September Moths

Below are some adult moths (and one caterpillar) that are active during September. ‘Foodplant’ and ‘breeding plant ‘refers to the plant eaten by the caterpillar. The following moth species are nocturnal.

All images J. Harding

This is the larva of the Pale Tussock moth. This larva can often be found on or near the ground in September. The tuft of pink hair marks the location of the tip of the abdomen. It pupates under or near the tree it fed on. The foodplants are varied and include native trees and shrubs such as birch, oak, and Common Hazel. The adult moth flies in May and June. The moth comes to light and does not feed, relying on fat stored by the larva.
Rosy Rustic. This light-attracted moth flies from August to October. It breeds on Broad-leaved Dock, Ribwort Plantain, horsetails, and Flag Iris.
Beaded Chestnut is a very numerous moth in wooded habitats, including gardens, during September. Its foodplants are buttercups, clovers, chickweeds, and, when larger, broadleaved trees and shrubs, especially Common Hawthorn. This moth comes to light in abundance.
Pink-barred Sallow. This has a short flight time, appearing in September and October. The adult comes to light and feeds on Common Ivy and over-ripe blackberries. The breeding plants are willows and poplars, and later on docks.
The Lunar Underwing is an abundant species. The adult flies mainly in September and October and takes nectar from ivy and sugar from over-ripe blackberries. It is named for its moon-white underwing, appropriately ‘clouded’ with hazy sooty markings, redolent of faint cloud partly obscuring the moon. It appears in some numbers at light traps, and its ground color varies from pale yellow to grey. The breeding plants are Yorkshire Fog and other grasses. 
The Willow Beauty has at least two generations, the first June-August the second from late August to October. Alder Buckthorn, birches, Common Yew, and Scots Pine are among its wide range of foodplants. It likes woods and mature gardens. The adult moth comes to light and enjoys nectar from Creeping Thistle and Common Ragwort.
Ripe and over-ripe blackberries are eagerly sought by September moths and butterflies.
Moths are a major prey item for birds, especially familiar garden birds like the Wren, Robin, and Blue Tit. Birds learn to forage for moths early in the morning in areas where outdoor lights are left on at night.

Calm, mild September nights often yield large populations of moths. A car journey home at around 10 pm along our rural hedged roads will often give an indication of the level of abundance, with ghostly wings flitting in and out of view in the car lights. A glance upward at hedge-top height might reveal bats that will dip down to snatch a moth meal. Some moths are disorientated by light and will flutter against the base of hedges, kerbs, or walls that the light falls on. If a car is parked with lights on for some minutes, these moths can fall prey to frogs and hedgehogs emerging from cover nearby.

Moths are very important food web components. A healthy moth population is a sign of environmental health. When you see plenty of moths in your headlights, feel good about the area you are in!

 

 

What to look for in September

September is an in-betweener, no longer summer but with some days bringing summer heat, not winter but with some sharp cold at night. The blooms of summer have mainly faded but new flowers offer sustenance to September moths and butterflies. Some fruits are available too; Rowan, bramble, and Elder offer their sweetness to species still in flight.

Oddly, September is a peak month for some species: Red Admiral, Comma and Speckled Wood are abundant in September.  Yesterday I counted 54 Speckled Woods in the rather cool, breezy Russellstown Wood, near Russborough House, in County Wicklow.  The butterfly is also extremely abundant in the scrublands of the Burren, where its copiousness astonishes, an apparently seasonally discordant richness.

Speckled Wood eggs being laid now are unlikely to produce pupae before winter, but the larvae will pupate next spring or early summer depending on development rates. The resulting adults are likely to fly in May and June after their aunts and uncles have flown in March, April, and early May. This prolonged emergence arising from individuals from more than one generation from the previous year provides a very long first brood, running from later in March to the end of June.  A gap occurs in the flight of this species during July, with few flying in this month. However, from August lasting well into September and even October, there is an extended emergence resulting from larvae produced during April-July. According to Thomas and Lewington (2014) not all spring larvae produce second-generation butterflies. Some develop slowly and form over-wintering pupae in autumn.

