Habitat preferences of Papilio alexanor Esper, [1800]: Implications for habitat management in the Italian Maritime Alps

Abstract

Papilio alexanor Esper, [1800] is a threatened European butterfly species listed in Annex IV of the Habitats Directive and in Appendix II of the Bern Convention, being considered extremely vulnerable to climate change. According to some projections (e.g., Bambu, a scenario of moderate climate change), it would be expected to lose 63% of its European climatic niche by the year 2050 and 77% by 2080. The few remaining populations are expected to become concentrated in the Maritime Alps. In 2009 and 2010, we studied what is probably the densest P. alexanor population in the Italian part of this area. It occurs in a series of dry, xerothermic grasslands, also partially occupied by an abandoned limestone quarry, where the larval food plant is Ptychotis saxifraga (L.) Loret & Barrandon. Females lay eggs in July, choosing patches where the food plants are higher and more abundant. The habitat preference, conservation status and survival of the early instars larvae of P. alexanor have been investigated in order to draw up conservation strategies for this species in the Italian Maritime Alps.

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Published on 27March 2015
169
ZOOSYSTEMA • 2015 • 37 (1)
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KEY WORDS
conservation,
butteries,
restoration ecology,
oviposition,
larval development.
urn:lsid:zoobank.org:pub:791830E1-C6B6-4064-984E-C9DBCA6816CD
Bonelli S., Barbero F., Casacci L. P.& Balletto E.2015. — Habitat preferences of Papilio alexanor Esper, [1800]: im-
plications forhabitat management in the Italian maritime Alps, in Daugeron C., Deharveng L., Isaia M., Villemant C. &
Judson M. (eds), Mercantour/Alpi Marittime All Taxa Biodiversity Inventory. Zoosystema 37 (1): 169-177. http://dx.doi.
org/10.5252/z2015n1a7
ABSTRACT
Papilio alexanor Esper, [1800] is a threatened European buttery species listed in Annex IV of the Habi-
tats Directive and in Appendix II of the Bern Convention, being considered extremely vulnerable to
climate change. According to some projections (e.g., Bambu, a scenario of moderate climate change),
it would be expected to lose63% of its European climatic niche by the year2050 and77% by2080.
e few remaining populations are expected to become concentrated in the Maritime Alps. In2009
and2010, we studied what is probably the densest P.alexanor population in the Italian part of this area.
It occurs in a series of dry, xerothermic grasslands, also partially occupied by an abandoned limestone
quarry, where the larval food plant is Ptychotis saxifraga (L.) Loret & Barrandon. Females lay eggs in
July, choosing patches where the food plants are higher and more abundant. e habitat preference,
conservation status and survival of the early instars larvae of P.alexanor have been investigated in order
to draw up conservation strategies for this species in the Italian Maritime Alps.
RÉSUMÉ
Préférences d’habitat de Papilio alexanor Esper, [1800]: implications pour la gestion de l’habitat dans les
Alpes-Maritimes Italiennes.
Papilio alexanor Esper, [1800] est un papillon diurne menacé en Europe et inscrit à l’Annexe IV de la
Directive Habitats dans l’Appendice II de la Convention de Berne. Cette espèce est considérée comme
particulièrement sensible aux changements climatiques. Selon quelques projections (ex.Bambu, un
scénario de changement climatique modéré), elle serait susceptible de perdre63 % de sa niche cli-
matique européenne d’ici à l’année2050 et jusqu’à77 % d’ici à2080. On peut s’attendre à ce que
les quelques populations survivantes soient concentrées dans les Alpes maritimes. En2009 et2010,
nous avons étudié la population de P.alexanor probablement la plus dense parmi celles qui habitent
la portion italienne de ce territoire. Elle occupe une série de friches semi-naturelles aussi bien qu’une
Simona BONELLI
(Corresponding author)
Francesca BARBERO
Luca Pietro CASACCI
Emilio BALLETTO
Department of Life Science and Systems Biology, Turin University
Via Accademia Albertina13 10123, Turin (Italy)
simona.bonelli@unito.it
francesca.barbero@unito.it
luca.casacci@unito.it
emilio.balletto@unito.it
Habitat preferences of Papilio alexanorEsper,[1800]:
implications forhabitat management
intheItalianMaritimeAlps

170 ZOOSYSTEMA • 2015 • 37 (1)
Bonelli S. et al.
