main retina, monarch butterfly eyes contain orange filtering pigments that filter the light reaching some green-absorbing opsins, thereby making a fourth photoreceptor cell sensitive to longer-wavelength light. The combination of filtered and unfiltered green opsins permits the butterflies to distinguish yellow from orange colors. The ultraviolet opsin protein has also been detected in the dorsal rim region of monarch eyes. One study suggests that this allows the butterflies the ability to detect ultraviolet polarized skylight to orient themselves with the sun for their long migratory flight.
These butterflies are capable of distinguishing colors based on their wavelength only, and not based on intensity; this phenomenon is termed "true color vision". This is important for many butterfly behaviors, including seeking nectar for nourishment, choosing a mate, and finding milkweed on which to lay eggs. One study found that floral color is more easily recognized at a distance by butterflies searching for nectar than floral shape. This may be because flowers have highly contrasting colors to the green background of a vegetative landscape. On the other hand, leaf shape is important for oviposition so that the butterflies can ensure their eggs are being laid on milkweed.
Beyond the perception of color, the ability to remember certain colors is essential in the life of monarch butterflies. These insects can easily learn to associate color, and to a lesser extent, shape, with sugary food rewards. When searching for nectar, color is the first cue that draws the insect's attention toward a potential food source, and shape is a secondary characteristic that promotes the process. When searching for a place to lay its eggs, the roles of color and shape are switched. Also, a difference may exist between male and female butterflies from other species in terms of the ability to learn certain colors; however, no differences is noted between the sexes for monarch butterflies.


Monarch courtship occurs in two phases. During the aerial phase, a male pursues and often forces a female to the ground. During the ground phase, the butterflies copulate and remain attached for about 30 to 60 minutes. Only 30% of mating attempts end in copulation, suggesting that females may be able to avoid mating, though some have more success than others. During copulation, a male transfers his spermatophore to a female. Along with sperm, the spermatophore provides a female with nutrition, which aids her in laying eggs. An increase in spermatophore size increases the fecundity of female monarchs. Males that produce larger spermatophores also fertilize more females' eggs.
Females and males typically mate more than once. Females that mate several times lay more eggs. Mating for the overwintering populations occurs in the spring, prior to dispersion. Mating is less dependent on pheromones than other species in its genus. Male search and capture strategies may influence copulatory success, and human-induced changes to the habitat can influence monarch mating activity at overwintering sites.


The range of the western and eastern populations of D. p. plexippus expands and contracts depending upon the season. The range differs between breeding areas, migration routes, and winter roosts. However, no genetic differences between the western and eastern monarch populations exist; reproductive isolation has not led to subspeciation of these populations, as it has elsewhere within the species' range.
In the Americas, the monarch ranges from southern Canada through northern South America. It is also found in Bermuda, Cook Islands, Hawaii, Cuba, and other Caribbean islands the Solomons, New Caledonia, New Zealand, Papua New Guinea, Australia, the Azores, the Canary Islands, Madeira, continental Portugal, Gibraltar, the Philippines, and Morocco. It appears in the UK in some years as an accidental migrant.
Overwintering populations of D. p. plexippus are found in Mexico, California, along the Gulf Coast of the United States, year-round in Florida, and in Arizona where the habitat has the specific conditions necessary for their survival. On the East Coast of the United States, they have overwintered as far north as Lago Mar, Virginia Beach, Virginia. Their wintering habitat typically provides access to streams, plenty of sunlight (enabling body temperatures that allow flight), and appropriate roosting vegetation, and is relatively free of predators.
Overwintering, roosting butterflies have been seen on basswoods, elms, sumacs, locusts, oaks, osage-oranges, mulberries, pecans, willows, cottonwoods, and mesquites. While breeding, monarch habitats can be found in agricultural fields, pasture land, prairie remnants, urban and suburban residential areas, gardens, trees, and roadsides – anywhere where there is access to larval host plants.


