Arctia plantaginis

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Wood tiger
Parasemia plantaginis.jpg
Male
Parasemia plantaginis01.jpg
Mounted
Scientific classification OOjs UI icon edit-ltr.svg
Domain: Eukaryota
Kingdom: Animalia
Phylum: Arthropoda
Class: Insecta
Order: Lepidoptera
Superfamily: Noctuoidea
Family: Erebidae
Subfamily: Arctiinae
Genus: Arctia
Species:
A. plantaginis
Binomial name
Arctia plantaginis
Synonyms
List
  • Phalaena plantaginisLinnaeus, 1758
  • Phalaea alpicolaScopoli, 1763
  • Bombyx hospita[Denis & Schiffermüller, 1775]
  • Bombyx matronalisFreyer, 1843
  • Nemeophila plantaginis floccosaGraeser, 1888
  • Parasemia plantaginis uralensisKrulikowsky, 1904
  • Parasemia plantaginis insularumSeitz, 1910
  • Parasemia plantaginis carpathicaDaniel, 1939
  • Chelonia plantaginis var. kamtschaticaMénétriès, 1857
  • Parasemia plantaginis kamtschadalusBryk, 1942
  • Parasemia plantaginis paramushiraBryk, 1942
  • Parasemia plantaginis passanauriensisAlberti, 1973
  • Parasemia plantaginis hospita f. interruptaSchawerda, 1910
  • Parasemia plantaginis jezoensisInoue, 1976
  • Nemeophila macromera var. leucomeraButler, 1881
  • Parasemia plantaginis japonicaInoue & Kobayashi, 1956
  • Nemeophila plantaginis ab. melasChristoph, 1893
  • Nemeophila caespitisGrote & Robinson, 1868
  • Nemeophila caespitisBoisduval, 1869
  • Nemeophila cichoriiGrote & Robinson, 1868
  • Nemeophila cichoriiBoisduval, 1869
  • Platarctia modestaPackard, 1864
  • Nemeophila alascensisStretch, 1906
  • Eupsychoma geometricaGrote, 1865
  • Nemeophila geddesiNeumoegen, 1884
  • Platarctia scudderiPackard, 1864
  • Nemeophila selwyniiH. Edwards, 1885
  • Parasemia plantaginis macromera ab. sachalinensisMatsumura, 1927
  • Parasemia plantaginis altaicaSeitz, 1910
  • Parasemia plantaginis stoetzneriO.Bang-Haas, 1927
  • Parasemia plantaginis(Linnaeus, 1758)

Arctia plantaginis, the wood tiger, is a moth of the family Erebidae. Several subspecies are found in the Holarctic ecozone south to Anatolia, Transcaucasus, northern Iran, Kazakhstan, Mongolia, China, Korea and Japan. One subspecies is endemic to North America.

Contents

This species was formerly a member of the genus Parasemia, but was moved to Arctia along with the other species of the genera Acerbia, Pararctia, Parasemia, Platarctia, and Platyprepia. [1] [2]

P. plantaginis males occur predominantly in two distinct color phenotypes: yellow and white. They are aposematic, meaning their colorations serve to deter predators from attacking. In populations of aposematic species, it is common to have a single coloration phenotype dominate, because predators better learn to avoid the more common phenotype and rare phenotypes suffer higher predation. Rare phenotypes are often selected against because predators are less familiar with their aposematic signal. Thus, other selective pressures exist to perpetuate weaker aposematic signals in exchange for other adaptive benefits. [3] P. plantaginis has become a common model for studying the counteracting selective pressures of predation, mate choice, immune function, thermoregulation, and more.

