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Large-Heath-davus Whixall Moss 13-June-11 03C8780
Wingspan
35 - 40mm
Photo © IainLeach
Large Heath

Coenonympha tullia
Number: 59.004
B&F No.: 1628
Family:Nymphalidae (Swainson, 1827)
Subfamily:Satyrinae (Boisduval, 1833)
Tribe:Coenonymphini (Tutt, 1896)
Genus:Coenonympha (Hübner, 1819)
Subgenus: 
Species:tullia (Müller, 1764)
Subspecies:davus (Fabricius, 1777)
 polydama (Haworth, 1803)
 scotica (Staudinger & Rebel, 1901)
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  Introduction  

Found in the north of the British Isles, the Large Heath is unique in that it is more or less confined to boggy areas. The Large Heath lives on the British mainland in isolated colonies from central Wales in the south to Orkney in the north, and also in scattered colonies throughout Ireland. It is absent from Shetland. The best colonies can be very large in good years, where the number of adults emerging is measured in thousands. Large colonies used to exist in the mosses around Manchester and Liverpool, but these have long since disappeared.

The eye spots on the underside of this species vary considerably. Those in the north have almost no spots at all with adults looking like a large Small Heath, while those in the south have very distinctive spots. This has given rise to 3 named subspecies. Those with the least distinct spots are referred to as ssp. scotica, those with the most distinct spots as ssp. davus and those that are intermediate as ssp. polydama. This species forms a typical cline and, unsurprisingly, intermediates occur between the 3 named subspecies. For example, Ford (1945) writes: "It appears that in the island of Islay scotica predominates, but that the intermediate sub-species tullia [= polydama] is not uncommon and that even specimens closely approaching philoxenus [= davus] occur".

Ford (1945) also describes the situation in Ireland: "The sub-species philoxenus [= davus] does not occur, but scotica and tullia [= polydama], with their intermediates, fly together in the same location". Nash (2012) concurs with Ford (1945) that both scotica and polydama are found in Ireland. Riley (2007), however, suggests that only polydama is found in Ireland, although no explanation is given for this position.

Brakefield (1992) describes, in detail, the reason for this variation, which is believed to be natural selection based on predation by birds. The cooler climate in the north, along with fewer hours of daylight, results in less-active adults whose plain undersides make them difficult to find while at rest. Adults further south, on the other hand, are much more active and are more-likely to attract the attention of birds as a result. In this case, the distinct eye spots deflect the bird's attention away from the body.

Coenonympha tullia ssp. tullia

The species was first defined in Müller (1764) as shown here (type locality: Frederiksdal, Copenhagen, Denmark). The nominate subspecies has not been recorded in the British Isles.

Coenonympha tullia ssp. davus

This subspecies was first defined in Fabricius (1777) as shown here (type locality: Germany). This subspecies can be found in north-west England and central England near the border with Wales. This is the darkest and most colourful of the subspecies.

Coenonympha tullia ssp. davus (Fabricius, 1777)

Original (Latin)

alis integerrimis fuluis: anticis ocellis duobus, posticis sex coecis, subtus pupillatis.

Habitat Hamburgi Dr. Schulz, Kilonii Sehestedt.

Medius. Alae anticae supra fuluae ocellis duobus atris coecis tertioque minutissimo vix distincto; subtus fascia alba ocellis duobus pupilla alba. Posticae obscuriores ocellis quinque aut sex coecis; subtus griseae fascia interrupta alba ocellis sex atris pupilla alba posteriore didymo.

Translation

Wings entire, reddish yellow: the forewings with two eyespots, the hindwings with six, blind (above), pupilled beneath.

Lives in Hamburg (Dr Schulz), Sehestedt in the Kiel region.

Medium. Forewings yellowish red above with two blind, black eyespots and a tiny, barely discernible, third one; a white band beneath with two eyespots with white pupils. Hindwings dark with five or six blind eyespots; grey beneath with a broken white band, with six black, white-pupilled eyespots, the last one double.