Although larvae eat only milkweed, adult monarchs feed on the nectar of many plants, including:
Apocynum cannabinum – Indian hemp
Asclepias spp. – milkweeds
Aster sp. – asters
Cirsium sp. – thistles
Daucus carota – wild carrot
Dipsacus sylvestris – teasel
Echinacea sp. – coneflowers
Erigeron canadensis – horseweed
Eupatorium maculatum – spotted Joe-Pye weed
Eupatorium perfoliatum – common boneset
Hesperis matronalis – dame's rocket
Liatris sp. – blazing stars
Medicago sativa – alfalfa
Solidago sp. – goldenrod
Syringa vulgaris – lilac
Trifolium pratense – red clover
Vernonia altissima – tall ironweed
Monarchs obtain moisture and minerals from damp soil and wet gravel, a behavior known as mud-puddling. The monarch has also been noticed puddling at an oil stain on pavement.


In North America, monarchs migrate both north and south on an annual basis, in a long-distance journey that is fraught with risks. This is a multi-generational migration, with individual monarchs only making part of the full journey. The population east of the Rocky Mountains attempts to migrate to the sanctuaries of the Mariposa Monarca Biosphere Reserve in the Mexican state of Michoacán and parts of Florida. The western population tries to reach overwintering destinations in various coastal sites in central and southern California. The overwintered population of those east of the Rockies may reach as far north as Texas and Oklahoma during the spring migration. The second, third, and fourth generations return to their northern locations in the United States and Canada in the spring.
Captive-raised monarchs appear capable of migrating to overwintering sites in Mexico, though they have a much lower migratory success rate than do wild monarchs (see section on captive-rearing below). Monarch overwintering sites have been discovered recently in Arizona. Monarchs from the eastern US generally migrate longer distances than monarchs from the western US.
Since the 1800s, monarchs have spread throughout the world, and there are now many non-migratory populations globally.
Flight speeds of adults are around 9 km/h (6 mph).


In both caterpillar and butterfly form, monarchs are aposematic, warding off predators with a bright display of contrasting colors to warn potential predators of their undesirable taste and poisonous characteristics. One monarch researcher emphasizes that predation on eggs, larvae or adults is natural, since monarchs are part of the food chain, thus people should not take steps to kill predators of monarchs.
Larvae feed exclusively on milkweed and consume protective cardiac glycosides. Toxin levels in Asclepias species vary. Not all monarchs are unpalatable, but exhibit Batesian or automimics. Cardiac glycosides levels are higher in the abdomen and wings. Some predators can differentiate between these parts and consume the most palatable ones.
Butterfly weed (A. tuberosa) lacks significant amounts of cardiac glycosides (cardenolides), but instead contains other types of toxic glycosides, including pregnanes. This difference may reduce the toxicity of monarchs whose larvae feed on that milkweed species, as a naturalist and others have reported that monarch caterpillars do not favor the plant. Some other milkweeds have similar characteristics.


While monarchs have a wide range of natural predators, none of these is suspected of causing harm to the overall population, or are the cause of the long-term declines in winter colony sizes.
Several species of birds have acquired methods that allow them to ingest monarchs without experiencing the ill effects associated with the cardiac glycosides (cardenolides). The black-backed oriole is able to eat the monarch through an exaptation of its feeding behavior that gives it the ability to identify cardenolides by taste and reject them. The black-headed grosbeak, though, has developed an insensitivity to secondary plant poisons that allows it to ingest monarchs without vomiting. As a result, these orioles and grosbeaks periodically have high levels of cardenolides in their bodies, and they are forced to go on periods of reduced monarch consumption. This cycle effectively reduces potential predation of monarchs by 50% and indicates that monarch aposematism has a legitimate purpose. The black-headed grosbeak has also evolved resistance mutations in the molecular target of the heart poisons, the sodium pump. The specific mutations that evolved in one of the grosbeak's four copies of the sodium pump gene are the same as those found in other milkweed butterflies like the common crow that also evolved to resist cardiac glycosides. Other bird predators include brown thrashers, grackles, robins, cardinals, sparrows, scrub jays, and pinyon jays.
The monarch's white morph appeared in Oahu after the 1965-1966 introduction of two bulbul bird species, Pycnonotus cafer and Pycnonotus jocosus. These are now the most common avian insectivores in Hawaii, and probably the only ones that eat insects as large as monarchs. Although Hawaiian monarchs have low cardiac glycoside levels, the birds may also be tolerant of that toxin. The two species hunt the larvae and some pupae from the branches and undersides of leaves in milkweed bushes. The bulbuls also eat resting and ovipositing adults, but rarely flying ones. Because of its color, the white morph has a higher survival rate than the orange one. This is either because of apostatic selection (i.e., the birds have learned the orange monarchs can be eaten), because of camouflage (the white morph matches the white pubescence of milkweed or the patches of light shining through foliage), or because the white morph does not fit the bird's search image of a typical monarch, so is thus avoided.
Some mice, particularly the black-eared mouse (Peromyscus melanotis), are, like all rodents, able to tolerate large doses of cardenolides and are able to eat monarchs. Overwintering adults become less toxic over time making them more vulnerable to predators. In Mexico, about 14% of the overwintering monarchs are eaten by birds and mice and black-eared mice can eat up to 40 monarchs per night.
In North America, eggs and first-instar larvae of the monarch are eaten by larvae and adults of the introduced Asian lady beetle (Harmonia axyridis). The Chinese mantis (Tenodera sinensis) will consume the larvae once the gut is removed thus avoiding cardenolides. Predatory wasps commonly consume larvae, though large larvae may avoid wasp predation by dropping from the plant or by jerking their bodies.


