(Quartau & Boulard, 1995) is a small-sized cicada
species endemic to the southern Iberian Peninsula,
whose ecology and current distribution is still
poorly known. As a result of recent fieldwork in
southern Portugal and Spain we report here an update
on the distribution range and habitat preferences of
T. mariae. The distribution of this species
is mostly restricted to the coast of central Algarve
(Portugal) and Huelva province (Spain), being
heavily fragmented and discontinuous. The species
appears to be habitat-specific, with a clear
preference for habitat with stone pine and at a
close distance from the sea. The conservation of
T. mariae populations is threatened,
particularly in Algarve, by anthropogenic changes in
land use and habitat loss and by interspecific
competition with its sibling and wider distributed
T. argentata. We advocate that T. mariae
should be considered as an endangered species and
should be subjected to annual monitoring.
Cicada, conservation, distribution range, habitat,
land use, Portugal.
(Quartau & Boulard, 1995) é uma espécie de cigarra
de pequeno porte endémica do sul da Península
Ibérica, com ecologia e distribuição ainda pouco
conhecida. Em resultado de intensivo trabalho de
campo desenvolvido recentemente no sul de Portugal e
Espanha, reportamos aqui uma atualização da
distribuição e preferências de habitat de T.
mariae. A distribuição desta espécie encontra-se
praticamente restrita à costa da região central do
Algarve (Portugal) e da província de Huelva
(Espanha), sendo bastante fragmentada e descontínua.
Esta espécie parece ser especialista em relação ao
habitat, com uma clara preferência por zonas com
pinheiro manso e perto do mar. A conservação das
populações de T. mariae está ameaçada,
principalmente no Algarve, por mudanças
antropogénicas no uso o solo e perda de habitat e
pela competição com a espécie-irmã mais largamente
distribuída, T. argentata. Defendemos que
T. mariae seja considerada uma espécie ameaçada
e sujeita a monitorizações anuais das suas
Palavras-chave: Cigarra, conservação, área de
distribuição, habitat, uso do solo, Portugal.
are common in Mediterranean countries and quite
famous for their conspicuous calling songs in the
summer. Cicadas have long larval stages spent
underground and a short winged adult stage that
lasts for a few weeks (Claridge 1985,
Boulard & Mondon 1995, Williams & Simon
1995). The calling songs are produced by adult males
to attract females for reproduction and are usually
species-specific (Boulard 2006). The biology and
accurate distribution range of most Mediterranean
cicadas is still poorly known. In fact, several
cicada species were recognized only recently,
highlighting the importance of the Iberian Peninsula
as a hotspot for cicada diversity and endemism (Boulard
Quartau & Boulard 1995,
Puissant & Sueur 2010).
A recent study confirmed the occurrence of 13 cicada
species in Portugal (Sueur
et al. 2004).
Most Portuguese cicadas have currently a scattered
distribution, which in some cases makes them
cicadas are believed to have very restricted
distribution ranges but extensive field surveys are
missing and accounts of occurrence in literature
mostly result from the accumulation of singular
observations by different authors.
(Quartau & Boulard, 1995) is a small-sized cicada
that was described from Quinta do Lago, Algarve, and
was thought to be endemic to this region of
Portugal, until its recent discovery in the province
of Huelva, in Spain (Simões et al. 2013). This
species may occur in sympatry with other
Tettigettalna species, namely T. argentata
(Olivier, 1790) and T. josei (Boulard, 1982)
(Quartau & Boulard 1995, Sueur et al. 2004, Nunes et
al. 2014). Tettigettalna mariae and its
sibling T. argentata are morphological and
genetically very similar but their calling songs are
distinctive and species-specific (Quartau & Boulard
1995, Nunes et al. 2014, Mendes et al. 2014).
As an outcome from intensive fieldwork for species
of genus Tettigettalna in both southern
Portugal and Spain during the summers of 2011-2013,
we report here an update on the distribution range
and habitat preferences of T. mariae. The
results confirm the restricted and fragmented
distribution of the species, being particularly
vulnerable to habitat loss in Algarve, where human
pressure is higher.
Materials and methods
Several field surveys were conducted in the southern
Iberian Peninsula (regions of Algarve and Andalusia)
from the end of June until mid-August during the
summers of 2011-2013. The fieldwork took place from
10:00 am to 07:00 pm with sunny weather and with
temperatures ranging from 24º to 39ºC. Initial
searches in 2011 were conducted by driving a car at
low speed to allow the detection of the calling
males. In 2012 and 2013, searches were targeted to
areas of potentially suitable habitat. Specimens
were located and identified through their
species-specific calling song (Fig. 1). Geographical
coordinates of each location were determined with a
GPS (Garmin, Oregon series 550t). Several male songs
were recorded using a Marantz PMD 661 Portable SD
recorder (20Hz – 24kHz) connected to a Telinga Pro 7
Dat-mic microphone (Twin Science) following the
procedures given in Simões et al. (2000). Species
confirmation was accomplished with time and
frequency analysis of sound recordings using the
software Avisoft Sas-Lab Pro (Specht 2012) as
described in Simões et al. (2013).
