| 107. Melitaea athalia  (Rottemburg,  1775) / Heath fritillary / Nymphalidae – Melitaeinae NL:  bosmelitea,  bosparelmoervlinder / D: Wachtelweizen-Scheckenfalter / F: melitée du mélampyre,  damier athalie      Photographs:  Jeroen Voogd ©.
 Small or  medium-sized, wing length 19 (16-21) mm. In the Benelux, this species is a  woodland inhabitant, at edges between heath and wood or in open woodland with  an undergrowth of a vegetation containing common cow-wheat (Melampyrum pratense). It occurs in  Luxemburg and Wallonia but has disappeared from Flanders (Maes & Van Dyck  1999: 312-313) and in the Netherlands it has disappeared from many places and  is now present only in a few localities on the Veluwe. Butterfly  is on the wing from end-May until mid-July and peaks mid-June. The species is  known from maritime and continental climates, amplitude 6 to 17. Required heat  sum 600°d and maximum tolerated 2000°d, corresponding climate windows 22 and 38  weeks. At present  it seems that the occurrence of the species is correlated with the presence of Melampyrum pratense but in fact the  larvae feed on various plants containing iridous glycosides (Van Genderen,  Schoonhoven & Fuchs 1996: 221-228).In England  this species occurs mainly in coppiced woods or copses and the main  conservation practise is to re-establish this historical exploitation. However,  on the Exmoor hills, this species inhabits open heathland, feeding on foxglove  (Digitalis  purpurea).
 In the  Netherlands some reintroduction trials were carried out in the 1980’s and  1990’s in the province Drenthe, the last one in 1995. However, none were  successful (Bos et al. 2006: 288).
 In the subarctic climate in the north of  Scandinavia there occurs the subspecies Melitaea  athalia norvegica,which must  have a quite different lifecycle because of the short season there.
 Ecological characteristics
 Behaviour over timeOverwintering: small larva in third or fourth instar in a  small group together under a wilted leaf in the litter layer.
 Reproduction: oviposition starts after 7-10 days when the  body contains 191 ± 60 eggs (n=7), estimated potential production 1.5 times as  much.
 Larval feeding periods: in summer about 32 days in period from  mid-July until late-August, in next spring 40 (28-51) days from end-March until  early-June.
 Generations: one.
 Spreading of risk: stage of overwintering, in third or fourth  instar, and the occurrence of a second diapause up to 8%.
 Life cycle: egg about 16 days; larva 47 or 99 weeks; pupa  15-26 days.
 Life  span of adult:  males short, 2 weeks, females rather longer, 3 weeks, 20.2 ± 5.4 days (n=5).
    Photographs:  Frits Bink ©.
 Behaviour in spaceFrom stay-at-home to migrant: stay-at-home, spatial requirement  modest.
 Finding a mate: male patrols or meets females when nectaring.
 Orientation in the landscape: gradient between wood and tall  grass or heath. Also an open wood with a grassy undergrowth.
 Oviposition: in batches of 75 (20-130) eggs on underside of  a mature leaf.
 DefenceThreats from other organisms: larva is armoured with stiff  bristles.
 Threats  from the environment: vulnerable  to heat and drought.
 Feeding habitsAdult: all kind of flowers, such as creeping thistle  (Cirsium arvense) and high up in the  canopy on a flowering common lime (Tilia  vulgaris).
 Larva: when young it lives gregariously in silken  nest, in spring more solitarily and wanders to look for a new host-plant.
  Larval foodplantsPlant species: Plantaginaceae,  Digitalis purpurea, Plantago lanceolata, Veronica chamaedrys, Orobanchaceae, Melampyrum pratense.
 JournalRearing  experiments:
 1. Based on  specimen from national park Hoge Veluwe, Netherlands:
 29 June 1980:  one female captured.
 8 July:  eggs laid.
 24 July:  eggs hatched.
 4 August:  larvae second instar 2-3 mm in length.
 17 August:  larvae unequal in size, most in third instar, 6.5 mm in length.
 [Accepted  plant species Veronica chamaedrys, Melampyrum arvense, M. pratense, Plantago  lanceolata. Refused: Veronica  officinalis, Succisa pratense.]
 Overwintered  outdoors.
 1 April 1981:  larvae had been active for 10 days, fed a little, basked socially on a leaf. Of  the original 80 larvae there were only eight survivors.