Figures 3-7, wood tiger moth forms Moths of the British Isles Plate080.jpg
Figures 3–7, wood tiger moth forms

Description

This moth is extraordinarily variable. The wingspan is 32–38 mm. Normally, it has a black forewing in both sexes, with moderately broad, ivory yellow bands. In the male, the hindwing is yellow or white with an irregular marginal band, which is often interrupted, and two or three submarginal spots. The basal portion of the hindwing bears black streaks at the margin of the cell and before the anal margin. In the female, the hindwing is red above with the base strongly black. Numerous aberrations have been found and named, which often occur predominantly, and only exceptionally among typical specimens. Major aberrations are listed by Seitz, 1913. [4]

Geographic range

There are populations throughout the globe, but most common in northern latitudes [5] of North America and Eurasia. [6] The North American populations range from Alaska to Manitoba, and south through the Rocky Mountain region to southern New Mexico, with isolated populations occurring in Arizona and the Sierra Nevada mountains of California and Nevada. [6]

Habitat

P. plantaginis prefer slightly moist areas, like meadows with nearby streams. Adults like to spend time close to lupine stands, which are meadows of plants from the genus Lupinus . It is estimated that over 250 annual and perennial species of this genus Lupinus are distributed throughout both montane and lowland habitats, with hugely diverse regions found in North and South America. [7]

Home range and territoriality

Genetic population structure

A two-year study of populations of P. plantaginis throughout the Alpine regions of Italy, Austria and Switzerland indicated a single whole population. Pairwise Fst values, AMOVA and COl results showed little to no differentiation between populations during the two sampling years of 2009 and 2010. This overall high genetic diversity and low differentiation between populations suggests much gene flow and high population density in P. plantaginis populations. Though this extreme gene flow would be thought to lead to fixation of a single morphological phenotype, the differential selective pressures experienced by various populations of the species likely leads to the maintenance of its widespread polymorphism. [8]

Food resources

Wood tiger moths are polyphagous, meaning their diet can vary significantly. Eating different host plants can result in different immune function and overall life history traits; one example of this is shown by wood tiger moth caterpillars that feed on ribwort plantain. These plant contain high levels of iridoid glycosides, which help caterpillars produce defensive chemicals. A 2015 study showed that the iridoid glycosides present in plantain-eating larvae is sufficient to deter both ants and parasitoids. [9]

Caterpillars

Though it may benefit caterpillars to intake more plant compounds that can help produce defensive chemicals (depending on the plant), this process can be costly and energy intensive for caterpillars. As polyphagous larvae, this process of detoxification and toxin sequestration can be especially costly if their physiology has to support detoxification processes for different types of plants and compounds. Investing more in detoxification as larvae results in lower reproductive output as adults. [10]

Lupine, a common host plant for Arctia plantaginis Necedah Lupine (5447887639).jpg
Lupine, a common host plant for Arctia plantaginis

Adults

P. plantaginis are capital breeders, which means that they do not feed as adults, and thus the larval diet is incredibly important component in adult fitness. [10]

Wood tiger adult Parasemia.plantaginis.jpg
Wood tiger adult

Life history

Males are, on average, smaller than females but experience a relatively similar rate of development. Generally a longer development time correlates with a larger pupal mass, and in females, pupal mass correlates with total lifetime eggs produced. [11]

As a polyphagous species, the life history traits of P. plantaginis depend on its habitat and diet. In P. plantaginis, high anti-oxidant intake from their diet significantly increases their ability to encapsulate pathogens. [11] Encapsulation is an important, innate immune response that occurs in invertebrates to protect against a variety of parasites and pathogens. [12] The antioxidants serve to protect cells from damage incurred by the creation of free radicals resulting from the encapsulation reaction. [13] In environments where the pathogen load is likely to be high, the food ingested by an individual moth is important in building its defense mechanisms. [11]

Enemies

Birds and ants are the most common predators of P. plantaginis, to which the moth has both general and specialized defense mechanisms. [14] The blue tit (Cyanistes caeruleus) is a well known predator.