carrière de roches calcaires aujourd’hui désaectée. Dans notre aire de travail la plante nourricière
des chenilles est Ptychotis saxifraga (L.)Loret & Barrandon. Les femelles pondent leurs œufs au mois
de juillet, choisissant les sites où les plantes nourricières sont particulièrement hautes et abondantes.
Les préférences écologiques, l’état de conservation et la survie des premiers stades du développement
larvaire ont été étudiés, en vue de proposer une stratégie pour la conservation durable de cette espèce
dans les Alpes maritimes italiennes.
MOTS CLÉS
conservation,
papillons,
écologie de la
restauration,
oviposition,
développement larvaire.
INTRODUCTION
Papilio alexanor Esper, [1800] is a swallowtail buttery with
a highly fragmented Euro-Central Asiatic range. It occurs
in the pre-Alps and sub-Mediterranean areas of SE France,
from Provence to the French Maritime Alps, while in Italy it
is restricted to a few sites of the Ligurian and Italian Maritime
Alps. Outside of this range, its distribution shows a very wide
gap and it occurs again in the southern Balkans, thereafter
extending eastwards as far as Central Asia (see Kudrna etal.
2011). e rst records of P.alexanor in the Italian Alps date
back to the 1970s, when it was found in the Ligurian Alps
(Balletto& Toso1976). In the Maritime Alps, apart from
an old, unconrmed report by Turati& Verity (1911-12;
“Col de Fenestre”) and a vaguely localized record by Barajon
(1957: “upper Val Tanaro”), its occurrence was conrmed
only much later (Baldizzone1971; Balletto etal. 1982). More
recent reports include those by Ortali& Bertaccini (1987),
Prola& Prola (1990), Sala& Bollino (1991), Audisio& De
Biase (1993), David& Sanetra (1994) and Arnscheid (2000).
e presence of this species in the general area had, however,
already been known to amateur collectors for a number of
years. Populations from the Italian Maritime Alps were later
described as “ssp. radighierii” Sala& Bollino, 1991. Elsewhere
in Italy, although P.alexanor was sometimes observed in some
parts of Calabria and NE Sicily (see Fig.2), it does not have
any stable populations there and the records probably refer
to stray adults having reached the Italian Peninsula by ying
from the Balkans.
In NW Italy, the species is found in mountainous or
hilly regions, from around500 to1200m a.s.l., but at least
occasionally reaching up to2100m, at Col de Tende (E.
Balletto, personal observation). Papilioalexanor is a ther-
mophilous species that prefers calcareous areas, on south-
facing, sometimes steep and rocky slopes. is is probably
a consequence of the ecological needs of its food plants,
some of which are pioneering species selecting eroded and
poor soils. e larvae feed on various Umbelliferae mainly
Ptychotis saxifraga (L.) Loret& Barrandon in the Maritime
Alps, but eggs can also be laid on Trinia glauca ssp. glauca
(L.) Dumort (C. Forte, pers. comm.; Nel& Chauliac1983;
Bollino& Sala2004). At lower elevations (500-800m) of
the southern slopes of the Ligurian Alps, caterpillars are
found on Opoponax chironium (L.) Koch (see Reche1978),
while populations from the Balkans and Central Asia may
use several species of Ferula (see de Freina1996). As observed
by Nel (1991), each population (or group of populations)
of P.alexanor generally shows a selected trophic preference
for a single Umbelliferous species, although the use of up to
three species, each having dierent blooming time, has been
reported locally for some populations from central Greece
(Pimpinellasp., O.chironium, Ferulagosp., see Bollino&
Sala2004). Similar local plant-shifts are also reported for
French populations, where larvae of P.alexanor, occurring
on opposite slopes of the same hill, select O.chironium on
the northern side and P.saxifraga on the southern. e same
authors (Bollino& Sala2004) argue that the capability of
exploiting various food plants having spaced-out blooming
times may represent an important adaptation in the case of
species having a prolonged emergence period. Papilioalexanor
is invariably monovoltine: adults may y from late March to
late July, depending on local climatic conditions (Bollino&
Sala2004). Papilioalexanor overwinters as a pupa, and is
known to stay in diapause for up to threeyears (Nakamura&
Ae1977; Bollino& Sala2004).