Monarchs are toxic and foul-tasting because of the presence of cardenolides in their bodies, which the caterpillars ingest as they feed on milkweed. Monarchs and other cardenolide-resistant insects rely on a resistant form of the Na+/ K+-ATPase enzyme to tolerate significantly higher concentrations of cardenolides than nonresistant species. By ingesting a large amount of plants in the genus Asclepias, primarily milkweed, monarch caterpillars are able to sequester cardiac glycosides, or more specifically cardenolides, which are steroids that act in heart-arresting ways similar to digitalis. It has been found that monarchs are able to sequester cardenolides most effectively from plants of intermediate cardenolide content rather than those of high or low content. Three mutations that evolved in the monarch's Na+/ K+-ATPase were found to be sufficient together to confer resistance to dietary cardiac glycosides. This was tested by swapping these mutations into the same gene in the fruit fly Drosophila melanogaster using CRISPR-Cas9 genome editing. These fruit flies-turned monarch flies were completely resistant to dietary ouabain, a cardiac glycoside found in Apocynaceae, and even sequestered some through metamorphosis, like the monarch.
Different species of milkweed have different effects on growth, virulence, and transmission of parasites. One species, Asclepias curassavica, appears to reduce the symptoms of Ophryocystis elektroscirrha (OE) infection. The two possible explanations for this include that it promotes overall monarch health to boost the monarch's immune system or that chemicals from the plant have a direct negative effect on the OE parasites. A. curassavica does not cure or prevent the infection with OE; it merely allows infected monarchs to live longer, and this would allow infected monarchs to spread the OE spores for longer periods. For the average home butterfly garden, this scenario only adds more OE to the local population.
After the caterpillar becomes a butterfly, the toxins shift to different parts of the body. Since many birds attack the wings of the butterfly, having three times the cardiac glycosides in the wings leaves predators with a very foul taste and may prevent them from ever ingesting the body of the butterfly. To combat predators that remove the wings only to ingest the abdomen, monarchs keep the most potent cardiac glycosides in their abdomens.


The monarch is the state insect of Alabama, Idaho, Illinois, Minnesota, Texas, Vermont, and West Virginia. Legislation was introduced to make it the national insect of the United States, but this failed in 1989 and again in 1991.
Homeowners are increasingly establishing butterfly gardens; monarchs can be attracted by cultivating a butterfly garden with specific milkweed species and nectar plants. Efforts are underway to establish these monarch waystations.
An IMAX film, Flight of the Butterflies, describes the story of the Urquharts, Brugger, and Trail to document the then-unknown monarch migration to Mexican overwintering areas.
Sanctuaries and reserves have been created at overwintering locations in Mexico and California to limit habitat destruction. These sites can generate significant tourism revenue.
However, with less tourism, monarch butterflies will have a higher survival rate because they show more protein content and a higher value of immune response and oxidative defense.
Organizations and individuals participate in tagging programs. Tagging information is used to study migration patterns.
The 2012 novel by Barbara Kingsolver, Flight Behavior, deals with the appearance of a large population in the Appalachians.