Selection by predation can impact host immune defense, as demonstrated by an experiment measuring the virulence of a pathogen Serrate marcescens in Arctia plantaginis larvae. Larvae with smaller warning signals had higher survival rates than those with larger warning signals, suggesting that developing a warning signal comes at the expense of immune function. Basically, there is a trade off between immune function and predatory defense. Thus predation is an import factor when considering the evolution of pathogen virulence and host immunity. [15]

Protective coloration and behavior

Aposematism

Aposematism is common in many Lepidoptera species; it is an adaptive mechanism in which prey produce conspicuous warning signals. In the wood tiger moth, conspicuous coloration patterns communicate a poisonous, toxic, or otherwise unpalatable or unprofitable effect to predators. Typically, aposematic species experience strong selection favoring monomorphic populations. As a specific warning signal phenotype becomes more common in an environment, more and more predators learn to avoid individuals bearing such signals. In P. plantaginis, a distinct hindwing pattern of bands and splotches of white or yellow on black warns predators of its chemical defenses. Populations of P. plantaginis, are, however, almost always polymorphic, with males exhibiting varying degrees of either yellow or white melanized banding patterns. Yellow morphs show stronger warning signals and experience lower predation rates and longer predator-hesitation. White morphs are preyed upon significantly more by birds than yellow morphs, but persist as a frequent phenotype in many populations, suggesting there are other selection pressures favoring white morphs. [3]

A possible explanation for the persistence of white morphs despite their higher predation rates is selection heterogeneity; in other words, due to the wide geographic distribution of the wood tiger moth, different populations experience vastly different selective pressures. One consideration is immune variation. A 2013 study demonstrated that male yellow and white larvae saw different survival rates when reared in aggregations; yellow male survived to pupate better in aggregations than white males did, which may reflect different immune investments. In aggregations, white males saw better ability to encapsulate pathogens, while yellow males had higher hemolymph (equivalent to insect 'blood') lytic activity (virus attacking). Thus the two types of wood tiger moth may be maintained in populations because they have different immune investments. This is advantageous in thriving in heterogeneous environments with differential risk factors for immune challenges. [16]

Feigning death

Though the aposematic signal of a wood tiger moth is highly conspicuous against vegetative scenery, its patterning is less easy to detect when it drops to the ground. Disruptive coloration is when a pattern creates an illusion that makes discerning the edges of an object difficult. It essentially destroys the appearance of any outline the object may have. Though the idea of disruptive coloration, which is clearly a camouflage technique, seems counterintuitive in aposematism, it has been demonstrated that the same coloration pattern on a moth can act as either a warning signal or a camouflage depending on the backdrop. Wood tiger moths exhibit a behavior where they essentially 'feign death' by dropping suddenly on to the ground in the presence of a predator and taking on a specific, rigid posture with folded legs. Once on the ground, the moths are much more difficult to detect. This suggests that the hindwing pattern of the wood tiger moth can switch instantly from conspicuous to camouflage, which has obvious adaptive advantages. [17]

Target specific chemical defense mechanisms

A 2017 study highlighted the ability of P. plantaginis to secrete two different chemical fluids as defense mechanisms in response to two different types of predators. Along with its colorful, conspicuous hindwing color patterns, these moths secrete defense fluids from their abdomen and thoracic glands. The abdominal fluids deterred ants and not birds, while thoracic fluids deterred birds but not ants, suggesting that a single species is capable of producing target-specific chemical defense fluids in response to predation threats. [14]

Genetics

Subspecies

Genetics of color patterns

Warning signals show no phenotypic plasticity in adult wood tiger moths. The shapes and patterning of adult warning signals are entirely determined during resource allocation of the larval stage. Once an adult metamorphoses, their warning signal phenotype can no longer change. [18]

Mating

Yellow morphs are able to avoid predation more readily than white morphs; however, a laboratory study showed that yellow males had lower mating success compared to white males. This trade-off between reproductive success and predator avoidance could explain why two polymorphisms exist. [19]

Females tend to attract males during the day, and they group together at dusk. It has been observed that once attracted to a group of females, male P. plantaginis will readily mate with females of a related species Arctica villa, most likely due to their similar sex pheromones. Similarly, female P. plantaginis are also attracted to male Artica villas. [20]