Although P.alexanor is listed in Annex IV (species of
community interest) of the Habitats Directive (H.D.) and
in Appendix II (strictly protected species) of the Bern Con-
vention, no immediate or major threats to its survival have
been identied at the European level and it is considered a
species of “least concern” (van Swaay etal. 2010). In2007
its conservation status under Article17 (H.D.) was assessed
as “favourable” for the Alpine region and “unknown” for the
Mediterranean area.
Papilioalexanor, however, is known to be particularly vulner-
able to climate change and Settele etal. (2008) have listed it
as an “Extremely high climate change risk (HHR)” species, in
their Climate Risk Atlas of the European butteries. Indeed,
climate change will soon represent a real threat for P.alex-
anor, which, according to currently available climatological
scenarios, is expected to lose63% of its European climatic
niche by2050, and77% by2080 (Bambu scenario). Given
its good dispersion ability, it is likely that P.alexanor will
respond to climate change by colonizing new, climatically
suitable areas, as suggested by recent observations of isolated
individuals far north from its actual range along the Rhone
Valley, especially during the warmest years. However, sites
in the SW Alps will be among the very few to oer suitable
conditions for its long-term survival, so that it is extremely
important to protect all populations in the Maritime and
Ligurian Alps, in order to mitigate the severe eects that
climate change will have on this species.

171
ZOOSYSTEMA • 2015 • 37 (1)
Papilio Alexanor in the Italian Alps
Even though no population extinction is known to have
aected P.alexanor on Italian territory (Bonelli etal. 2011a)
and all known Italian populations occur within protected areas
(Bonelli etal. 2011b), we recently estimated as “inadequate”
the overall conservation status of this species, on the basis of
its “future prospects”. Apart from climate, the other threats
that Italian populations are facing, and which may eventu-
ally undermine the long-term persistence of this species,
are mainly connected with the abandonment of traditional
pastoral systems and the return of natural forestation (H.D.
Article17 assessment; Balletto etal. in press).
Perhaps the most severely threatened Italian population of
P.alexanor occurs in Valdieri, in the Italian Maritime Alps
(Maritime Alps natural Park – SCI and SPZ IT1160056).
Adults occur there in a semi-natural habitat (calcareous rocky
slopes, Annex I Habitats Directive, 8210) and in an aban-
doned quarry. Among the most important factors threatening
this population we can cite: 1) over-collecting of both adults
and larval stages; 2) habitat loss, as a consequence of natural
forestation; and 3) possible opening of new quarrying activi-
ties in the area.
We investigated this population in the years2009 and2010
within the framework of the programme “Inventorying biodi-
versity in the Mercantour/Marittime”, funded by the European
Distributed Institute of Taxonomy (EDIT).
e main aims of our study are listed below:
–eld study of the local larval food-plant. ere is growing
evidence of polyphagous buttery species that are monopha-
gous at the local level. is would have strong inuence for
in situ conservation plans and general habitat management
(see Dolek etal. 2013);
–assess habitat preferences and food-plants use. We wanted
to evaluate whether females select oviposition sites on the
basis of microhabitat characteristics or of some specic plant
features, as observed in other buttery species (e.g., Patricelli
etal. 2011);
–assess the conservation status of P.alexanor at Valdieri sites.
Since swallowtails are large butteries and require substantial
amounts of food to reach their nal instars, it will be useful
to estimate the food plant density and the area of favourable
habitat needed to guarantee the long term survival of this
population.
