Pyrgus Malvae
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''Pyrgus malvae'', the grizzled skipper, is a
butterfly Butterflies are winged insects from the lepidopteran superfamily Papilionoidea, characterized by large, often brightly coloured wings that often fold together when at rest, and a conspicuous, fluttering flight. The oldest butterfly fossi ...
species from the family
Hesperiidae Skippers are a group of butterflies placed in the family Hesperiidae within the order Lepidoptera (moths and butterflies). They were previously placed in a separate superfamily, Hesperioidea, but have since been placed in the superfamily Papilion ...
. It is a small skipper (butterfly) with a chequered pattern on its wings that appears to be black and white. This butterfly can be found throughout Europe and is common in central and southern regions of England. The butterfly prefers three major types of habitat:
woodland A woodland () is, in the broad sense, land covered with woody plants (trees and shrubs), or in a narrow sense, synonymous with wood (or in the U.S., the '' plurale tantum'' woods), a low-density forest forming open habitats with plenty of sunli ...
,
grassland A grassland is an area where the vegetation is dominance (ecology), dominated by grasses (Poaceae). However, sedge (Cyperaceae) and rush (Juncaceae) can also be found along with variable proportions of legumes such as clover, and other Herbaceo ...
, and industrial. Referenced as a
superspecies In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
, ''Pyrgus malvae'' includes three semispecies: ''malvae'', ''malvoides'', and ''melotis''. Eggs are laid on plants that will provide warmth and proper nutrition for development. As
larvae A larva (; : larvae ) is a distinct juvenile form many animals undergo before metamorphosis into their next life stage. Animals with indirect developmental biology, development such as insects, some arachnids, amphibians, or cnidarians typical ...
, their movement is usually restricted to a single plant, on which they will build tents, unless they move onto a second host plant. Larvae then spin cocoons, usually on the last host plant they have occupied, where they remain until spring. Upon emerging as adult butterflies, grizzled skippers are quite active during the day and tend to favour blue or violet-coloured plants for food. They also possess multiple methods of communication; for example, vibrations are used to communicate with
ant Ants are Eusociality, eusocial insects of the Family (biology), family Formicidae and, along with the related wasps and bees, belong to the Taxonomy (biology), order Hymenoptera. Ants evolved from Vespoidea, vespoid wasp ancestors in the Cre ...
s, and chemical secretions play a role in mating. Exhibiting territorial behaviour, males apply perching and patrolling strategies to mate with a desired female.


Taxonomy/phylogeny

In terms of a
species complex In biology, a species complex is a group of closely related organisms that are so similar in appearance and other features that the boundaries between them are often unclear. The taxa in the complex may be able to hybridize readily with each oth ...
, ''Pyrgus malvae'' is considered a superspecies that consists of three semispecies, which exhibit geographic variations in the genitalia of both male and female butterflies. These three semispecies are considered to be the ''Pyrgus malvae,'' '' Pyrgus malvoides'', and '' Pyrgus melotis'' types. This classification can also be described as a monophyletic clade. Significant isolation mechanisms exist to accentuate the division between the ''malvae'' type and ''melotis'' type, more than the difference between the ''malvae'' and ''malvoides'' types. In fact, interbreeding has been observed between the ''malvae'' and ''malvoides'' types, indicating their close relation - namely that they are both part of the same species.


Description

With its characteristic chequered black-and-white pattern, the grizzled skipper is quite distinctive. It is small, with an average forewing diameter of 12 millimeters, and closely resembles moths in appearance. Males and females can be differentiated by the shape of their wings: males have slightly more angular wings, while females have a more rounded wing shape. Larvae are coloured green and light brown with darker brown stripes. Seitz describes ''malvae'' "The terminal row of white dots developed, at least on the hindwing. Underside of hindwing reddish, with distinct white dots, those of the subterminal band being rounded. Veins bordered with yellowish white or white. In ab. ''taras'' Bergstr.(86a) the white spots of the forewing are united to form bands; occurs singly among ordinary specimens. Europe, Asia from the Mediterranean Sea to the Amur; Mongolia. — Larva yellowish grey, minutely dotted with greenish, the dots bearing short thin hairs, dorsal line darker, spiracles yellowish; in June and October on ''Potentilla'', ''Dipsacus'', Strawberry, Raspberry, and other plants. The butterflies are on the wing in April and May and again from the end of July onwards, on sunny slopes, roads among fields and clearings in woods, being common everywhere in Central Europe.