Physiology

Flight

Flight behavior in populations of P. plantaginis vary between color morphs and are under frequency dependent selection. In outdoor cage experiments of populations with various frequencies of yellow and white male morphs, researches found that white morphs were significantly more active and had longer periods of sustained activity than yellow morphs across all treatment groups. In groups with higher frequencies of yellow morphs, overall flying activity for both morphs was considerably lower. The data suggest that white male morphs invest less in producing costly warning signals and thus have more energy to invest in flight for both avoiding predation and finding mates. Yellow males, which in previous studies have been shown to be less sexually favored by females than whites, tend to be most active at peak female-calling periods. [21]

Thermoregulation

Wood tiger moths have a limited amount of resources to allocate to different life history traits and adaptive strategies; thermoregulation is an important part of their physiology, especially in the cooler climates of North America and Eurasia. As latitude increases, populations of P. plantaginis show higher melanization (conversion of resources into melanin). This melanin confers thermoregulatory advantages by increasing a male moth's ability to absorb radiation. This increased melanization comes at a cost, however, as it is costly to produce, and thus male moths with more melanization suffer increased predation as their warning signals are weaker. Thus it is thought that due to the various climatic conditions of populations of wood tiger moth, there are different costs and benefits to produces more melanin, which serves to maintain the global diversity of warning signals that we see throughout the species. In both yellow and white male phenotypes, individuals with more melanin had a heightened ability to trap heat but an increased predation rate due to its weaker and less effective signal. [22]

Related Research Articles

Frequency-dependent selection is an evolutionary process by which the fitness of a phenotype or genotype depends on the phenotype or genotype composition of a given population.

<span class="mw-page-title-main">Poison dart frog</span> Family of amphibians

Poison dart frog is the common name of a group of frogs in the family Dendrobatidae which are native to tropical Central and South America. These species are diurnal and often have brightly colored bodies. This bright coloration is correlated with the toxicity of the species, making them aposematic. Some species of the family Dendrobatidae exhibit extremely bright coloration along with high toxicity — a feature derived from their diet of ants, mites and termites— while species which eat a much larger variety of prey have cryptic coloration with minimal to no amount of observed toxicity. Many species of this family are threatened due to human infrastructure encroaching on their habitats.

<span class="mw-page-title-main">Müllerian mimicry</span> Mutually beneficial mimicry of strongly defended species

Müllerian mimicry is a natural phenomenon in which two or more well-defended species, often foul-tasting and sharing common predators, have come to mimic each other's honest warning signals, to their mutual benefit. The benefit to Müllerian mimics is that predators only need one unpleasant encounter with one member of a set of Müllerian mimics, and thereafter avoid all similar coloration, whether or not it belongs to the same species as the initial encounter. It is named after the German naturalist Fritz Müller, who first proposed the concept in 1878, supporting his theory with the first mathematical model of frequency-dependent selection, one of the first such models anywhere in biology.

<span class="mw-page-title-main">Aposematism</span> Honest signalling of an animals powerful defences

Aposematism is the advertising by an animal to potential predators that it is not worth attacking or eating. This unprofitability may consist of any defenses which make the prey difficult to kill and eat, such as toxicity, venom, foul taste or smell, sharp spines, or aggressive nature. These advertising signals may take the form of conspicuous coloration, sounds, odours, or other perceivable characteristics. Aposematic signals are beneficial for both predator and prey, since both avoid potential harm.

<span class="mw-page-title-main">Garden tiger moth</span> Species of moth

The garden tiger moth or great tiger moth is a moth of the family Erebidae. Arctia caja is a northern species found in the US, Canada, and Europe. The moth prefers cold climates with temperate seasonality, as the larvae overwinter, and preferentially chooses host plants that produce pyrrolizidine alkaloids. However, garden tiger moths are generalists, and will pick many different plants to use as larval host plants.