Fig.1.—Papilio alexanor Esper, [1800]. Photograph: Davide Piccoli.

172 ZOOSYSTEMA • 2015 • 37 (1)
Bonelli S. et al.
MATERIALS AND METHODS
During the spring-summer period of2008, preliminary in-
spections were made to verify the persistence of the Papilio
alexanor population and conrm that Ptychotis saxifraga is the
larval food-plant in the study area.
Study area
Papilio alexanor adults and eggs occur in both natural and
semi-natural/replacement habitats.
e buttery was detected in the abandoned quarry of Valdieri
and in the nearby Natural Reserve for Juniperus phoenicea.
e limestone banks that extend to the East of Valdieri were
quarried for about35years (1962-1997) for the preparation
of cement (Ansaldi etal. 2006).
e Phoenician juniper (Juniperus phoenicea L.) is a very rare
plant in this part of Italy, where it reaches its northernmost
limit. A Natural Reserve for the protection of an important
stand of this shrub was therefore created in1984. It extends
over the limestone and dolomitic banks occurring close to
the summit of Mt Saben (1670m), where the southern
exposition and the presence of carbonate rocks favour the
establishment of a microclimate that is also suitable for the
survival of P.alexanor.
Sampling the food plantS and the larval inStarS
Papilioalexanor is a fast and high-ying swallowtail buttery
that always occurs at low densities, which makes studying its
adult’s population dynamics by the Mark-Release-Recapture
method unfeasible. During the summers of2009 and2010,
we surveyed the spatial distribution of P.saxifraga and focused
our eorts on nding the pre-imaginal stages of P.alexanor.
We randomly chose47quadrats (25m2) in which at least
one food plant existed. In each geo-referenced plot, we col-
lected data on the number and height of P.saxifraga plants. We
measured the density of the vegetation cover and the propor-
tion of bare soil, according to the Braun-Blanquet’s method
(Braun-Blanquet1932). For each plot, we noted the number
of eggs, larvae and/or pupae of P.alexanor. In2010, the same
data were collected once a week for the whole ight period
of P.alexanor adults, to assess the in-eld larval development
and the microhabitat requirements of the early stages. Finally,
we measured the minimum distance of each plot centre from
the nearest thistle plant (e.g., Carduus, Cirsium), since these
plants represent the only nectar sources potentially available
for P.alexanor adults at that time of the year.
In addition to eld sampling, we performed a study of the
survival of larval instars in the laboratory. Ten plants bear-
ing visible P.alexanor eggs were collected, carried and set in
laboratory conditions. We measured the length of each larval
stage during development and the number of days between
each moulting until pupation.
StatiStical analySiS
Data on plot parameters collected over the two-year samplings
were not normally distributed. erefore, we used non-
parametric tests to compare the ecological factors between
occupied and unoccupied plots, in separate tests.
We used the Wilcoxon test to compare the numbers and
heights of P.saxifraga plants, the percentage of bare soil and
vegetation cover, between plots occupied by P.alexanor in2009
and/or in2010. e Mann-Whitney-U test was used to assess
dierences in plot distances from the rst colonised square
and the rst nectar source (thistle) between plots, either oc-
cupied or unoccupied by P.alexanor.
In-eld larval survival was calculated for each plot, using
data collected in2010, as the number of larvae divided by
the initial number of eggs. Dierences in larval survival be-
tween plots occupied by increasing numbers of food-plants
were analysed with the Friedman two-way analysis of vari-
ance by ranks test.
RESULTS
During the summer of2008, we conrmed the persistence
of a P.alexanor population in the surroundings of an aban-
doned quarry in the Valdieri area. Larvae were feeding only
on P.saxifraga.
Survey of P. saxifraga
In2009, we surveyed the food-plants and counted P.alexanor
eggs, larvae and pupae. We identied47quadrats, spanning
in altitude from700to950m, where P.saxifraga was pre-
sent. At the base of the quarry, the food-plant was located
Fig.2.— Stable range of P.alexanor Esper,[1800] in Italy (Balletto etal. 2007),
observations from S Italy are interpreted as being based on vagrant specimens
from the Balkans, the arrow indicates the Valdieri study area.