 17 April:  first prepupa, larvae hid at night and during cold days.
 9 May: last  larva pupated.
 13 May:  first adult appeared.
 2. Based on  specimen from the Loenermark, de Imbos, Netherlands:12 July 1990:  female captured, laid 184 eggs.
 Overwintered  outdoors.
 7 March 1991:  larvae basked but had not feed.
 10 March:  larvae started to eat Veronica spicata, V. chamaedrys, Digitalis purpurea.
 13 March:  there were eleven fourth instar and twelve third instar larvae.
 14 March: when  the ambient temperature had reached 29°C most larvae moved into shade and  appeared again when temperature dropped to 25°C.
 24 March:  larvae moulted, basked up to a temperature of 32°C, when it became warmer they moved  into the shade. They ate alternating Digitalis, Plantago, Veronica.
 28 March:  larvae mostly in moult, three in last instar.
 2 April: larvae  refused Veronica officinalis, Plantago maritima, but accepted Veronica agrestis, V. austriaca, Plantago  coronopus.
 7 April: first  pupa.
 5 May:  first adult appeared.
 14 May:  male and female appeared.
 19 May: another  female hatched.
 24 May: last pupa hatched.
 Table  107-1. Results of dissections 
 Table 107-2. Collection  and observation localities
 B, Ethe, 240 m, 49° 36’ 25”N – 5° 35’ 50”E; 11 July 1984. D, Meetschow, 53° 02’ 32”N – 11° 23’ 32”E; 22 June 1984.
 D, Arnstein, 50° 01’ 57”N – 9° 47’ 41”E; 22 July 1984.
 D, Steigerwald, 49° 39’N – 10° 23’E; 24 July 1984.
 EST, Kuresoo, 58° 28’ 00”N – 25° 11’ 59”E; 8 July 1999.
 F, Dommartin-la-Montagne, 49° 01’ 51”N – 5° 36’ 53”E;  24 June 2006.
 F, la Garde Freinet, 100 m, 43° 19’N – 6° 30’E; 29  April 1997.
 F, Lorraine, Rupt devant Saint-Mihiel, 280 m, 48° 53’  01”N – 5° 24’ 10”E; 28 June 2006.
 NL, Hoge  Veluwe, 52° 05’N – 5° 51’E; 29 June 1980, 11 July 1983, 30 July 1984.
 NL, Hoge Veluwe, 52° 04’ 47”N – 5° 49’ 47”E; 27 July 2000,  31 August 2000.
 NL, Hoge  Veluwe, 52° 03’ 41”N – 5° 50’ 46”E; 31 May 2002.
 NL,  Loenermark, de Imbos, 52° 06’ 09”N – 5° 58’ 36”E, 15 June 2001.
 S, Gotland, Buttle, 57° 25’ 09”N 18° 34’ 38”E; 25 June  2004, 6 July 2004, 16 July 2004.
 S, Gotland, Folhammar, 57° 20’ 49”N – 18° 44’ 12”E; 19  June 2004.
 S, Gotland, Ljugarn, 57° 22’ 03”N – 18° 41’ 31”E; 13  July 1982, 21 June 2004.
 S, Gotland, Grodde, 57° 52’ 01”N – 18° 49’ 02”E; 2  July 2004.
 S, Gotland, Hummelbosholm, 57° 11’ 48”N – 18° 32’  56”E; 8 July 2004.
 S, Gotland, Klinteklinten, 57 40’ 14”N – 18 46’ 26”E;  13 July 2004.
 S, Gotland, Lickershamn, 57° 50’ 17”N – 18° 34’ 54”E;  3 July 2004.
 S, Gotland, Lickershamn, 57° 51’ 05”N – 18° 37’ 46”E;  17 July 2004.
 S, Gotland, Östergarn, 57° 24’ 43”N – 18° 43’ 27“E; 15  July 2004.
 S, Gotland, Russväter, 57° 23 29”N – 18° 43’ 36”E; 16  July 2004.
 S, Gotland, Stygmyr, 57° 50’ 10”N – 18° 38’ 32”E; 18  July2004.
 S,  Öland, Ismantorp borg, 56° 44’ 44”N – 16° 38’ 33”E; 8 July 1982, 13 July 2004.
 Fig. 10-1. Melitaea athalia, phenogram  adapted from Fichefet et al. 2008: 235. 
 Fig. 107-2. Melitaea athalia, habitat characteristics. 
 Fig. 107-3. Melitaea athalia athalia, climate matrix, heat-sums 600 - 2000°d.Melitaea athalia norvegica, heat-sums 150-400°d (dark part).
 
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