Biology Reference
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6. DIAPAUSE CAN MODIFY LARVAL/PUPAL GROWTH
Modulations of larval/pupal developmental rate are also programmed
to adjust the growth on diapause trajectory. As an example, the swallowtail
butterfly Sericinus montelus triggers a pupal diapause linked to the modifica-
tions of larval/pupal development. In fact, larvae slow down development
for more than 20 days upon perception of short days (“diapause-inducing”),
while they accelerate growth ( < 19 days) under long days (direct develop-
ment). Pupal growth is also modified: diapausing pupae are fatter and heavier
than developing ones ( Wang, Yang, Zhou, Zhao, & Lei, 2007 ). Similarly,
the geometrid moths Cabera exanthemata and Chiasmia clathrata trigger pupal
diapause after a prolonged larval life in which the larval growth rates is lower
than those of larvae set for direct development ( Kivel¨,V¨lim¨ki, &
M¨enp¨¨, 2012; V¨lim¨ki et al., 2013 ).
The nymphalid Pararge aegeria diapauses either as a third (penultimate)
instar larva or as a pupa, depending by photoperiod ( Nylin, Wickman, &
Wiklund, 1989; Wiklund & Friberg, 2011 ). Larvae exposed to long days
( > 17 h of light) develop directly without diapause, those reared under inter-
mediate days (14-16 h) induce pupal diapause and those perceiving short
days ( < 13 h) trigger larval diapause ( Nylin et al., 1989 ). Such trajectories
reflect changes in larval/pupal growth rate since “diapausing” larvae develop
slower than developing ones and diapausing larvae develop into heavy
pupae, diapausing pupae have intermediate weight and “nondiapausing” lar-
vae develop into small pupae ( Gotthard & Berger, 2010; Nylin et al., 1989 ).
In this species, diapause trajectory is also linked to the appearance of seasonal
butterfly morphs (eyespots on the dorsal side of the wings) ( Gotthard &
Berger, 2010; Van Dyc & Wiklund, 2002; Wiklund & Friberg, 2011 ).
In contrast to the slowing of development in some species, diapause is
linked to a fast larval growth when larvae have to quickly reach the dormant
stage when developing in proximity of an imminent winter (i.e., late in the
season of “rapidly changing” environments) ( Gotthard, 2008 ). Such a phe-
nomenon is also reported for two nymphalids, Lasiommata maera and Lopinga
achine , whose larvae hatch in late summer and diapause as half-grown larvae
in autumn. Such larvae accelerate their growth to quickly reach diapause
when exposed to short days. Once diapause ends, the modulation of devel-
opmental time is reversed: larvae resume postdiapause development, but
grow at a slower rate under short days (spring—meaning more time available
to reach pupation) and a faster one under long days (summer—meaning less
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