173
Butteries of the Italian maritime Alps
ZOOSYSTEMA • 2015 • 37 (1)
mostly in the low-lying areas colonised by pioneer vegetation,
while at higher altitudes it occurred at the edge of the rocky
terraces and on the south facing rocky slopes. e data col-
lected in2009 allowed us to estimate P.saxifraga coverage:
about1.8hectares out of a total of29.5hectares surveyed,
while in2010 the proportion of habitat occupied by the food-
plant had slightly decreased, to1.5ha. In2009, 594plants
were scanned: 538of them (90.5%) did not present any trace
of P.alexanor eggs, while56 (9.5%) were exploited by the
buttery and occurred in28out of47quadrats.
Within each quadrat we counted from a minimum of
oneplant up to a maximum of50. e average height per
plot of the P.saxifraga plants varied between20 and 60cm.
In2009 we counted a total of65eggs, as well as70rst,
43second, 12third andfournal (IV and V) instar larvae,
and5pupae of P.alexanor.
In2010, we monitored the same47patches surveyed
in2009. e distribution of P.alexanor was similar to that
observed in2009, butnineplots were no longer occupied
in2010, whilevepatches unexploited in2009 were colo-
nized by P.alexanor in2010.
e comparison between the quadrats monitored in our
two-year study showed that occupancy depends on the number
of P.saxifraga plants (Wilcoxon test: Z = –2.745, p = 0.006)
as well as on the percentage of plant cover (Wilcoxon test: Z
= –2.251, p = 0.024) (Fig.3).
Quadrats colonized by P.alexanor also showed a statistically
signicant larger average height of the food-plants (Wilcoxon
test: Z = –2.506, p = 0.012) (Fig.3). e percentage of bare
soil showed no statistically signicant correlation, while the
percentage of other vegetation was lower in the quadrats oc-
cupied by the buttery (Wilcoxon test: Z = –2.254, p = 0.024).
In2010, 21patches were exploited by P.alexanor while26were
unoccupied. A total of423P.saxifraga plants were examined,
356of which (84%) did not bear any pre-imaginal P.alexanor
pre-imaginal instars, while67 (16%) were occupied by the
buttery. Inside quadrats we surveyed a minimum of oneplant
up to a maximum of32 plants. In quadrats with P.alexanor
we examined an average (± SD) of11.2 ± 8.6 plants.
Comparing distances between quadrats, we found that
plots where P.alexanor was absent were much more isolate
than those where the buttery was present (Mann-Whitney
U = 84.000, p = 0.18), the distance between the occupied
quadrats and the nearest quadrat was lower (15.81 ± 4.23m,
mean ± SE) than the distance between unoccupied quadrats
(27.22m±4.50; mean±SE) (Fig.3). Distances from the
nearest potential nectar source (thistles) were similar for plots
where P.alexanor was present and those where it was absent
(Mann-Whitney U = 155.5, p = 0.950).
larval Survival in the laBoratory
On average, larval development lasted22days and the transi-
tion from one stage to the next took4.1 ± 0.7days (Fig.4).
Larval body length changed from0.4 ± 0.15cm (rst instar
larvae) to a maximum of4.3 ± 3.5cm (nal instar larvae).
e development of the last instars was faster than that
of rst instar larvae: P.alexanor took about a week to pass
fromthird instar larva to pupa (Fig.4). Two butteries hatched
in the laboratory and were transported, along with the reared
pupae, to their site of origin.
present presen
tp
resent
absent absen
ta
bsent
Occurrence of P. alexanor Occurrence of P. alexanor Occurrence of P. alexanor
Height of P. saxifraga
plants (cm)
50
40
30
20
10
0
Number of P.saxifraga
plants
0
5
10
15
20
Percentage
of vegetation
60
50
40
30
20
10
0
A
BC
0510 15 20 25
0
5
4
3
2
1
e
I
II
III
IV
p
Days
Lenght (cm)
F
ig
.3.— Comparison of mean number (A) and height (± SD) (B) of P.saxifraga (L.) Loret& Barrandon and average percentage (± SD) of vegetation cover
(C) between occupied and unoccupied plots by Papilio alexanorEsper,[1800].