Speckled Wood in Russellstown Wood, Co. Wicklow.

The understanding of the Speckled Wood’s complex brood structure comes from studies done in Britain. Perhaps this applies to our Speckled Woods, but until breeding studies are undertaken in Ireland we cannot confirm this. While our records of the adult butterfly suggest that the butterfly has a similar brood structure here, when I reared the species from eggs laid in late June, all of these reached adulthood in late August and during September. If some spring larvae do not produce butterflies in the same year, do some summer larvae undergo slower development, producing over-wintering larvae or over-wintering pupae in autumn?

I suspect that there is more to learn about the Speckled Wood’s brood structure in Ireland, potentially including regional differences in its development rates.

Another species with a complex brood structure is the Comma, and in parts of Ireland where it is established, it is present in good numbers, feeding on autumn fruits and late flowers to store up fat for its long sleep. Unlike the Speckled Wood which over-winters as a larva or pupa, the Comma passes the autumn and winter in just one life stage, the adult stage.  With its dark, bark-coloured undersides and scalloped oak-leaf outline, it is perfectly placed to conceal itself in woods, on tree trunks, and among heaped leaf litter. By contrast, the bright orange uppersides now illuminate hedged country lanes, wood edges, clearings, and gardens. When a vivid example is perched on blackberries, the luminosity is striking. When the sun is obscured and sometimes when it detects approach, it shuts its wings and becomes a dead leaf trapped in the bramble.

A male Comma feeding on a blackberry, Mulhussey, Co. Meath.
A male Comma underside. The female Comma’s underside (see below) is much less variegated and may indicate a different choice of over-wintering site than that used by the male.
A female Comma feeding on Common Ivy.

The tiny Small Copper is still adorning our grasslands and the warm sunny August and pleasant September might see the third generation later this month. Two broods are seen each year with the third generation weather-dependent. Today (September 17th) I saw the pristine female shown in the photograph below. Shortly after I took the photograph, she was courted by two males. Her offspring will fly next year’s first generation, typically in May and June.

A newly emerged female Small Copper, Mulhussey, Co. Meath.

Another gorgeous butterfly, the Red Admiral, is present in our countryside and gardens, and in some abundance in flower-rich areas.  It has a broad taste, taking advantage of nectar, fruit juice, and tree sap. By contrast with the previous butterflies profiled here, this will leave our shores this autumn to breed further south, in Europe. A few stay behind to breed in coastal areas but this applies to a small number of individuals, as far as is known.

For now, enjoy this black and scarlet butterfly. It will fly throughout September and October, relying increasingly on ivy nectar as other food sources disappear.

Red Admiral basking on birch, Lullybeg, Co. Kildare.

The Small Tortoiseshell rounds off this article. This bright, friendly-looking butterfly is busy feeding up to spend the winter as an adult butterfly. Unlike its relative, the Comma, the Small Tortoiseshell will seek out buildings as well as natural places to over-winter. Like the Comma, its undersides are muted in colour blending it with the dark places chosen for over-wintering. All over-wintering butterflies close their wings when sleeping, so colourful uppersides are not on show. Like all species that over-winter as adults, the Small Tortoiseshell does not enter this state synchronously; rather individuals enter the over-wintering state over several weeks, so the individuals seen at any time during August-October represent those yet to close their wings for the year. This elucidates the paradox of seeing more individuals in spring than appear to have been active during the previous late summer and autumn.