Geographic range

''Pygrus malvae'' can be found throughout Western Europe in northern Scandinavia, parts of Greece, and some of the Mediterranean Islands. Its populations in many European countries appear to be quite stable. It is also present in Korea, as well as throughout the Mediterranean up to Middle Finland, and rarely in parts of Germany.


Habitat

Although grizzled skippers occupy three major forms of habitats, they tend to settle in environments with spring nectar plants, larval food plants (
agrimony ''Agrimonia'' (from the Greek ), commonly known as agrimony, is a genus of 12–15 species of perennial plant, perennial herbaceous flowering plants in the family Rosaceae, native to the temperate regions of the Northern Hemisphere, with one sp ...
,
creeping cinquefoil ''Potentilla reptans'', known as the creeping cinquefoil, European cinquefoil or creeping tormentil, is a flowering plant in the family Rosaceae. Description A creeping perennial plant which can reach heights of up to 20 cm. Its trailing ...
, wild strawberry,
tormentil ''Potentilla'' is a genus containing over 500 species of annual, biennial and perennial herbaceous flowering plants in the rose family, Rosaceae. Potentillas may also be called cinquefoils in English, but they have also been called five finge ...
), ranker vegetation, and edges with scrub or woodland. Host plants are from the family ''
Rosaceae Rosaceae (), the rose family, is a family of flowering plants that includes 4,828 known species in 91 genera. The name is derived from the type genus '' Rosa''. The family includes herbs, shrubs, and trees. Most species are deciduous, but som ...
'' with a focus on ''
Agrimonia eupatoria ''Agrimonia eupatoria'' is a species of agrimony that is often referred to as common agrimony, church steeples or sticklewort. The whole plant is dark green with numerous soft hairs. The soft hairs aid in the plant's seed pods sticking to any a ...
'' as well as ''
Potentilla ''Potentilla'' is a genus containing over 500 species of Annual plant, annual, Biennial plant, biennial and Perennial plant, perennial herbaceous plant, herbaceous flowering plants in the rose family (biology), family, Rosaceae. Potentillas m ...
.'' Woodland: This mainly consists of sparsely distributed vegetation and can have regions of bare ground that result from cutting or windblow. Grassland: These can result from three different patterns that involve animal grazing, scrub cutting, or disturbance by animals: 1) Scrubby grassland that includes
bramble ''Rubus'' is a large and diverse genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae, most commonly known as brambles. Fruits of various species are known as raspberries, blackberries, dewberries, and bristleberries. I ...
and wild strawberry 2) Unimproved grassland that include creeping cinquefoil 3) Unimproved grassland that includes agrimony. Industrial: Sparse vegetation with mainly wild strawberry or creeping cinquefoil depending on whether the environment is along a railway or clay working. These environments have typically been abandoned fairly recently. Other possible environments for the butterflies are
heath A heath () is a shrubland habitat found mainly on free-draining infertile, acidic soils and is characterised by open, low-growing woody vegetation. Moorland is generally related to high-ground heaths with—especially in Great Britain—a coole ...
land, shingle,
sand dunes A dune is a landform composed of wind- or water-driven sand. It typically takes the form of a mound, ridge, or hill. An area with dunes is called a dune system or a dune complex. A large dune complex is called a dune field, while broad, flat ...
, and acidic, neutral and marshy grassland.


Food resources


Adult food preference

Studies support a concentrated preference for specifically coloured flowers by grizzled skippers. They are most attracted to blue and violet while showing little or no attraction to white, yellow, or red. Butterflies have ultraviolet and blue receptors that may be responsible for Hesperiidae butterflies favoring blue. This evidence indicates that when butterflies from this species forage for food, they are particularly attentive to short-wavelength light that is reflected off flowers. In fact, this particular preference aligns with a prominent attentiveness that members of Hesperiidae have for blue. This preference may be a result of phylogenetic adaptations, foraging signals, and learning abilities. Specifically, the prevalence of blue flowers in lowlands could further intensify this preference, especially for grizzled skippers that tend to be found in lowland grasslands.