F
ig
.4.— Pre-imaginal development of Papilio alexanor Esper,[1800] under
laboratory conditions; box plots illustrate increases in larval length; vertical
lines: median larval length; box: 25th-75th percentiles; whiskers: minimum
and maximum observed values;  outliers; dark grey band width represents
the standard deviations of mean development intervals (days). Abbreviations:
e,egg; I-IV, larval instars; p, pupa.

174 ZOOSYSTEMA • 2015 • 37 (1)
Bonelli S. et al.
development of P. alexanor in field conditionS
During2010, we observed a total of369eggs and268larvae.
P.alexanor females laid on average2.4 ± 0.99eggs (mean ±
SD) on each plant, but in quadrats where the availability of
plants was more limited (oneor few P.saxifraga plants), we
found a higher number of eggs per plant (up to20 eggs per
plant). Within quadrats, depositions occurred on an average
of2.6 ± 1.6 plants (min1; max5), even when plant avail-
ability was high.
As can be seen in Fig.5, the highest number of eggs was
found in the rst week of sampling (6.81 ± 1.02 eggs per quad-
rate), whereas this value decreased during July, as the number
of caterpillars increased. e highest number of larvae was
observed during the third sampling (on average5.04±0.73
larvae per quadrat). In total, 24 larvae atfourth orfth instar
were found during the last two sampling events, as well as a
single pupa (Fig.5).
We calculated the survival rates of three larval stages, from
the second to the nal instar larva. Survival was similar for
second and the third instar larvae, while it was signicantly
lower for nal instar larvae. Comparing the average larval
survival within each quadrat with the available number of
food-plants, we observed that larval survival increased when
the number of plants was higher than15 (Friedman’s test:
Z= 6.000; p = 0.05) (Fig.6).
DISCUSSION
Our results show that P.alexanor is locally monophagous: in
the study area it strictly exploits Ptychotis saxifraga, whereas
at a locality situated 70km away (above Latte, in the prov-
ince of Imperia) it exclusively feeds on Opoponax chironium.
Local exploitation of a single food-plant species commonly
occurs in otherwise polyphagous butteries (e.g., Finke&
Scriber1988), including Papilionidae Latreille, 1802. For
example, Papilio glaucus (Linnaeus, 1758) is considered one of
the most polyphagous species among Papilionidae (563species
in the world), using more than14dierent families as food-
plants in North America, yet this species is monophagous in
Florida (Scriber1986). Studies carried out under laboratory
conditions have shown that the larvae of Papilio troilus (Lin-
naeus, 1758) are associated with various species of Lauraceae,
depending on the geographical area. More specically, they can
survive on three dierent plants, but growth rates are inu-
enced by the essence selected (Nitao etal. 1991). Euphydryas
maturna (Linnaeus, 1758) is another example of a buttery
locally using dierent plants. Females lay their eggs on the
leaves of Fraxinus excelsior L. and the whole pre-hibernation
development takes place on this plant. After winter diapause,
larvae are known to feed on various common herbaceous
species, such as Pulmonaria ocinalis L., Lonicera coerulea
L., Veronica longifolia L., Violasp. and Plantago lanceolata
L. (see Freese etal. 2006). Nonetheless, at the local level the
buttery selects only one plant species to feed on, even where
many more are available (Dolek etal. 2013).
eoretical studies predict that butteries, like many other
phytophagous insects, will tend to specialise, moving from
polyphagy to monophagy, since any new mutation that in-
creases tness on the primary host plant, and thereby lowers
the relative tness on secondary host plants, will be favoured
by selection (Bernays& Graham1988, Futuyma& More-
no1988, Futuyma2008). Furthermore, at least in some cases,
additional advantages may be achieved by entering relatively
competitor-free new spaces (Wiklund& Friberg2008).