Large Heath - Whixall Moss 2-July-2011
Male
Photo © ronniethepoo
Large Heath - male - Meathop Moss - 19-Jun-14-2
Male Underside
Photo © Pete Eeles
Large Heath female - Whixall Moss 10-July-2013
Female
Photo © MikeOxon
Large-Heath-davus Whixall Moss 19-June-11 03C2778
Female Underside
Photo © IainLeach

Coenonympha tullia ssp. polydama

This subspecies was first defined in Haworth (1803) as shown here (type locality: Yorkshire, England). This subspecies is found in parts of England, Wales, Scotland and Ireland. In Wales it is found in central and north-west areas. It is also found in the north of England in Cumberland, North Northumberland, South Northumberland, North-east Yorkshire and North Lincolnshire. It is found in southern Scotland from the border with England up to a line that runs between Renfrewshire in the west to South Aberdeenshire in the east, being replaced further north by the subspecies scotica. In Ireland this widespread in the north, but more isolated elsewhere.

This subspecies is considered to be intermediate between davus and scotica. It differs from the subspecies davus in being somewhat paler on both upperside and underside, and with fewer eyespots on the underside. Those eyespots that are present are smaller in size and often lacking the white pupil.

Coenonympha tullia ssp. polydama (Haworth, 1803)

Original (Latin)

P.D.F. (The Marsh Ringlet) Alis fulvis, anticis subtus ocellis duobus; posticis 6 albo cinctis quarum 3 dimidiatis.

Pap. Polydama Scop. Carn. 434?

HABITAT rarissime comitatu Eboracense. Semel capta et ad me missa amicissimo meo P.W. Watson. Imago mense Junio Paludosis.

EXPANSIO alarum 1 unc. 7 lin.

DESCRIPTIO. Imago. Alae antlcae griseo-fulvae ocellis duabus posticis caecis. Alae posticae fuscae sed ad latus interius late albicantes, puncto ocellari caeco parvo postico versus angulum ani. Subtus anticae fulvo-fuscae, basi nigricantes, apice cinereae, fascia postica albida abbreviata transversa; inter hanc et marginem posticum ocelli 2 remoti pupilla obsoleta alba, iride nigra albo cincta. Posticae basi fascia lata nigricante extus dentata, fasciola albida irregulari terminata; pone hanc cinereae; ocellis 6 parvis quarum 3 dimidiatis et fere obliteratis, omnibus circulo albo cinctis.

OBS. Simillima praecedenti magnitudine et statura; differt supra magis fulva, subtus magis cinerea; ocelli minores inaequales albo nec fulvo cincti.

Translation

P.D.F. (The Marsh Ringlet) Wings reddish yellow, the forewings with two spots beneath; the hindwings with 6, circled in white, of which three are reduced.

Pap. Polydama Scop. Carn 434?

Very rare in the county of Yorkshire. I was once sent one captured by my very good friend P.W. Watson. Adult in the month of June, in bogs.

WINGSPAN 1 inch. 7 lin.

DESCRIPTION. Adult. Forewings greyish-fulvous with two blind eyespots towards the outer [edge]. Hindwings dark but whitening broadly towards the inside, with a small, blind, eyelike rear spot towards the anal angle. Beneath, the forewings are dark reddish yellow, blackish towards the base, grey at the apex and with a short, white transverse band; between this and the outer margin are 2 distant eyespots with vestigial white pupils, the iris black, encircled in white. Hindwings with a broad, blackish region at the base, outwardly toothed, bounded by a white, irregular strip; beyond this [they are] grey; with 6 small eyespots of which 3 are reduced and almost lost, all of them encircled with white.

OBS. Similar to the previous one in size; differs above by having more reddish yellow and beneath by more grey; eyespots smaller and unequal, circled with white not reddish yellow.


