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Melanargia titea  (Klug, 1832)         Levantine Marbled White



                    The most common  Melanargia in the Levant, where    micropyle. L1 hatches after 12-14 days, nibbles around
                    hundreds may be seen at the height of the short (two to   the egg top and pushes upwards to emerge. L1 is hairy,
                    three week) flight season. With no geographical barrier   2.5 mm long, light brown with five, thin longitudinal
                    and with much morphological variability,  titea  may be   brown lines, and with body tapering to the anal end. The
 #
                    present as lighter or darker specimens. The following,   eggshell may be eaten, after which the newly emerged
                    sometimes overlapping, ssp./forms have been described   larva enters a six-eight months summer aestivation,
                    from the Levant:                                    commencing feeding towards the end of December or
                    D  the nominate ssp.  titea from Beirut, Lebanon, flies   later at higher elevations. Larvae feed at night and hide
                    in the Levant species’ centre of distribution. This is the   at the base of the hostplant during the day, moving
 21603-BRACHA-PARPAR - 21603-BRACHA-PARPAR | 5 - B | 21-12-30 | 11:34:47 | SR:-- | Magenta
 21603-BRACHA-PARPAR - 21603-BRACHA-PARPAR | 5 - B | 21-12-30 | 11:34:47 | SR:-- | Yellow
                    darkest ssp./form, with a broad submarginal area of   slowly to avoid detection. Larvae grow to 28-35 mm,                        © Feza  Can          © Dubi Benyamini                                    © Dubi Benyamini
 #21603-BRACHA-PARPAR - 21603-BRACHA-PARPAR | 5 - B | 21-12-30 | 11:34:47 | SR:-- | Black
 21603-BRACHA-PARPAR - 21603-BRACHA-PARPAR | 5 - B | 21-12-30 | 11:34:47 | SR:-- | Cyan
                    the male uph where the ocelli are within the black area.   and are decorated with longitudinal, light, pastel green-
                    The cell in the basal area of the upf is white or slightly   brown-reddish-pinkish stripes. When alerted, larvae
                    suffused with black. The female is dark-brown and the   elevate from their prolegs and lower the head. Larvae
                    underside has a yellow-orange tint (Bozano, 2002: 12);   pupate, ventral side up, beneath litter on the ground,
                    E  ssp./form titania which is lighter, with ocelli open to   with no silken web. Another  Melanargia sp., i.e.  M.
                    the white uph and has the widest distribution in Israel, N   galathea, was reported to pupate ‘at the surface of the
                    Jordan, S Lebanon, Syria and S Hatay;               ground, under soil or moss’ (Thomas & Lewington, 2014:
                    F  ssp./form standfussi is the lightest of the Levant titea,   253); and head up at bottom of grasses (Lafranchis et
                    distributed from N Hatay north-eastwards, out of the   al., (2015: 683); Adam Warecki, pers. comm.). The light-
                    Levant to S Malatya and C Adiyaman districts of Turkey.   brown pupa, has a thin, dense dark net painting over its
                    However, its  ethology is identical everywhere: M. titea is   wing cases, and is 15 mm in length. Adults hatch after
                    a weak flyer, fluttering close to the ground, usually near   2-3 weeks.                                                                        © Dubi Benyamini          © Dubi Benyamini                 © Dubi Benyamini
                    its hostplants (Poaceae spp.) and adults are attracted
                    to flowers of Dipsaceae,  Centaurea spp. and other   $GXOW  FKHPLFDO  GHIHQFH  Melanargia  spp. are typified
                    composites. Specimens from high elevations sometimes   by a slow, fluttering and gliding flight, their coloration
                    have a yellow tint to the extensive white ground colour.   and patterning not typically aposematic, but such a
                    M.  titea is the most common of the mainland Levant   warning signalling might be conveyed by the fluttering
                    satyrines. Night-roosting was reported by Larsen in   black and white wings, allowing nectaring (e.g. on
                    Lebanon (1973b). They congregate to sip nectar together   Centaurea and Dipsaceae spp., the favoured nectar