If you are lucky to host a Small Tortoiseshell this autumn or winter, do not release it. Relocate any woken by the central heating to a cool place such as a shed or even a kitchen roll-lined Tupperware box with the lid on, placed in the fridge (never the freezer). Release the butterfly in warm, sunny weather in March. The individual in the photo below has traces of gossamer on its wings resulting from an encounter with a spider’s web. It overcame this survival challenge. Will it survive our long, dark foodless winter? For Small Tortoiseshells that take refuge behind our cupboards, wardrobes, picture frames, mirrors, and within curtain folds, that is for you to decide…

A Small Tortoiseshell on Rough Hawk-bit, Lullymore, Co. Kildare.

All photos by Jesmond Harding.

Key Reference

Thomas, J. & Lewington, R. (2014) The Butterflies of Britain and Ireland. (Revised edition) British Wildlife Publishing, Dorset.

 

The Importance of Scrub and Grassland Mosaics for the Butterflies and Moths of the Burren

Clooncoose Valley, County Clare.

The Burren is a region in North County Clare and South Galway. It is a karst landscape containing exposed limestone. The Burren as it appears today is mainly the result of the last glaciation and the impact of human activity, especially farming. It is a remarkable landscape, containing a variety of habitats that belies the seeming uniformity of appearance when viewed from a distance, particularly during winter. In addition to eye-catching geological features, the area has a range of habitats that add enormously to the region’s biodiversity and international reputation. This article focuses mainly on the value of scrub and limestone grassland for butterflies and moths, particularly in areas where these habitats exist close together.

Scrub is a broad term and includes areas that are at least 50% covered by shrubs, stunted trees or brambles. The term includes dense growth with little ground vegetation. The canopy height is usually below 5m. The scrub of most value to butterflies is open scrub on lightly grazed, herb-rich grassland that receives good sunlight. Scrub management is carried out by occasional burning, manual cutting or cutting using machinery, sometimes followed by herbicide application. The most important habitat in Ireland for butterflies is the open scrub on calcareous grassland in the Burren in County Clare and Galway. The main scrub components present in the Burren are Common Hazel Corylus avellana, Common Hawthorn Crataegus monogyna, Common Blackthorn Prunus spinosa, Bramble Rubus fruticosus agg. and erect or scrambling roses Rosa spp., in addition to several willows Salix spp., Ling Calluna vulgaris, Guelder Rose Viburnum opulus, Common Holly Ilex aquifolium, occasional Juniper Juniperus communis and Common Yew Taxus baccata.

Fahee North, County Clare. The closely cropped grassland is the result of year-round grazing on this site. The scrub and grassland here provide excellent habitats for rare Lepidoptera.

The grassland in the Burren clearings is often rich in flora, especially clovers Trifolium spp., violets Viola spp., Common Knapweed Centaurea nigra, Devil’s-bit Scabious Succisa pratensis, Selfheal Prunella vulgaris, Common Bird’s-foot Trefoil Lotus corniculatus, Cat’s-ear Hypochoeris radicata, Lady’s Bedstraw Galium verum and Oxeye Daisy Leucanthemum vulgare. The more calcareous grasslands are characterised by broadleaved herbs such as Cowslip Primula veris, Greater Knapweed Centaurea scabiosa, Kidney Vetch Anthyllis vulneraria, Mountain Everlasting Antennaria dioica, Yellow-wort Blackstonia perfoliata, Salad Burnet Sanguisorba minor and Carline Thistle Carlina vulgaris, and may also be important for orchids, including Ophrys and Orchis spp.

Flower-rich habitat (dry calcareous grassland) at Clooncoose, County Clare.

Grasses present often include fescues Festuca spp., Sweet Vernal-grass Anthoxanthum odoratum, Crested Dog’s-tail Cynosurus cristatus, Cock’s-foot Dactylis glomerata and Yorkshire-fog Holcus lanatus. Grasses that are indicative of strongly calcareous soils include Downy Oat-grass Avenula pubescens, Yellow Oat-grass Trisetum flavescens, Blue Moor-grass Sesleria caerulea and Quaking-grass Briza media.