Large Heath - Steng Moss, Harwood Forest, Northumberland 6-July-2014
Male
Photo © citybirding
Large-Heath-polydama- 5D34991 Crowle, Lincs 21 June 2013
Male Underside
Photo © IainLeach
Female
Large-Heath-polydama-Crowle-23-June-2010- 03C3190
Female Underside
Photo © IainLeach

Coenonympha tullia ssp. scotica

This subspecies was first defined in Staudinger & Rebel (1901) (type locality: Scotland). This subspecies is found in northern Scotland, north of a line between the Clyde Isles in the west and North Aberdeenshire in the east. It is found in most of the western isles and is also present in Orkney. As described above, Ford (1945) considers both polydama and scotica to occur in Ireland, as does Nash (2012). However, Riley (2007) suggests that only polydama is found in Ireland and that, therefore, scotica is absent. Whatever the correct status, if scotica does occur in Ireland, then it follows the distribution of polydama and is widespread in the north, but more isolated elsewhere.

In comparison with the subspecies polydama this subspecies is paler with minute, often absent, underside eyespots. Apart from its larger size, it appears very similar to its close cousin, the Small Heath.

Coenonympha tullia ssp. scotica (Staudinger & Rebel, 1901)

Original (Latin)

al. supra [al. post. latius] cinereo-marginatis, subt. obscurior ocellis subnull.

Translation

Wings grey margined above [the hindwings more broadly], the eyespots on the underside being more obscure, virtually absent.


Male
Large Heath ssp. scotica 11th July 2014  Nr Loch Garten Scotland  imago 1
Male Underside
Photo © Reverdin
Female
Large Heath (ssp. scotica) - Braemore, Wester Ross  5-July-2012
Female Underside
Photo © nomad

  Phenology  

There is one generation each year, with adults emerging at the end of June at most sites, peaking in early July. In northern Scotland and at high altitude adults emerge a little later, at the beginning of July.

Coenonympha tullia ssp. davus


Coenonympha tullia ssp. polydama


Coenonympha tullia ssp. scotica


The chart(s) above have been correlated with the phenology plot below, taken from the UK Butterfly Monitoring Scheme. The blue line gives average counts over the full data set from 1976 to date, and the red line gives the average for the last year.


  Habitat  

This butterfly is found in flat wetland areas such as bogs, waterlogged peat mosses and damp moorland where the foodplant, normally Hare's-tail Cottongrass, and appropriate nectar sources are also found. Sites are often clothed in Heather.

  Larval Foodplants  

The primary larval foodplant is Hare's-tail Cottongrass (Eriophorum vaginatum). Common Cottongrass (Eriophorum angustifolium) and Jointed Rush (Juncus articulatus) are also used.

  Nectar Sources  

Adults feed primarily on Cross-leaved Heath (Erica tetralix), Hawkweeds (Hieracium/Hypochoeris), Heather (Calluna vulgaris / Erica spp.), Tormentil (Potentilla erecta) and White Clover (Trifolium repens).

  Imago  

Despite its slow and lumbering flight, the butterfly can be difficult to follow due to the boggy ground underfoot. Furthermore, when disturbed, the butterfly will launch itself into the air, often flying some distance before landing again. All in all, this is not always the easiest species to see, let alone photograph!

The adults remain somewhat active even in dull weather, but will remain tucked away in vegetation in strong winds and cold weather. Males are more-often seen than females, which tend to stay hidden away in grass tussocks unless nectaring or egg laying. Adults always rest with their wings closed and regulate their temperature by orientating their wings at an appropriate angle to the sun. Both sexes take nectar, Cross-leaved Heath being a particular favourite that often grows alongside the foodplant.