Parental care


Host plant preference/selection

An overarching theme in
behavioral ecology Behavioral ecology, also spelled behavioural ecology, is the study of the evolutionary basis for ethology, animal behavior due to ecology, ecological pressures. Behavioral ecology emerged from ethology after Niko Tinbergen outlined Tinbergen's f ...
can be seen through female grizzled skipper investment in host plant selection. Females tend to lay eggs on host plants that are viewed as larger and more nutritionally rich. However, this nutritional advantage for caterpillars must be balanced with the presence of a warm microclimate that is suitable for the species. Warm microclimates align with ''P. tabernaemontani'' plants, but these may also have an increased chance of desiccation. On the other hand, ''A. eupatoria'' is a larger plant when near
molehill A molehill (or mole-hill, mole mound) is a conical mound of loose soil raised by small burrowing mammals, including moles, but also similar animals such as mole-rats, and voles. The word is first recorded in the first half of the 15th century. ...
s and can be found in more suitable environments. As a result, ''A. eupatoria'' provides an appropriate balance for both of these requirements and is preferred as a host plant by ''P. malvae,'' particularly near molehills. Females consider optical conspicuousness, or plant visibility, through prominence (height differences between host plants and vegetation). More prominent host plants, like ''A. eupatoria'' over ''P. tabernaemontani'', are favored. However, these two different habitats are used significantly and may be evolutionary adaptations to offset the grizzled skippers’ risk of extinction imposed by a polarized weather pattern.


Life history


Egg

Eggs are mainly found on agrimony, creeping cinquefoil, and wild strawberry plants, which provide nourishment for larvae. Other plants that can be considered include barren strawberry, tormentil,
salad burnet ''Sanguisorba minor'', the salad burnet, garden burnet, small burnet, burnet (also used for ''Sanguisorba'' generally), :wiktionary:pimprenelle, pimpernelle, Toper's plant, and burnet-bloodwort, is an edible perennial plant, perennial herbaceou ...
,
bramble ''Rubus'' is a large and diverse genus of flowering plants in the rose family, Rosaceae, subfamily Rosoideae, most commonly known as brambles. Fruits of various species are known as raspberries, blackberries, dewberries, and bristleberries. I ...
,
dog rose ''Rosa canina'', the dog rose, is a variable climbing, wild rose species native to Europe, northwest Africa, and western Asia. Description The dog rose is a deciduous shrub normally ranging in height from , though it can scramble higher into the ...
, and
wood avens ''Geum urbanum'', also known as wood avens, herb Bennet, colewort, clove root and St. Benedict's herb (Latin: ''herba benedicta''), is a perennial plant in the rose family (Rosaceae), which grows in shady places (such as woodland edges and near ...
. Plants favored by mothers are typically located in bare ground or short vegetation environments, can have high nitrogen contents, and tend to be located in warmer microclimates. The eggs for grizzled skippers are laid one at a time, although females can lay them on more than two species of plants. Preferred plants have been observed to contain up to 22 eggs.


Larva

This stage continues for two to three months in its entirety. At first, larvae live and spend the vast majority of their time within ‘tents’ that they build on the host leaf, and this partially feeds into their restricted mobility. The two reasons for leaving this shelter are 1.) to feed on another leaf, or 2.) to create another tent. Later on in this stage, larvae expand their mobility and are no longer restricted in diet or habitat. Plants that are higher in nitrogen content and nutrition are more highly incorporated into the diets of larger larvae, as are coarser shrubs. Host plants located on molehills are favored for early larval stages due to open vegetation that is available as well as bare ground and warmer microclimates.


Pupa

Pupal cocoons spun by larvae can be located on or within low vegetation. Generally, these cocoons are not found near the larvae's last tent. The location of these cocoons can consequently affect adult emergence patterns. Pupal cocoons on shorter vegetation will facilitate adult emergence earlier in the spring than pupal cocoons on longer vegetation.


Adult

Grizzled skippers produce one brood per season and are in flight from the middle of March to the middle of July. Active during the day and focused on feeding or basking, they roost on tall vegetation like
marjoram Marjoram (, ''Origanum majorana'') is a cold-sensitive perennial plant, perennial herb or undershrub with sweet pine and citrus flavours. In some Middle Eastern countries, marjoram is synonymous with oregano, and there the names sweet marjoram ...
,
knapweed ''Centaurea'' () is a genus of over 700 species of herbaceous thistle-like flowering plants in the family Asteraceae. Members of the genus are found only north of the equator, mostly in the Eastern Hemisphere; the Middle East and surrounding ...
, and St John's-wort. In particular, adults spend days basking on bare ground and afternoons on taller vegetation. This provides them favorable microclimates and visibility.