                    (group nectaring).                                  sources in the Levant), seemingly without risk of
                                                                        predation. The ‘aposematic’ appearance derives from
                                                                        flavonoids, which are responsible for the yellowish and
                                                                        white colour of the wings, but as flavonoids are largely
                    %LRORJ\                                             non-toxic, the hostplants are, as stated by Dennis &
                                                                        Shreeve (2004) ‘not leading candidates for chemical
                    )OLJKW  SHULRG   early March around the Sea of Galilee   defence’. The nature of the suspected chemical
                    (-200 m, bsl), near Beit She’an upper Jordan Valley   defence (that had remained unknown for nearly two
                    (Graves, 1925a) and the Samarian Desert; April on the   and a half centuries since the initial description of
                    coast to the third week of June at 1200 m in the C & S   the type species,  Papilio galathea Linnaeus, 1758),                                              © Dubi Benyamini                                 © Ali Atahan
                    Mediterranean zone (Benyamini, 2002: 136); May-June   was revealed by Nash (in Rothschild, 2001) as Loline,
                    from 300 to 1200 m in Hatay (Atahan et al., 2018: 72);   a toxic pyrrolizidine alkaloid (PA). The toxin was found
                    mid-May at 1000 m to the third week of July in S Lebanon   to be present (at least in British  M. galathea reared
                    (Gebel Barouk, 1960 m), Mt Lebanon Cedar range and   on Agropyron sp., Poaceae) in both adults and pupae,
                    Anti-Lebanon Mt Hermon (both 2000 m) (Larsen, 1974:   and was shown to have been sequestered from grasses
                    130-131 & DB pers. obs.); 1545-2400 m in Al-Lazzab   infected by an endophytic, symbiotic fungus of the
                    reserve Syrian Anti-Lebanon (Zarikian & Ghrejyan, 2018).     genus  Acremonium (see also Porter, 1994). Whether
                                                                        grasses in the Levant are similarly infected, is not
                    /LIH KLVWRU\  univoltine. Attempted mating was observed   known, therefore we cannot extrapolate that  M.  titea
                    in early May 2014, near Adar Mt 10 km NW Jerusalem   uses PAs as a primary defence (Benyamini, 2003e).
                    (850 m), where a male was seen in pursuit of an     Certainly, the aposematic appearance did not prevent
                    unreceptive female. Upon landing, several rejection   the devouring by a chameleon of an entire M. titea, seen
                    postures were used by the female to repel the male:   nectaring on  Centaurea crocodylium in the garden of
                    initially wings were held open and vibrated, but when the   DB, while a second chameleon appeared ready to follow
                    male tried to land, the female elevated her abdomen and   suit (Benyamini, 2012c). Chemical defence is much
                    closed her wings, before turning head to head towards   better known in pierine butterflies (e.g. Pieris rapae, P.
                    the male. When mated, the female carries the male in   brassicae that sequester toxic mustard oils from their
                    cop. (DB field obs. & breeding notes). Females oviposit,   Brassicaceae hostplants), than in  Melanargia spp.
                    mostly without settling, while sipping nectar or fluttering   that, behaviourally and morphologically, more closely
                    over the hostplants; in captivity, females oviposited   resemble pierines than the satyrines.
                    while nectaring on Centaurea crocodylium flowers. Eggs
                    are white, spherical 1-1.25 mm in diam. (with tiny warts   RHFRUGHG  KRVWSODQWV  Poaceae (Gramineae) –  Avena
                    on 60-70% of the lower half) and slightly conical, with 15   sterilis,  Brachypodium distachyon (Feingold, 2005b),
                    vertical ribs upwards to the nearly flat top bearing the   Bromus spp., Piptatherum blancheanum & P. miliaceum.                                              © Ali Atahan                                   © Ali Atahan

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