Coenonympha tullia ssp. davus


Large Heath - Meathop Moss, Cumbria 11-July-07
Photo © Vince Massimo
11-Jul-2007
Large Heath - imago - Meathop Moss - 29-Jun-06 (0427)
Photo © Pete Eeles
29-Jun-2006
Large Heath - imago - Meathop Moss - 29-Jun-06 (0429)
Photo © Pete Eeles
29-Jun-2006
Large Heath - imago - Meathop Moss - 29-Jun-06 (0433)
Photo © Pete Eeles
29-Jun-2006
Large Heath [Nick Sampford]
Photo © Nick Sampford
Large Heath - Whixall Moss 2-July 2011
Photo © ronniethepoo
Large Heath - Whixall Moss 2-July-2011
Photo © ronniethepoo
Large-Heath-davus Whixall Moss 19-June-11 03C2591
Photo © IainLeach
Large-Heath-davus Whixall Moss 19-June-11 03C2658
Photo © IainLeach
Large-Heath-davus Whixall Moss 19-June-11 03C2778
Photo © IainLeach
Large-Heath-davus Whixall Moss 13-June-11 03C8731
Photo © IainLeach
Large-Heath-davus Whixall Moss 13-June-11 03C8780
Photo © IainLeach
Large-Heath-davus Whixall Moss 13 June 2011  03C8430
Photo © IainLeach
Large-Heath-davus Whixall Moss 13 June 2011  03C8490
Photo © IainLeach
Large-Heath-davus Whixall Moss 13 June 2011  03C8618
Photo © IainLeach
Large-Heath-davus Whixall Moss 19 June 2011  03C2699
Photo © IainLeach
Large Heath ssp. davus - Whixall Moss Shropshire 04.07.2012
Photo © nfreem
04-Jul-2012
Large Heath ssp. davus - Whixall Moss Shropshire 04.07.2012
Photo © nfreem
04-Jul-2012
Large Heath ssp. davus - Whixall Moss Shropshire 04.07.2012
Photo © nfreem
04-Jul-2012
Large Heath female - Whixall Moss, Shropshire 19-June-2010
Photo © Neil Hulme
19-Jun-2010
Large Heath - Whixall Moss,Shropshire 18-June-2010
Photo © Neil Hulme
18-Jun-2010
Large-Heath-davus- 5D30741 Whixall Moss July 2013
Photo © IainLeach
Large-Heath-davus- 5D30931 Whixall Moss July 2013
Photo © IainLeach
Large-Heath-davus- 5D31439 Whixall Moss July 2013
Photo © IainLeach
Large-Heath-davus- 5D31586 Whixall Moss July 2013
Photo © IainLeach
Large Heath female - Whixall Moss 10-July-2013
Photo © MikeOxon
Large Heath - Meathop Moss Cumbria 14.06.2014
Photo © nfreem
14-Jun-2014
Large Heath - male - Meathop Moss - 19-Jun-14-2
Photo © Pete Eeles
19-Jun-2014
Large Heath - male - Meathop Moss - 19-Jun-14
Photo © Pete Eeles
19-Jun-2014

Coenonympha tullia ssp. polydama


Large Heath - imago - Kirkconnell Flow, Dumfries-shire - Unknown date [Adrian Riley]
Photo © Adrian Riley
Large Heath - imago - Peatlands Country Park, Northern Ireland - 14-Jun-06 [Graham Smith]
Photo © Graham Smith
Large Heath - Mullenakill Bog, Peatlands Park, Co Armagh, Northern Ireland 25-June-2011
Photo © Dave McCormick
25-Jun-2011
Large Heath, Dogden Moss, Berwickshire, Scotland. July 1st 2011.
Photo © IAC
01-Jul-2011
Large-Heath-polydama-Crowle-23-June-2010- 03C2213
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C2416
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C2951
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C3037
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C3127
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C3190
Photo © IainLeach
Large-Heath-polydama-Crowle-23-June-2010- 03C3651
Photo © IainLeach
Large Heath ssp polydama Rothbury, Northumberland
Photo © Graham Beckwith
03-Jul-2011
Large Heath ssp polydama Upperwings Rothbury, Northumberland 28th June 2011
Photo © Graham Beckwith
28-Jun-2011
Large-Heath-polydama- 5D34640 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D34751 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D35360 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D35525 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D35605 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D35687 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D34991 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large-Heath-polydama- 5D35212 Crowle, Lincs 21 June 2013
Photo © IainLeach
Large Heath - female - Thatcham - 13-Jul-13 (1) [REARED]
Photo © Pete Eeles
13-Jul-2013
Large Heath - female - Thatcham - 13-Jul-13 (2) [REARED]
Photo © Pete Eeles
13-Jul-2013
Large Heath - female - Thatcham - 13-Jul-13 (3) [REARED]
Photo © Pete Eeles
13-Jul-2013
Large Heath - female - Thatcham - 16-Jul-13 (1) [REARED]
Photo © Pete Eeles
16-Jul-2013
C. tullia spp. polydama North Yorkshire June 2014
Photo © Reverdin
21-Jun-2014
Large Heath - Steng Moss, Harwood Forest, Northumberland 6-July-2014
Photo © citybirding
06-Jul-2014