Migration


Location or regional dispersal

Butterflies move away from open spaces to the case of increasing wind speeds to seek shelter. Shelter may be particularly important because of vegetation; for example, they can support herbs and bramble. They are also useful in connecting multiple separated habitats. In fact, shelters are associated with higher activity amongst the butterflies.


Enemies


Predators

Grizzled skippers belong in family Hesperiidae, but another group of butterflies (family
Lycaenidae Lycaenidae is the second-largest family (biology), family of butterflies (behind Nymphalidae, brush-footed butterflies), with over 6,000 species worldwide, whose members are also called gossamer-winged butterflies. They constitute about 30% of ...
) uses the mechanism of wing expansion to produce vibrations and communicate with ants. Similar to this behavior, grizzled skippers also produce vibrations upon expanding their wings. This resembles the ‘ant-attendant’ behaviour that is specific to butterflies in the Lycaenidae. Ants are recognized as potential predators for lycaenid caterpillars, therefore, vibration signaling most likely functions as an anti-predatory survival mechanism. Similarly, grizzled skippers also exhibit this kind of vibratory communication. Although the behavior is absent during the caterpillar stage, grizzled skippers are able to produce vibrations upon wing expansion. Akin to lycaenids, grizzled skippers may use these vibrations as signals to communicate with ants and potentially as an attempt to temper aggression.


Mating behaviour


Female/male interactions


Pheromones

Grizzled skippers are known to contain organs called
androconia The external morphology of Lepidoptera is the physiological structure of the bodies of insects belonging to the order Lepidoptera, also known as butterflies and moths. Lepidoptera are distinguished from other orders by the presence ...
that are responsible for producing chemicals within the
thorax The thorax (: thoraces or thoraxes) or chest is a part of the anatomy of mammals and other tetrapod animals located between the neck and the abdomen. In insects, crustaceans, and the extinct trilobites, the thorax is one of the three main di ...
as well as the
abdomen The abdomen (colloquially called the gut, belly, tummy, midriff, tucky, or stomach) is the front part of the torso between the thorax (chest) and pelvis in humans and in other vertebrates. The area occupied by the abdomen is called the abdominal ...
. They are found in males at the forewing costal fold. The organs release
sex pheromone Sex pheromones are pheromones released by an organism to attract an individual of the same species, encourage them to mate with them, or perform some other function closely related with sexual reproduction. Sex pheromones specifically focus on ind ...
s that can be used as a sexual recognition mechanism and drive evolution. The process involves the male first locating a female visually, then using low concentrations of the pheromones at a relatively close proximity to indicate its viability as a mate. While courting the female, male butterflies have specialized hairlike structures called ‘tibial tufts’ on their hind legs that can be used to steer these chemicals directly to the female. However, this chemical communication cannot be differentiated between ''P. malvae'' and ''P. malvoides'', a close relative considered a subspecies of ''P. malvae'' and not separated by isolation reproductive barriers''.''


Courting

There are two main mating strategies that are used by grizzled skippers, which ultimately illustrate mechanisms of
sexual selection Sexual selection is a mechanism of evolution in which members of one sex mate choice, choose mates of the other sex to mating, mate with (intersexual selection), and compete with members of the same sex for access to members of the opposite sex ...
by this species. In both, males demonstrate territorial behaviours. In the case of perching, males wait on taller plants for females to come to them in order to begin courting. In the case of patrolling, males locate a desired female and fly down to her. The type of behaviour that they use in mating depends upon the pattern of food plant availability in their respective habitats. * Perching Approach: This is used in habitats that are poor for food plants. Males will engage in more of a perching approach where they await females above scrub edges that afford them with shelter, warmth, and visible range. * Patrolling Approach: This is used in habitats that are rich for food plants. Males will engage in more of a patrolling strategy by mingling with females.


Climate effects on distribution

''Pyrgus malvae'' is particularly receptive to warmer and drier climates. Warmer summers are more favorable for the grizzled skipper and are positively correlated with the species. This could result from relationships between warmth and the success of the mother's ability to lay the egg as well as of larvae survival. Warmer climates also tend to hasten the development of larvae and allow for earlier onset of the pupal stage. Cooler northern weather may explain the concentration of this butterfly in southern regions. As a result of climate warming, the grizzled skipper appears to show decreases in northern range margin, distribution area, and abundance. This response appears to be dependent upon climatic as well as nonclimatic driving forces.