<span class="mw-page-title-main">Unkenreflex</span>

Unkenreflex – interchangeably referred to as unken reflex – is a defensive posture adopted by several branches of the amphibian class – including salamanders, toads, and certain species of frogs. Implemented most often in the face of an imminent attack by a predator, unkenreflex is characterized by the subject’s contortion or arching of its body to reveal previously hidden bright colors of the ventral side, tail, or inner limb; the subject remains immobile while in unkenreflex.

<i>Arctia</i> Genus of moths

Arctia is a genus of tiger moths in the family Erebidae. Therein, it belongs to the subtribe Arctiina in the tribe Arctiini in the subfamily Arctiinae. Species are well distributed throughout North America, Palearctic, India, and Sri Lanka.

<i>Arctia matronula</i> Moth genus and species

Arctia is species of tiger moth in the family Erebidae. It was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. It can be found in central and eastern Europe, Kazakhstan, southern Siberia, northern Mongolia, Amur Region, Primorye, Sakhalin, Kunashir, northern and northeastern China, Korea and Japan.

<i>Arctia virginalis</i> Species of moths

Arctia virginalis, the Ranchman's tiger moth, is a species of tiger moth in the family Erebidae. It was first described by Jean Baptiste Boisduval in 1852.

<i>Arctia aulica</i> Species of moth

Arctia aulica, the brown tiger moth, is a moth of the family Erebidae. The species was first described by Carl Linnaeus in his 1758 10th edition of Systema Naturae. It is found in the temperate areas of central Europe up to the area surrounding the Amur River to the east and up to the Balkans and the Black Sea to the south.

<i>Arctia alpina</i> Species of moth

Arctia alpina is a moth of the family Erebidae. It is found in northern Scandinavia, northern Siberia, high mountains of southern Siberia and northern Mongolia; also in Alaska and northwestern Canada.

Arctia seitzi is a moth of the family Erebidae. It was described by Andreas Bang-Haas in 1910. It is found in central Asia, including Kazakhstan and Kirghizia.

<span class="mw-page-title-main">Deimatic behaviour</span> Bluffing display of an animal used to startle or scare a predator

Deimatic behaviour or startle display means any pattern of bluffing behaviour in an animal that lacks strong defences, such as suddenly displaying conspicuous eyespots, to scare off or momentarily distract a predator, thus giving the prey animal an opportunity to escape. The term deimatic or dymantic originates from the Greek δειματόω (deimatóo), meaning "to frighten".

Riitta Johanna Mappes is an evolutionary ecologist based in Finland. Her research focuses on interspecific interactions, such as those between predators and prey. She is known for her work on the evolution of aposematic signals and mimicry in chemically defended prey, the evolution of signal polymorphism, the evolution of bacterial virulence, and the evolution of sexual and asexual reproduction. Her main study species include the wood tiger moth, vipers (Viperidae), the Colorado potato beetle and the drumming wolf-spider.

The novel world method is a technique used in animal behaviour experiments that address questions on the evolution of warning signals that chemically defended prey use to deter predators, and also on warning signal mimicry.

Arctia thibetica is a species of tiger moth in the family Erebidae, found in the northwestern Himalayas of India.

Arctia yarrowii, or Yarrow's tiger moth, is a moth of the family Erebidae. It was described by Richard Harper Stretch in 1874. It is found in North America from Hudson Bay to British Columbia and northern Arizona. The habitat consists of barren rocky fellfields and slides above the timberline. These moths are also found in the Pacific Northwest.

Arctia murzini is a moth in the family Erebidae. It was described by Vladimir Viktorovitch Dubatolov in 2005. It is found in Shaanxi, China.

<i>Arctia parthenos</i> Species of moth

Arctia parthenos, the St. Lawrence tiger moth, is a moth in the family Erebidae. It was described by Thaddeus William Harris in 1850. It is found in boreal North America, ranging from Alaska to Labrador, south to New Mexico and Arizona in the Rocky Mountains and to North Carolina in the Appalachian Mountains. The habitat consists of riparian areas and mixed hardwood-conifer forests at middle to high elevations.

References

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