Coenonympha tullia ssp. scotica


Large Heath - imago - Creag Meagaidh, Inverness - Unknown date [Adrian Riley]
Photo © Adrian Riley
Large Heath (ssp. scotica) - Braemore, Wester Ross  5-July-2012
Photo © nomad
above Braemore, Wester Ross
05-Jul-2012
Large Heath (ssp. scotica) - Fannich Mountains, Wester Ross 5-July-2012
Photo © nomad
Moorland below the Fannich Mountains, Wester Ross
05-Jul-2012
Large Heath (ssp. scotica) - Fannich Mountains, Wester Ross 5-July-2012
Photo © nomad
Moorland below the Fannich Mountains, Wester Ross
05-Jul-2012
Large Heath ssp. scotica 11th July 2014  Nr Loch Garten Scotland  imago 1
Photo © Reverdin
Large Heath ssp. scotica, seen Nr Loch Garten, Scotland 11/07/2014
11-Jul-2014
Large Heath ssp. scotica 11th July 2014  Nr Loch Garten Scotland  imago 2
Photo © Reverdin
Large Heath ssp. scotica, seen Nr Loch Garten, Scotland 11/07/2014
11-Jul-2014

  Aberrations  

Description to be completed.

Click here to see a full list of aberrations for this species.

Unclassified Aberrations


Large Heath - Crowle Moors - 23.06.2010
Photo © PhiliB
23-Jun-2010
Large Heath ssp polydama Dark Specimen Rothbury, Northumberland 3rd July 2011
Photo © Graham Beckwith
An unusually dark specimen with entirely orange tips to antennae.
03-Jul-2011

ab. lanceolata (Arkle.Entom.1913.46.p.93.)

On the underside the spots of both wings lanceolated.


Large Heath, Davus ab.lanceolata. 3/7/2011 Meathop Moss.
Photo © badgerbob
03-Jul-2011

  Ovum  

The spherical eggs are laid singly on the foodplant, often on dead leaves at the base of the plant, and are pale yellow when first laid, although brown blotches develop after several days, the egg growing even darker as the larva develops within. This stage lasts around 2 weeks.


Large Heath - ovum - Unknown Location - Unknown Date (2) [REARED] [Reg Fry]
Photo © Reg Fry
Large Heath - ovum - Meathop Moss - 19-Jun-14
Photo © Pete Eeles
19-Jun-2014

  Larva  

The larva feeds on the tender leaf tips of the foodplant and remains hidden away deep within the tussock when not feeding. The larva hibernates while in the 3rd instar and it has been known for larvae to pass two winters before pupating, particularly in northern colonies. The larva can also survive long periods under water and even being frozen - both distinct possibilities in their boggy habitat. There are 4 moults in total.


Large Heath - larva - Thatcham - 17-Apr-05 (3) [REARED]
Photo © Pete Eeles
15-Apr-2005
Large Heath - larva - Thatcham - 17-Apr-05 (4) [REARED]
Photo © Pete Eeles
15-Apr-2005
Large Heath - larva - Thatcham - 28-May-05 (2) [REARED]
Photo © Pete Eeles
28-May-2005
Large Heath - larva - Thatcham - 28-May-05 [REARED]
Photo © Pete Eeles
28-May-2005
Large Heath - larva - Thatcham - 07-Sep-12 (1) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 22-Aug-12 (55) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 10-May-13 (1) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 10-May-13 (6) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 18-May-13 (1) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 18-May-13 (6) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 25-May-13 (1) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 25-May-13 (2) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 25-May-13 (5) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 25-May-13 (6) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 27-Apr-13 (2) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 27-Apr-13 (13) [REARED]
Photo © Pete Eeles
Large Heath - larva - Thatcham - 16-Jun-13 (4) [REARED]
Photo © Pete Eeles
16-Jun-2013

  Pupa  

The pupa hangs head down, attached by the cremaster to the foodplant or other vegetation. This stage lasts around 3 weeks.