Physiology


Wing structure effect on flight behaviour

The
wings A wing is a type of fin that produces both lift and drag while moving through air. Wings are defined by two shape characteristics, an airfoil section and a planform. Wing efficiency is expressed as lift-to-drag ratio, which compares the bene ...
of ''Pygrus malvae'' are only mildly adhesive, rough, and highly hydrophobic. These qualities prevent their wings from sticking to external surfaces, support their resistance to water damage, and allow the butterfly to self-clean its wings without risking desiccation. Additionally, these properties enable the grizzled skipper to avoid excess weight bearing, consequently promoting secure and efficient flight patterns.


Conservation


Concerns


Past

1- Decreased woodland clearings, unimproved grassland, scrub habitats 2- Unimproved grasslands and scrub habitats that have been changed 3- Woodland and grassland sites that have been broken apart or deserted 4- Artificial habitats that have been deteriorated


Current/Future

1- Further changes that reduce open areas and spread apart clearings 2- Reductions in coppice produce 3- Further deteriorating unimproved grassland, scrub, and artificial habitats 4- Further breaking apart and deserting habitats


Prevalence decline

The grizzled skipper has notably been decreasing in its prevalence in several other European countries over the past 25 years. The species has experienced significant declines (ranging from 25% to over 50%) in
Britain Britain most often refers to: * Great Britain, a large island comprising the countries of England, Scotland and Wales * The United Kingdom of Great Britain and Northern Ireland, a sovereign state in Europe comprising Great Britain and the north-eas ...
,
Croatia Croatia, officially the Republic of Croatia, is a country in Central Europe, Central and Southeast Europe, on the coast of the Adriatic Sea. It borders Slovenia to the northwest, Hungary to the northeast, Serbia to the east, Bosnia and Herze ...
,
Belgium Belgium, officially the Kingdom of Belgium, is a country in Northwestern Europe. Situated in a coastal lowland region known as the Low Countries, it is bordered by the Netherlands to the north, Germany to the east, Luxembourg to the southeas ...
,
European Turkey East Thrace or Eastern Thrace, also known as Turkish Thrace or European Turkey, is the part of Turkey that is geographically in Southeast Europe. Turkish Thrace accounts for 3.03% of Turkey's land area and 15% of its population. The largest c ...
, and the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
. Less severe decline can be seen in the
Czech Republic The Czech Republic, also known as Czechia, and historically known as Bohemia, is a landlocked country in Central Europe. The country is bordered by Austria to the south, Germany to the west, Poland to the northeast, and Slovakia to the south ...
,
Germany Germany, officially the Federal Republic of Germany, is a country in Central Europe. It lies between the Baltic Sea and the North Sea to the north and the Alps to the south. Its sixteen States of Germany, constituent states have a total popu ...
,
Denmark Denmark is a Nordic countries, Nordic country in Northern Europe. It is the metropole and most populous constituent of the Kingdom of Denmark,, . also known as the Danish Realm, a constitutionally unitary state that includes the Autonomous a ...
,
Finland Finland, officially the Republic of Finland, is a Nordic country in Northern Europe. It borders Sweden to the northwest, Norway to the north, and Russia to the east, with the Gulf of Bothnia to the west and the Gulf of Finland to the south, ...
,
Luxembourg Luxembourg, officially the Grand Duchy of Luxembourg, is a landlocked country in Western Europe. It is bordered by Belgium to the west and north, Germany to the east, and France on the south. Its capital and most populous city, Luxembour ...
,
Romania Romania is a country located at the crossroads of Central Europe, Central, Eastern Europe, Eastern and Southeast Europe. It borders Ukraine to the north and east, Hungary to the west, Serbia to the southwest, Bulgaria to the south, Moldova to ...
,
Sweden Sweden, formally the Kingdom of Sweden, is a Nordic countries, Nordic country located on the Scandinavian Peninsula in Northern Europe. It borders Norway to the west and north, and Finland to the east. At , Sweden is the largest Nordic count ...
,
Slovakia Slovakia, officially the Slovak Republic, is a landlocked country in Central Europe. It is bordered by Poland to the north, Ukraine to the east, Hungary to the south, Austria to the west, and the Czech Republic to the northwest. Slovakia's m ...
, and Asian Turkey. Once found across England as well as regions in
Wales Wales ( ) is a Countries of the United Kingdom, country that is part of the United Kingdom. It is bordered by the Irish Sea to the north and west, England to the England–Wales border, east, the Bristol Channel to the south, and the Celtic ...
, the grizzled skipper is now concentrated in Central and Southern England only. In fact, five of England's counties have already experienced extinction of the species, and the butterfly can only be found in less than five sites in another five English counties. However, between 1995 and 1997, evidence of recolonization by the grizzled skipper has emerged, specifically in several counties in England.