Local adaptations to single host plants may have favoured
dierentiation between populations, which might eventually
culminate in speciation. In fact, the two populations from
Latte (IM) and from Valdieri have been described as separate
subspecies (respectively P.alexanor alexanor Esper, 1799and
P.alexanor radighierii Sala& Bollino, 1991) and should at
least be considered separate ESUs (Casacci etal. 2013). Dier-
ences between them may have become xed and maintained
by the temporal shift (about15days) in the blooming of the
Number of P. alexanor larvae
and eggs (mean ± SE)
Sampling dates
9
8
7
6
5
4
3
2
1
0
9 July 2010 15 July 21 July 28 July
I instar
II instar
III instar
IV+V instars
eggs
tot. larvae
Fig.5.— Pre-imaginal development of P.alexanor Esper, [1800] in the eld. Mean (± SE) number of eggs and larvae of P.alexanor collected during thefour sam-
pling events (2010).

175
Butteries of the Italian maritime Alps
ZOOSYSTEMA • 2015 • 37 (1)
two locally selected host plants. Nevertheless, in standard
eld guides the descriptions of many buttery species are
accompanied by long lists of food-plants. Exact information
on regional food-plant variation is, in contrast, relatively
scanty, or provided independently from information on the
ecological context.
In our study area, oviposition took place at the end of June,
in agreement with the phenology of P.saxifraga, and larval
development occurred in July. e females laid few eggs per
plant (2.4 ± 0.99 on average), as previously reported (Bol-
lino& Sala2004), and used on average2.6plants close to each
other for oviposition (no more than veplants per plot), even
when many plants occurred in5× 5m plots. Females scattered
their eggs and this may increase their ospring’s tness. As
observed in eld and laboratory conditions, P.alexanor larvae
need to feed abundantly to exponentially increase their body
mass in about20days. Moreover, it is likely that females lay
few eggs per plot to prevent cannibalistic behaviour, which
is very frequent among buttery caterpillars.
If we assume that: 1) the sex ratio in the P.alexanor population
is balanced; 2) a single P.alexanor female lays about70/80eggs
during a season (C. Forte, pers. comm.); and 3) the area
covered by P.saxifraga was of1.5ha, having sampled the egg
density, the adult population size in the study area can be
estimated at about 170individuals. Nevertheless, it seems
that the Valdieri population, and therefore also its ovipos-
iting preferences, may be considered representative of the
whole “subspecies radighierii”, which often occurs in small,
but constant and isolated areas, supported by a few hectares
of suitable habitat (Bollino& Sala2004).
Our results indicate that the number of food-plants and their
height are among the factors that aect the number of eggs
laid by P.alexanor females on Ptychotis saxifraga plants. Female
preference for plots with high numbers of plants is conrmed
by the fact that larval survival is higher in those plots where
the number of P.saxifraga was more than15plants (Fig.6),
probably because larvae need to consume a great amount
of food. Selection for oviposition of higher plants, usually
having a larger number of owers, is common in butteries
(e.g., Myers etal. 1981; Courtney1984; Bonelli etal. 2005;
García-Barros& Fartmann2009). e most likely explana-
tion is that gravid females choose visually prominent plants
as their main targets. e fact that the distances of occupied
plots from the rst patch with P.saxifraga plants were lower
than between unoccupied plots suggested that, as for other
Lepidoptera (ompson& Pellmyr1991), P.alexanor females
accept or reject certain plants as oviposition sites on the basis
of their relative position in the food-plant community. On
the contrary, we did not observe any dierence in distances
from the nearest nectar source between occupied and unoc-
cupied patches. is is consistent with the high vagility of
P.alexanor adults, which can easily move from nectar sources
to oviposition sites, as well as with the relative abundance of
thistle plants, which were probably sucient to support the
whole P.alexanor population.