Figure 1. Sonograms (bottom right) of typical
male calling song of Tettigettalna mariae and
from its sibling Tettigettalna argentata. The
stone pine (Pinus pinea) wood photo was taken
in July 2013, near Cartaya (Huelva, Spain) and
corresponds to the location where the largest
population of T. mariae was found so far.
Field prospections in Algarve were at first
conducted near the coast, along road N125, from the
Atlantic west coastline, near Vila do Bispo, until
river Guadiana bank at Vila Real de Santo António.
Singing males of Tettigettalna mariae were
found only in central Algarve, confirming its
presence in all locations previously reported in the
literature (Sueur et al 2004), except for Castro
Marim. Searches in and around Castro Marim were
unsuccessful in both 2011 and 2012 summers. We
refined our fieldwork in central Algarve to
understand the real extent of the species
was found in several locations around Quinta do Lago,
Vale do Lobo, Quarteira, Vale Navio and Vale Judeu
We found Sesmarias as
the westernmost location and the pine woods at the
west vicinity of Faro city as the easternmost
occurrence (Long. 8º18’ to 7º57’W). The latitudinal
range is small (37º00’ to 37º07’N). It goes from the
coastline and does never exceed 7km inland along its
Map of the southwest of the Iberian Peninsula with
occurrence records for Tettigettalna mariae
in the summers of 2011-2013.
In Spain, the occurrence of the species beyond the
east bank of Guadiana river, in the province of
Huelva (Andalusia), was first noticed in 2012 (Simões
et al. 2013) and confirmed in the summer of 2013.
Our searches were extended to other Andalusian
provinces and Murcia, but T. mariae
distribution remains restricted to the province of
Huelva and partially to the province of Sevilla
(Fig. 2), between Cartaya and Coria del Rio.
We noted a remarkable coincidence between the
distributions of T. mariae and of stone pine
(Pinus pinea) in south western Iberia (Fig.
3). These pine woods are mostly present in sandy
soils near the coast. At some locations, T.
mariae males were found singing on almond (Prunus
carob trees (Ceratonia siliqua), in shrubs
such as acacia (Acacia longifolia), ladanum (Cistus
common blackberry (Rubus fruticosus) and on a
small abandoned orange (Citrus sinensis) tree
In all these cases, patches of stone pine forest or
dispersed or isolated stone pines could be found in
the immediate vicinity. The majority of males were
usually perched on pine branches while singing and
the location with the highest number of T. mariae
males singing in allopatry was found in Spain,
within a stone pine forest near El Portil (Lat.
37º13’32.0’’N, Long. 07º02’05.4’’W). This forest
was partially planted, with T. mariae singing
on relatively young trees, with 3-4m height and
about 2m wide crowns, producing little shade. Shrubs
and ladanum (C. ladanifer) in particular were
abundant under pine trees, but T. mariae
males were consistently singing perched on pine
Figure 3. Distribution of stone pine (Pinus
pinea) in the Iberian Peninsula (source: Benito
Garzón 2006, available at
Distribution of stone pine in Algarve (Portugal).
The map dates from 1980 and is available at
Distribution of stone pine in the southwest of Spain
Auñón and D. Sanchez de Ron, available at.
was found in sympatry or parapatry (< 500m) with
singing males of T. argentata in a high
proportion of occurrence points (Fig. 4),
particularly in Algarve, around Vale do Lobo and
Quinta do Lago (Fig.
The coastline of Algarve is heavily urbanized, and
most habitat patches where T. mariae occurs
are surrounded by touristic villages or golf
courses, mainly in Vale do Lobo and Quinta do Lago (Fig. 5).
The occurrence of T. mariae in Algarve is
particularly sparse between the localities of
Sesmarias and Vale do Lobo. The occurrence of stone
pine in this area is also quite dispersed, the
landscape being dominated by agriculture (small
crops of almond, carob, fig or olive trees),
uncultivated scrublands and human occupation.
Figure 4. a) Distribution map of
Tettigettalna mariae (light red circles)
overlapped with occurrence points of the sibling
species, T. argentata (blue triangles). b)
Pie chart of occurrence points of T. mariae
in Portugal (n = 48) and Spain (n = 13) classified
as allopatric or as sympatric or parapatric with
Figure 5. Land cover in the area between
Vale do Lobo and Quinta do Lago, in central Algarve
coastline, where most occurrence points of
Tettigettalna mariae were registered (source: ©
Google Earth). Most of the area is heavily
humanized, with touristic villages, golf courses and
unnatural green landscapes. Isolated stone pines are
maintained in humanized areas for its ornamental
value. Small forest areas with natural stone pine or
mixed stone and maritime pine persist between Vale
do Lobo and Quinta do Lago and harbour sympatric and
parapatric populations of T. mariae and T.