Grazing effect on population

Grazing has been applied as an approach to conservation, especially in the
Netherlands , Terminology of the Low Countries, informally Holland, is a country in Northwestern Europe, with Caribbean Netherlands, overseas territories in the Caribbean. It is the largest of the four constituent countries of the Kingdom of the Nether ...
where it is used to monitor the biodiversity of open habitats. ''Pyrgus malvae'' is important in these studied because it is considered as a threatened to susceptible species in the Netherlands. Grazing appears to be positively associated with populations of ''Pyrgus malvae'' that are concentrated more in habitats with grazed vegetation rather than those with cut and ungrazed regions. The trend may be reflective of the fact that grizzled skippers prefer open grassland habitats. With its endangered status in the Netherlands, the grizzled skipper may benefit from extensive grazing as a nature restoration project.


See also

*
List of butterflies of Great Britain This is a list of butterfly, butterflies of Great Britain, including extinct, naturalised species and those of dubious origin. The list comprises butterfly species listed in ''The Moths and Butterflies of Great Britain and Ireland'' by Emmet ''et a ...


References


Literature

* * Brereton, T. M.; Bourn, N. A. D., and Warren, M. S. (1998).
Species action plan. Grizzled Skipper
'. * Dover, J. W., T. H. Sparks, and J. N. Greatorex-Davies. "The importance of shelter for butterflies in open landscapes." ''Journal of Insect Conservation'' 1.2 (1997): 89–97. * Elfferich, Nico W. "Is the larval and imaginal signalling of Lycaenidae and other Lepidoptera related to communication with ants." ''Deinsea'' 4.1 (1998): 91–96. * Fang, Yan. "Anisotropic Characteristic of Insect (Lepidoptera) wing Surfaces." (2015). * Hammond, Nicholas, et al. ''Concise Butterfly & Moth Guide''. Bloomsbury, 2014. * Hernández-Roldán, Juan L., et al. "Morphological and chemical analysis of male scent organs in the butterfly genus Pyrgus (Lepidoptera: Hesperiidae)." ''Organisms Diversity & Evolution'' 14.3 (2014): 269–278. * Jong, Rienk. ''Superspecies Pyrgus malvae (Lepidoptera: Hesperiidae) in the East Mediterranean, with notes on phylogenetic and biological relationships''. Rijksmuseum van Natuurlijke Historie, 1987. * Kociková, Lenka, et al. "Is colour an important factor influencing the behaviour of butterflies (Lepidoptera: Hesperioidea, Papilionoidea)?." ''European Journal of Entomology'' 109.3 (2012): 403. * Krämer, Benjamin, et al. "Microhabitat selection in a grassland butterfly: a trade-off between microclimate and food availability." ''Journal of Insect Conservation'' 16.6 (2012): 857–865. * Pollard, E. "Temperature, rainfall and butterfly numbers." ''Journal of Applied Ecology'' (1988): 819–828. * Roy, David B., et al. "Butterfly numbers and weather: predicting historical trends in abundance and the future effects of climate change." ''Journal of Animal Ecology'' 70.2 (2001): 201–217. * Streitberger, Merle, and Thomas Fartmann. "Molehills as important larval habitats for the grizzled skipper, Pyrgus malvae (Lepidoptera: Hesperiidae), in calcareous grasslands." ''European Journal of Entomology'' 110.4 (2013): 643. * Wallisdevries, Michiel F., and Ivo Raemakers. “Does Extensive Grazing Benefit Butterflies in Coastal Dunes?” ''Restoration Ecology'', vol. 9, no. 2, 2001, pp. 179–188., doi:10.1046/j.1526-100x.2001.009002179.x. *


External links


Lepiforum.de

''Pyrgus malvae'' at Caterpillar Hostplants Database
{{Taxonbar, from=Q1109803 Pyrgus Butterflies of Asia Butterflies of Europe Butterflies described in 1758 Taxa named by Carl Linnaeus