In agreement with observations by Bollino& Sala (2004),
eggs took about sevendays to hatch in the eld, but developed
slightly faster under laboratory condition (aboutve days).
As noted in the literature, and observed in the eld, larvae
were easy to observe, not only because of their aposematic
colours, but also for their strong heliotropism, which causes
them to remain on a ower stem and rarely abandon it. Just
before pupation, the caterpillar usually abandons the plant
and pupates near the ground, indicating a negative photo-
taxis (Nakamura& Ae1977). e low number of individu-
als observed belonging to the last two stages may be related
to the diculty, already noted by other authors (Bollino&
Sala2004), of nding the last instar larvae and the mimetic
pupae of P.alexanor in eld conditions.
conServation remarkS
In order to ensure the persistence of the population in the
study area, we suggest that it is important to take into account
both the ecological features of the resident P.alexanor popula-
tion (e.g., the correlation between food plant’s phenology and
larval development) and the most signicant local threats. In
particular it will be crucial to discourage collectors of both
adults and larvae. Sampling could be authorized only for
justied scientic reasons and surveillance of the area should
be ensured, at least during June-July.
At the same time, to ensure the long-term persistence of the
species it would be essential to create new habitats, or restore
old ones, where suitable growth condition for P.saxifraga can
be assured. In this context, the restoration of the abandoned
quarry would generate very good opportunities for P.alexanor.
Butteries frequently benet from areas transformed by
human activity, such as pastures or areas under non-intensive
cultivation, and suer rapid decline when these habitats
are abandoned or altered (Balletto etal. 2007; Skorka etal.
2007). Less common, but relatively well known, are cases of
colonisation of quarry areas. A study performed in Bedford-
shire (UK) (Turner etal. 2009) has shown the settlement of
a riodinid buttery, Hamearis lucina (Linnaeus, 1758), at an
abandoned chalk quarry, fostered by the availability of its
larval food plants. e latter benet from disturbance caused
by occasional landslides on unstable ground and prefers steep
slopes, in the lee of the wind. us, as also demonstrated by
0
0.1
0.2
0.3
0.4
0.5
0.6
Larval survival
Number of P. saxifraga
<10 10-15 >15
L2 L3 L4 + 5
F
ig
.6.— Comparison of mean values of larval survival rates calculated for
the21 (occupied) plots ranked by increasing total number of P.saxifraga (L.)
Loret& Barrandon plants.

176 ZOOSYSTEMA • 2015 • 37 (1)
Bonelli S. et al.
the present study, abandoned quarries may harbour a signi-
cant number of plants, many of which may be characterized
by peculiar ecological requirements. Results of a comparative
study of buttery communities, carried out in 21limestone
quarries at the Moravian Gate (Czech Republic), showed that
quarries can provide secondary habitats for xerophilous spe-
cies, replacing the calcareous grasslands once densely popu-
lated by butteries (Benes etal. 2003). Butteries beneting
from these areas are rare xerophilous species and/or sedentary
species. eir abundance, even during extraction activities,
conrms the idea that quarries can contribute signicantly
to the conservation of these butteries.
Unfortunately, the ecological requirements of these species
are rarely considered in the management commonly practiced
in the dry grasslands of Central Europe, which typically aims
at protecting sites where charismatic plants (e.g., orchids)
are growing (Balmer& Erhardt2000; Kahmen etal. 2002).
Acknowledgements
e All Taxa Biodiversity Inventory + Monotoring Mercantour/
Alpi Marittime was launched by the European Distributed
Institute of Taxonomy (EDIT) project (2006-2011).
We thank M.De Biaggi (Parco Alpi Marittime) and
M.-F.Leccia (Parc national du Mercantour).
We also thank the Municipality of Valdieri and L.Mavilla,
C.Forte and E.Rivella (Regional Agency for Environmental
Protection – ARPA Piemonte). We are also grateful to the
reviewers for their helpful comments.
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Submitted on 5March 2014;
accepted on 20October 2014;
published on 27March 2015.