The acoustic surveys of 2011-2013 confirmed that the
distribution of Tettigettalna mariae is
currently restricted to the southwest coast of the
Iberian Peninsula and is heavily fragmented and
discontinuous. The species appears to be
habitat-specific, with a clear preference for
habitat with stone pine and at a close distance from
the sea. The distribution of stone pine in the
Iberian Peninsula is fragmented as well and mainly
found on coastal and inland sands or on very poor
soils (García del Barrio et al. 2013). The province
of Huelva harbours the largest area of stone pine
forest (Fig. 3) but this Mediterranean pine species
is presently absent in more than 60% of the
landscapes or biogeoclimatic classes that it could
inhabit in Spain (García del Barrio et al. 2013).
Paleoecological evidence suggests that the
distribution of stone pine was more extensive in the
past (Martínez & Montero 2004, Benito Garzón et al.
2007, Valbuena-Carabaña et al. 2010).
Reconstructions of past distribution of stone pine
for 6000 years before present indicate that it was
nearly continuous between central Algarve and Huelva
(see Benito Garzón 2006).The present fragmentation
in T. mariae’s distribution probably reflects
the effect of long term decline of suitable habitat.
Previous records of T. mariae in Castro Marim
from late nineties (Sueur et al. 2004) were not
confirmed by our surveys in two consecutive years.
The two main distribution areas (central Algarve and
Huelva) are now separated by nearly 70 kilometers.
We predict that this distance is large enough to
keep populations of both areas nearly or completely
isolated. Simões & Quartau (2007) determined through
mark-recapture that males from a large-size cicada (Cicada
orni) survive for about two weeks and their
flight dispersal does not exceed 150m. Instead of
investing in dispersal, males of C. orni tend
to remain close to their emergence site and maximize
their chances of mating with a strong singing
activity. It is likely that Tettigettalna
cicadas use a similar strategy and the dispersal of
both sexes is expected to be small.
Habitat loss and fragmentation are considered great
threats to biodiversity. However, not all species
are equally vulnerable to extinction risk (reviewed
by McKinney 1997). This uneven vulnerability
probably results from the specific ecological
attributes/life-traits of a species, which determine
how well it is able to withstand the external
threats to which it is exposed (McKinney 1997). In
addition, a combination of more than one of these
traits further increases a species risk of
extinction (Lawton 1994). For instance, species that
are both rare and specialized can be especially
vulnerable to extinction (Davies et al. 2004).
Considering the distribution update and habitat
preference reported here, we consider that T.
mariae is highly vulnerable to habitat loss
caused by changes in land use or forest fires, which
often jeopardize pine woods during the summer, when
adult cicadas have their breeding season and lay
their eggs on trees or bushes. This risk is greater
in Algarve, where the anthropogenic pressure is more
intense, than in southern Spain, where large patches
of stone pine are maintained as forest areas.
However, both in Portugal and Spain, T. mariae
populations are frequently overlapped or
surrounded by populations of T. argentata, a
more generalist species and with a much wider
distribution. Species-specific calling songs are
probably determinant to restrict gene flow between
these sibling species. However, according to
sequence data from the mitochondrial gene cytochrome
oxidase I (Nunes et al. 2014), there is no genetic
differentiation of T. mariae from T.
argentata populations surrounding or overlapping
T. mariae’s distribution range. In fact, some
haplotypes are shared between these species and it
remains unclear whether haplotype sharing is due to
past or recent introgression
or to incomplete lineage sorting.
Interspecific competition between sibling species
with similar requirements might represent an
additional threat for the persistence of T.
We advocate that T. mariae, being one of the
rarest Portuguese cicadas and a
Iberian endemism, should be considered as an
endangered species. Since
we believe that
should be taken
and it should be considered
an endangered species
in the category of
Annual monitoring for T. mariae populations
should be implemented at more sensitive areas such
as Vale do Lobo and Quinta do Lago, as well as in
suitable habitats near Castro Marim, to evaluate the
species ability to expand its range and recolonize
from adjacent areas in Spain.
should be preserved,
planting of new
fire prevention method,
only be done
in mosaic and
without soil plowing,
which may expose
the underground nymphs
(Quartau 2009, Quartau & Mathias 2010).
This study was financially supported by the project
PTDC/BIA-BIC/115511/2009 of the Portuguese
Foundation for Science and Technology (FCT). We
thank Bruno M. Novais for help during fieldwork.
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