These scrub and grassland mosaics are used for breeding by scrub/woodland and grassland butterflies. The butterfly species recorded breeding on herbs and grasses in open scrub in the Burren are Dingy Skipper Erynnis tages, Wood White Leptidea sinapis, Common Blue Polyommatus icarus, Small Tortoiseshell Aglais urticae, Peacock Aglais io, Red Admiral Vanessa atalanta, Painted Lady Vanessa cardui, Pearl-bordered Fritillary Boloria euphrosyne, Dark Green Fritillary Speyeria aglaja, Silver-washed Fritillary Argynnis paphia, Marsh Fritillary Euphydryas aurinia, Speckled Wood Pararge aegeria, Wall Brown Lasiommata megera, Grayling Hipparchia semele, Meadow Brown Maniola jurtina, Ringlet Aphantopus hyperantus and Small Heath Coenonympha pamphilus.

The species recorded breeding on shrubs in open scrub in the Burren are Brimstone Gonepteryx rhamni, Brown Hairstreak Thecla betulae, Purple Hairstreak Favonius quercus, and Holly Blue Celastrina argiolus.

Separating the species breeding in open scrub into (a) herb/grass feeders and (b) species feeding on shrubs ignores the complex habitat associations of several of the species listed here. The shade, shelter, and leaf litter provided by scrub influence conditions in the grasslands. This is vital for Lepidoptera, especially for the life stages lasting several months. The role played by scrub varies according to species, the time of year, and features such as the height, structure, composition, and management of the scrub and sward.

The Marsh Fritillary is a locally distributed, generally scarce species that requires grassland containing at least a 25% density of the larval foodplant Devil’s-bit Scabious, with the plants growing close together. The grassland that is most favoured has a sward height of 12-25cm. Some occupied swards are taller and even quite rank, but the foodplant is unshaded. The sward usually has tussock-forming grasses and flowers. There is usually some light cattle or horse grazing to maintain the grassland. In swards that are strongly exposed to the weather and that are moderately grazed during the summer, some sheltering scrub is important for the Marsh Fritillary. This species dislikes exposed grasslands unless there is a tall, (c.25cm) well-developed sward that is grazed irregularly and extensively or throughout the year if grazed extensively. Litter from easily warmed dead vegetation, especially from grasses, but sometimes from mosses, scrub foliage, and bracken, must be present for the Marsh Fritillary to be able to make use of the sward for breeding.

The Dark Green Fritillary enjoys superabundance in several areas in the Burren.

The scrub is also used by the Dark Green Fritillary. This eye-catching, dynamic species flies mostly from mid-June to mid-August. Eggs are laid on or near violets growing in grassy places with a fairly tall, well-developed sward, in open, exposed grassland, in sunlit clearings in open scrub, but also at the immediate edge of scrub where a lower, patchier sward exists. After about two weeks, the newly hatched larva eats the eggshell and immediately enters diapause (a rest phase with no feeding and little development), remaining in this state until March of the following year. Unless it has the shading effect of a well-developed sward or scrub, the unfed larva will desiccate in the summer heat.

A Wood White laying an egg on Bitter Vetch Lathyrus linifolius close to scrub, Knockaunroe, Co. Clare.

The Wood White has a highly restricted distribution in Ireland, where it has been found only on carboniferous limestone, from near Newmarket-on-Fergus in south County Clare northwards to Lough Corrib, County Galway on its east and west shores. Its stronghold is the Burren. It is scrub-dependent, breeding on vetches, especially Meadow Vetchling and Tufted Vetch, that receive direct sunlight but that grow in intimate contact with scrub. The adult has a weak, fluttering flight, and the shelter afforded by scrub appears necessary for it. The larva needs access to direct sunlight, shelter, and shade throughout its development. When feeding is complete, it pupates in the scrub or on vegetation growing among the scrub. Medium-tall vegetation in intimate contact with scrub is crucial for this species.