Large Heath - pupa - Lincs - Jun-00 [REARED] [Graham Smith]
Photo © Graham Smith
Large Heath - pupa - Thatcham - 16-Jun-13 (4) [REARED]
Photo © Pete Eeles
16-Jun-2013
Large Heath - pupa - Thatcham - 19-Jun-13 (1) [REARED]
Photo © Pete Eeles
19-Jun-2013
Large Heath - pupa - Thatcham - 29-Jun-13 (1) [REARED]
Photo © Pete Eeles
29-Jun-2013

  Similar Species  

Small Heath

Description to be completed.

  Videos  

No videos are currently available for this species.

  Sites  

Click here to see the distribution of this species overlaid with specific site information. Alternatively, select one of the sites listed below.

Sites
Alemoor West Loch and Meadow SSSI, Allt Mhuic Nature Reserve, Ardderry Lough, Ballynahone Bog, Beinn Eighe, Bellart How Moss, Bellcrag Flow, Bowness Common, Clara Bog, Cors Goch, Creag Meagaidh, Creighton's Wood, Crowle Moors, Drumburgh Moss, Falstone Moss, Fen Bog, Feoch Meadows, Ffridd y Fawnog, Ford Moss, Forsinard, Glasdrum Wood, Glen Loy, Glen Nevis, Harbottle Crags, Hartside, Inversnaid, Kirkconnell Flow NNR, Knowetop Lochs, Lake Vyrnwy, Letterfrack, Loch Garten, Loch Mudle, Lochuisge, May Beck, Meathop Moss, Meldon Hills, Mouds Bog, Muckle Moss, Peatlands Country Park, Pont-ar-Gonwy, Rabley Hill, Roudsea Wood NNR, Threepwood Moss SSSI, Wester Moss, Whixall Moss, Winmarleigh Moss

  Conservation Status  

There has been a moderate decline of this species over the long term and it is therefore a priority species for conservation efforts. The primary cause of this decline has been the drainage of its habitat for industry or agriculture, rendering such sites unsuitable for this species.

UK BAP StatusDistribution TrendPopulation Trend
Priority Species
Click here to access the Biodiversity Action Plan (BAP) for this species.
DecreaseLarge Increase

From The State of Butterflies in Britain and Ireland and the UK Biodiversity Action Plan (UK BAP) (2007 review).


  Links  

The following links provide additional information on this butterfly.

  References  

The species description provided here references the following publications:

ReferenceDetails
Brakefield (1992) Brakefield, P.M. and Shreeve, T.G.: Case Studies in Evolution. The Ecology of Butterflies in Britain. 1992.
Fabricius (1777) Fabricius, J.C.: Genera Insectorum. 1777.
Ford (1945) Ford, E.B.: Butterflies. Edn.1. 1945.
Haworth (1803) Haworth, A.H.: Lepidoptera Britannica. 1803.
Müller (1764) Müller, O.F.: Fauna insectorum Fridrichsdalina: sive Methodica descriptio insectorum agri fridrichsdalensis . 1764.
Nash (2012) Nash, D., Boyd, T. and Hardiman, D.: Ireland's Butterflies: A Review. 2012.
Riley (2007) Riley, A.M.: British and Irish Butterflies: The Complete Identification, Field and Site Guide to the Species, Subspecies and Forms. 2007.
Staudinger & Rebel (1901) Staudinger, O. and Rebel, H.: Catalog Der Lepidopteren Des Palaearctischen Faunengebietes. 1901.

  Copyright © Peter Eeles 2002-2014
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