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In the “complete time-calibrated multi-gen phylogeny of localities; we can compare it, speculate which one is built clear case of epigenetics (presented in the 38th ILS annual meeting on 16 3 3 3 3 3 3 3 3
the European butterflies” (Wiemers et al., 2020). faster, simpler, more sophisticated (engineering-wise), July 2021 in Kibbutz Na’an, Israel & pers. comm. to DB).
“L. reducta is quite far from L. camilla. They are not but we still did not measure the final results – the % of * DB started to see annual single atalanta overwintering visitors to his
sister taxa!...(and)…have an approximate divergence of their success to improve their offspring’s survivability. garden in Beit Arye in October 2001 and after the winter from 17.4.2002
about 8 or 9 million years. See the detail image of the While looking for explanation for the different defense every day nectaring on violet flowers of Lantana montevidensis until 2 May
tree I attach (the light grey band represents 20 MY)” – strategies that may relate to different “seniority” in 2002 its last day before taking off northwards. The same ritual happened
see Fig A - Roger Vila (pers. comm. to DB). In addition, evolutionary history their phylogeny as presented above every year and quite accurately on very similar dates, never more than one
Niklas Wahlberg a co-author of Wiemers et al., (2020) revealed a parallel evolution and hence in different life observed specimen.
wrote to DB: “... One could say that reducta and camilla zones and under different evolutionary threats (see Additional data: their LHP in my garden are only a few stands of introduced © Michel Baudraz
are closely related, but that they have had some period below). Parietaria judaica; larvae were feeding mostly on the southern warmer side
of time of independent evolution. (No one can)…say one of the house. On warm days of mid-winter in December and January the 4 4 4 4 4 4 4 4
species is older than the other, they have a common 3UHGDWRUV young L1s that still did not start consuming Red Admiral appeared to nectar on the same large Lantana bush (illustr.
ancestor…(from which)…they diverged…at the same leaves, hence did not get their frass camouflage 7). in springs of 2006 and 2007 they appeared on 24.2.2006 and
time, i.e. they are of equal age in comparison to each defence may pay heavy toll of up to 75% prey to small 9.2.2007 respectively, were observed in the garden until 3.3.2006 and
other” (see also Dhungel & Wahlberg, 2018). spiders and not often also to assassin bugs (Rhinocoris 7.3.2007; during this period they laid eggs and fresh adults were observed
spp., Reduviidae)(DB, reducta pers. obs. in Israel). In on 28.4.2006 and 23.4.2007. A year later the breeding cycle started on
Poland Adam Warecki found that “the main threat to 1.3.2008 and the fresh adult appeared on 1.5.2008. I never observed
wintering L. camilla larvae (inside its hibernaculums) any adult in my garden between the 2nd week of May to the last week of
are spiders (pers. comm. to DB). During many years of September.
breeding southern white admirals in Israel DB never © Evyatar Feingold
20 MY found any parasitoid in its larvae (except one record of 7KH P\VWHU\ DQG LWV VROXWLRQ
Trichogramma sp. egg parasite); it certainly means that
the larval defence tactics, as presented in this paragraph 7KH TXHVWLRQ 5 5 5 5
Every year in its overwintering sites all over Israel different adults appear
are very efficient, is it the reason that the female lay only every autumn. So, how comes that these new Red Admirals knew precisely
a few tens of eggs sufficient to keep a steady population? the exact location of its predecessors years earlier? How the exact “GPS”
(Lafranchis et al., 2015: 413). In the UK L. camilla location of the permanent biotope with its promised hostplants is transferred
larvae are attacked by the parasitoid Cotesia sibyllarum from one Red Admiral to another one? It seemed to be another case of
(Wilkinson) (Braconidae: Microgastrinae) (Eeles, pers. epigenetics; “science fiction?” of data transfer genetically from parents to
comm.) but its toll is yet unknown. their offspring ! because no other way of known know-how transfer exist in
butterflies – unlike bees or other social insects.