In Ireland, the Pearl-bordered Fritillary breeds on Common Dog-violet Viola riviniana growing among open scrub and open woodland on the carboniferous limestone in the Burren, Counties Clare and Galway, and in similar habitat near the Burren. In Ireland, it is restricted to these areas. According to Nash et al (2012), it has been recorded in just twelve 10 km squares, 35 tetrads. In two of the 10km squares, on Inishmore and Inishmaan, where single individuals were seen, it appears that these were wanderers from the mainland.

The breeding habitat for the Pearl-bordered Fritillary is the same as that used by the Wood White. The larva requires shade and direct sunlight. Direct light is required by the larva for it to be active and for digestion while the shade is needed to cool it down. During hot sunny weather in summer, the larva feeds in strong sunlight but quickly retreats to a shaded location. Cooling shade is needed during the larva’s long diapause from mid to late summer until feeding resumes in March. Furthermore, unshaded violets wilt and turn yellow, which makes make them unsuitable. In part-shaded areas, the foliage remains green.

A male Pearl-bordered Fritillary basking on limestone at Fahee North, Co. Clare. During overcast conditions, the exposed stone helps this species to maintain its body temperature.

The question of why the Pearl-bordered Fritillary’s distribution in Ireland is limited to the Burren is an interesting one, with the answer likely to be found in the conditions needed by the larva. It needs the Common Dog-violet, usually with low herbs and dry moss on dry, free-draining, warm, sheltered, usually south-facing sunny habitats containing leaf litter build-up against scrub on open limestone rock. The larval micro-sites have low grass density; the grasses present are typically scattered tufts of Blue Moor-grass and fescue grasses containing standing and lying warm, dry litter, particularly in spring. Plant litter is used by larvae for basking, concealment, and over-wintering. It may be important for the larva to have Common Hazel leaf litter. Hazel produces a leaf litter that is substantial without shading the foodplant. Dead hazel leaves curl, producing hiding places for resting larvae. The dead leaf dries quickly, and the larva needs dry conditions (Harding 2021).

Two other features favouring the species are the frequent rainfall that prevents the foodplant from losing moisture while the free-draining limestone provides the dryness needed by the larva. Grazing by cattle or horses during winter and spring, and periodic scrub cutting, but not scrub clearance, is beneficial.

The adult feeds mainly in the clearings in the scrub, especially on Common Bird’s-foot-trefoil, Bugle Ajuga reptans, buttercups Rununculus spp., and Dandelion Taraxacum spp. Thus, the butterfly needs the grassy areas adjoining the scrub, as well as the shadier areas occupied by scrub.

Pearl-bordered Fritillary breeding habitat near Clooncoose, Co. Clare. This photo was taken in early May.

The Peacock, a species distributed throughout Ireland, uses the Stinging Nettle Urtica dioica as a foodplant, and it selects unshaded nettles growing close to scrub or woodland. It is possible that the scrub provides the warmth and shelter required for larval development. Fully fed larvae leave nettles to pupate on trees and scrub.

Speckled Wood, another widespread species, has a complex relationship with scrub. Known as a butterfly of hedges, scrub and woodland, it is our most shade-tolerant butterfly. The adult uses scrub for food (feeding on flowers such as bramble and Common Ivy Hedera helix, on ripe blackberries and aphid ‘honeydew’), for shelter from the rain and heat, resting and roosting, territorial perches, patrolling areas, mating stations, and dispersal routes. The butterfly breeds on native grasses growing at the edge of scrub, hedges, and woodland, but a seasonal difference exists in the egg-laying sites chosen. Early and late in the year, warm clumps of grasses in full sun are selected for oviposition, while during the summer heat, partly shaded lusher grasses in humid locations are selected. However, a female will adjust her choice of egg site according to temperature, so that grasses in shaded positions will be selected during hot weather in autumn.

