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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                                                                   5 5 5 5
                                                                       Trichogramma sp. egg parasite); it certainly means that
                                                                       the larval defence tactics, as presented in this paragraph    7KH TXHVWLRQ
                                                                                                                                     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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