5HFRUGHG KRVWSODQWV Caprifoliaceae - Lonicera etrusca, 6XJJHVWHG DQVZHU
L. arborea. Eisenstein (2003: 181) & Benyamini (1999e) Let’s look again at the facts:
in Israel and Lafranchis (2019 & 2015: 412-415) in N Checking carefully facebook observations by photographers of the ILS,
Peloponessos, C Greece, S France and N Spain recorded DB found that the autumn arriving southbound Red Admirals were a mix of
egg laying also on the common cultivated L. japonica. worn and fresh specimens
)LJ $ - Modified F1 (cut, enlarged & marked), time-calibrated tree of These specimens were photographed in warm sunny winter days in © Yitzhak Meyuhas
Ļ
European butterflies (Lepidoptera: Papilionoidae) with time scale and December to February when they came out of their shelters to feed on
taxonomic assignment to subfamilies and families from Wiemers et al., 'LVWULEXWLRQ 6 6 6 6 6 6
2020: 110 – prepared by Roger Vila (CSIC-UPF), Barcelona, Spain . available nectar sources.
Larvae that were observed in October - November are winter 1st brood;
TL: Hankynda (Nagorno - Karabakh, Armenia). The their adults will proceed overwintering until spring, to produce 2nd brood or
/DUYDO GHIHQFH VWUDWHJLHV RI L. camilla 8. DQG species’ distribution ranges from S and C Europe to
Limenitis reducta ,VUDHO Turkey, Caucasia, Cyprus, Syria, Lebanon, Israel, rare in fly northwards before winter (no evidence).
“The larva of the White Admiral butterfly, Limenitis N Jordan (Dibbin, Jerash & Rasum), (absent from Sinai), Larvae that were observed in January-February are 2nd or 1st brood,
camilla ..is…– a master builder”. Fox (2005) presented Iran and Iraq. Reaching its southern limit of distribution their adults will leave to the northbound from March until May.
L1-L3 two complementary defence strategies; 1) “pier” in Netiv HaLamed-Heh (300 m), 24 km SW Jerusalem, Fresh Red Admirals started to appear in mid-winter gradually replacing
extension of Lonicera’s leaf midrib - built by bonding 31°41.2’N, Israel. Two subspecies have been described their “parents” until April-May when only fresh specimens were observed.
and silking frass-pellets to serve L1-L3 as a safe refuge, from the Levant: Limenitis reducta herculeana Stichel, This new generation of fresh, full of energy (lipids) specimens “took off”
resting and moulting place (illustr. 1). 2) an aerial 1908; TL: Dalmatia, S. France – S. Turkey, Cyprus. L. to their northbound summer localities and from June practically no Red
latrine built of cut leaves attached by silk threads and r. schiffermuelleri Higgins, 1933; TL: not given, Syria, Admirals remain in S Levant. © Gadi Ish-Am
suspended from the midrib by a silk strand - it serves Lebanon, Israel, Jordan. DB concluded that a possible solution to this mystery is that the same 7 7 7 7
the larvae to hide their frass and presence on the leaf, Israeli-born specimens made the whole round trip. It means that they had
and also use these stored pellets to bond it on its body to live at least five months from May to October but practically up to double
for camouflage pretending to be a piece of dirt (illustr. of this figure – not unknown in butterflies (e.g. the overwintering long-lived
12). Same needs for protection exist also in L1-L3 of L. Gonepteryx spp., the Monarchs, etc.). In their northbound to S & C Europe
reducta but the solutions are not identical; 1) instead they will mate and produce local brood/broods that will migrate (in part) in
of the complicated and possibly longer time required to the autumn back to their overwintering southbound localities. These may
build the “pier” reducta’s L1 “produce” it by feeding on also be Levant-born specimens that return home; It is still amazing how every
both sides of the midrib exposing 3-10 mm it needs. 2) single specimen can navigate hundreds or thousands of kilometers and find
no sophisticated latrine is built; L1 uses it fresh frass to (supposedly) exactly the place where it was “born”. But this capacity is known
quickly cover its body with these pellets achieving its first in the return flights of the Monarch Butterfly from their overwintering sites
priority for safety – the “frass defence” – illustr. 2. Other in Mexico northwards to North America. We still lack a lot of knowledge to
pellets remain on the leaf adding more similar pieces of fully understand this puzzle because at least some of the return specimens
dirt (risk sharing?), or hang them with silk creating spider- grow in northern habitats and each one had to find it parental birthplace
like web illustr. 13. The evolution of the first instar larvae by its epigenetics. A mirror image of northern populations that migrate
defence strategies as described are the output of their southwards for overwintering may suggest that while in S Europe (e.g. Greece
local stress for survival under different treats in different © Leah Benyamini and Spain) there are permanent populations the Levantine populations are © Dubi Benyamini

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2Butterflies of the Levant danaidae satyridae.indd 146 12/30/2021 4:27:08 PM 2Butterflies of the Levant danaidae satyridae.indd 159 12/30/2021 4:27:43 PM