Four butterfly species that breed on shrubs, Brimstone, Brown Hairstreak, Purple Hairstreak, and Holly Blue, use foodplants that receive direct sunlight. While often regarded as woodland species, none of these can breed within shaded areas of woodland or scrub. In the Burren, the eggs of the Brown Hairstreak are found on young growth on Common Blackthorn plants at the edge of a hedge,  scrub patch or woodland edge that is unshaded and south-facing.

Some butterflies enter a prolonged rest phase in their adult state. Four of our butterfly species undergo reproductive diapause in summer; these feed to build fat reserves before entering dormancy (quiescence) until the following spring when conditions favour breeding. The Brimstone, Small Tortoiseshell, Peacock, and Comma pass the winter in cool, dry conditions in scrub and woodland. It is likely that the Small Tortoiseshell, Peacock, and Comma require dense scrub for over-wintering. For this part of their life cycle, scrub and woodland habitats are vital.

Grassland/scrub mosaic in the Burren. These grassland/scrub transitional zones are excellent places for many butterflies and moths, but the development of scrub presents a challenge to farmers and conservationists attempting to protect orchid-rich grassland.

The removal of large areas of scrub over an extended area can produce severe survival challenges for rarer, scrub-dependent butterflies. Any remaining areas may be too small to support populations. Where suitable habitat patches are isolated, a species may modify its behaviour to secure its survival. There may be selection for sedentary individuals. Highly localised, sedentary populations are vulnerable to extinction arising from causes such as habitat destruction, bad weather, parasitoids, or perhaps inbreeding. Selection for more sedentary races of the Pearl-bordered Fritillary is suspected to have taken place in England in areas where the butterfly depended largely or fully on coppicing for its habitat (Thomas and Lewington 2014).

Climate change is expected to produce warmer, drier summers in Ireland. Many of the warmest years on record have occurred since 2000. According to the Copernicus European State of the Climate report in 2020, 11 of the 12 warmest years occurred since 2000. The projected decreases in rainfall for Ireland are largest for summer, with reductions ranging from 0% to 13% and from 3% to 20% for the medium-to-low and high emission scenarios, respectively. Projections indicate an increase of 1–1.6°C in mean annual temperatures, with the largest increases seen in the east of the country. Warming is enhanced for the extremes (i.e., hot or cold days), with the highest daytime temperatures projected to rise by 0.7–2.6°C in summer and the lowest night-time temperatures to rise by 1.1–3°C in winter (Nolan 2015).

Burren vegetation showing the effect of prolonged hot, dry weather in June 2018.

During recent hot, dry summers such as in 2018, desiccation of larval foodplants was observed in open, unshaded grasslands. During the hot weather, some species that usually frequent open areas during the day were observed sheltering in scrub. Interestingly, the Common Blue, which breeds on plants such as Common Bird’s-foot-trefoil growing in open grassland, was observed laying eggs on the foodplant growing in the shade. There may be several reasons for this untypical behaviour; the plants in open areas were in poor condition while in shaded areas they were not shriveled, the plants in shaded places were probably growing at the correct temperature during the heat of that summer, while during a more ‘typical’ summer, shaded plants will not support the larva. Finally, stress from extreme heat may be driving the adults to choose plants in less suitable micro-sites.

Breeding on plants shaded by scrub may be an adaptive behaviour by open grassland breeders under a warming climate. The increasing heat and dryness over the coming years may increase the importance of scrub for some Lepidoptera and other invertebrates.

The vast majority of Irish Lepidoptera are moths. There are about 1567 moth species recorded in Ireland with about 970 of these occurring in the Burren (D. Allen pers.com.). About 600 macro (larger) moths have been recorded. The following figures show the foodplants used by macro-moths; note that some species are polyphagous and use several of the following scrub species. 121 species have been recorded on willows, 96 species recorded on birches Betula spp., 61 species recorded on oaks Quercus spp., (the latter two tree groups are uncommon in the Burren scrublands), 26 species recorded on Aspen, 29 species recorded on Common Blackthorn, 36 species recorded on Common Hawthorn, 44 species recorded on Common Hazel, six species recorded on Purging Buckthorn and five on Alder Buckthorn (Waring et al 2004).

Scrub and limestone grassland on limestone pavement in the Burren National Park.

Hedges, scrub, and woodland are vital for our moth species, and not simply for larval foodplants. Moths use wooded habitats in similar ways to butterflies. One example is provided in a study by Coulthard et al. (2016) which showed that hedges are very important flight paths for moths. 68% of moths in the study were observed at 1m from the hedge, and of these 69% were moving parallel to the hedge. Hedges are believed to provide the sheltered corridors needed by flying insects in our generally open, farmed landscapes. This is likely to be especially relevant in open, exposed areas like the Burren.

The Silver-washed Fritillary breeds in scrub where abundant light reaches violets, the larval foodplant.
Silver-washed Fritillary larva prepares to pupate in a clearing in hazel scrub containing abundant violets.

There is an overwhelming body of evidence that supports the idea that more heterogeneous habitats can support more species diversity (Broeker 2018). Habitat heterogeneity, or small-scale changes in resource composition and structural complexity, provides more possible niche space for organisms to occupy and exploit (Tews et al, 2004). The range of habitats in the Burren, especially in the eastern areas, where complex mosaics of fens, cutover bogs, grasslands, heaths, limestone pavement, woods, and scrub exist support more butterfly species than anywhere else in Ireland. The scale of the habitats supports connectivity/dispersal, and genetic diversity and provides a range of niches in the face of the impacts of climate change, offering the prospect of adaptation when species are confronted with survival challenges. Maintenance of these conditions not only supports biodiversity generally, it preserves the magnificence of the region, touching those who experience the Burren with “the joy of elevated thoughts” (Lines Composed a Few Miles above Tintern Abbey, On Revisiting the Banks of the Wye during a Tour. July 13, 1798).

References

Broeker, H. (2018) Habitat Heterogeneity, Morphospecies Richness, and Niche Exploitation in the Human Skin Microbiome. Online at https://www.lakeforest.edu/news/habitat-heterogeneity-morphospecies-richness-and-niche-exploitation-in-the-human-skin-microbiome, accessed 11 September 2022

Coulthard, E., McCollin, D. & Littlemore, J. (2016) “The use of hedgerows as flight paths by moths in intensive farmland landscapes”, Journal of Insect Conservation, vol. 20, no. 2, pp. 345-350.

European Commission (2020) Warming trend shows 11 of the 12 warmest years occurred since 2000, according to the Copernicus European State of the Climate report. Online at: https://climate.copernicus.eu/warming-trend-shows-11-12-warmest-years-occurred-2000-according-copernicus-european-state-climate, accessed 10 September 2022

Fossitt, J.A. & Heritage Council Ireland (2000) A guide to habitats in Ireland. Heritage Council/Chomhairle Oidhreachta, Kilkenny.

Harding, J. (2021) The Irish Butterfly Book. Privately Published, Maynooth.

Nash, D., Boyd, T. & Hardiman, D. (2012) Ireland’s Butterflies: A Review. Dublin Naturalists’ Field Club, Dublin.

Nolan, P. 2015. EPA Report: Ensemble of Regional Climate Model Projections for Ireland. EPA climate change research report no. 159. EPA: Wexford. (show me this citation)

Tews, J., Brose, U., Grimm, V., Tielbörger, K., Wichmann, M. C., Schwa­ger, M., & Jeltsch, F. (2004) “Animal species diversity driven by habitat heterogeneity/diversity: the importance of keystone structures.” Journal of Biogeography, 31(1), 79-92.

Thomas, J. & Lewington, R. (2014) The Butterflies of Britain and Ireland. (Revised edition) British Wildlife Publishing, Dorset.

Waring, P., Townsend, M., & Lewington, R. (2004) Field Guide to the Moths of Great Britain and Ireland. British Wildlife Publishing, Hampshire.