Content uploaded by Roberto Battiston
Author content
All content in this area was uploaded by Roberto Battiston on May 08, 2017
Content may be subject to copyright.
25
Onychium, 13: 25-30 Published 20 April 2017
ISSN: 1224-2669 www.onychium.it
http://dx.doi.org/10.5281/zenodo.546318
The silent spreading of a giant mantis: a critical update on the distribution of
Sphodromantis viridis (Forskål, 1775) in the Mediterranean islands
(Mantodea: Mantidae)
Roberto BATTISTON1,a, Simone ANDRIA2,b & Gianpaolo RUZZANTE3,c
1 Musei del Canal di Brenta, Palazzo Perli, via Garibaldi 27, I-36020, Valstagna (Vicenza), Italy;
2 Corso Umberto 99, I-09070, Seneghe (Oristano), Italy;
3 Via Giardini 132, I-09127, Cagliari, Italy.
E-mail: a roberto.battiston@museivalstagna.it; b seyesimoneandria@gmail.com; c ruzzpa@tiscali.it
Abstract. The recent discovery of new specimens of Sphodromantis viridis (Forskål, 1775) in Sardinia, together
with the increasing number of sightings reported in the literature for the Mediterranean islands, allow a first
investigation of the spreading dynamics of this mantis outside of its continental habitat in a biogeographical
perspective. An insect historically absent from the Mediterranean islands but often sinantropic and with spreading
dynamics compatible with an anthropogenic dispersal is confirmed here as a new species for Italy.
Riassunto. La silenziosa diffusione di una mantide gigante: un aggiornamento critico sulla distribuzione di
Sphodromantis viridis (Forskål, 1775) nelle isole del Mediterraneo (Mantodea : Mantidae). La recente scoperta di
nuovi esemplari di Sphodromantis viridis (Forskål, 1775) in Sardegna, congiuntamente al sempre maggior numero
di avvistamenti segnalati in letteratura per le isole mediterranee, consente una prima indagine sulle dinamiche di
diffusione di questa mantide al di fuori del suo habitat continentale in u na prospettiva biogeografica. Un insetto
storicamente assente dalle isole mediterranee, ma spesso sinantropico e con dinamiche di diffusione compatibili
con una dispersione antropogenica, viene qui confermato come una specie nuova per l’Italia.
Key words. Mantodea, distribution, taxonomy, biogeography, alien species, natural spread.
Introduction
Sphodromantis Stål, 1871 is a xerothermophilous mantid genus (Mantodea: Mantidae), widely
distributed in the African continent, and Sphodromantis viridis (Forskål, 1775) is the species with the
northernmost distribution of the genus, reaching Southern Europe and the Mediterranean coast of the
Middle East (LA GRECA, 1966). LA GRECA (1966) proposed a subdivision of the species into four
subspecies belonging to different geographic areas of the Mediterranean and North Africa: S. v.
viridis (Forskål, 1775) for the Egyptian area, where the typical specimens have been described by
Forskål, S. v. meridionalis La Greca, 1950 in Ethiopia and Sudan, S. v. barbara La Greca, 1967 in
Tunisia, Libya and the Algerian Sahara, S. v. occidentalis Werner, 1906 in the countries of the
Atlantic coast of Africa from Morocco to Cameroon. Sphodromantis viridis simplex La Greca &
Lombardo, 1987 is described in a second time by LA GRECA & LOMBARDO (1987), for Ethiopia and
Somalia. LA GRECA (1966) did not examine any specimens from the Middle East and examined only
few specimens from Spain, the only European country where this species was known at the time, and
this lead him to speculate over a possible European subspecies, never described because of the lack of
available material. ROY (2010) reviewed the positions of LA GRECA (1966) stating the impossibility to
identify all the specimens of West Africa and Europe as a single homogeneous group, questioning the
validity of S. v. occidentalis as subspecies. ROY (2010) rehabilitated the name “vischeri” for the
subspecies from Spain using the historical specimen Stagmatoptera vischeri described by WERNER
(1933), but left open the question on the West African populations.
Onychium, 13: 25-30
26
The recent discovery of individuals of this species, historically absent from the insular areas of the
Mediterranean and in particular the study of new specimens collected in Sardinia, the examination of
their external morphology and the shape of the male genitalia, allow to give an overview on
expansion dynamics of this species.
Material and methods
In this work, we considered the scarce but detailed literature of this species, with particular attention
to the records of the Mediterranean specimens. We examined, with a Leica Wild M3c
stereomicroscope and a 2mpxl 50-200x digital microscope, the external shape and genitalia of: 1) an
adult male captured in a mist-net in 2011 on the island of Spargi in Northern Sardinia (RUZZANTE &
LEO, 2012); 2) an adult male found by the authors in 2016 in the Sardinian hinterland, attracted by the
artificial lights of a parking lot in the countryside of Santa Giusta (Oristano). The morphology of
these two specimens was then compared with the ones reported in the literature and with the
specimens preserved in the private collections of the authors.
Results
Examined specimens. Sardinia: Zinnigas, Santa Giusta (Oristano), 39°51'45.9'' N 8°36'27.0'' E
(WGS84), 2 m a.s.l., 5.VIII.2016, S. Andria leg., 1 adult male, coll. Battiston; La Maddalena (Olbia-
Tempio), Isola Spargi, Cala Corsara, 41°13'47'' N 9°20'49'' E (WGS84), 4 m a.s.l., 19.X.2011, G.
Ruzzante leg., 1 adult male, coll. Ruzzante.
The examination of the genitalia of these two males showed a perfect compatibility with the
variability reported by LA GRECA (1966) for the Spanish specimens, in particular with those from
Cordoba and especially for the shape of the pronotum and of the pseudophallus (Fig. 1), the latter
with almost straight and sub-parallel margins, distinct from the curved ones from Africa. These two
characters, referred as diagnostic in the taxonomy of LA GRECA (1966), highly variable within the
occidentalis group show a remarkable homogeneity among the Sardinian specimens (Fig. 2) and
between the Sardinian and Iberian specimens. This similarity and the marked differences from all the
other known subspecies lead to assume the existence of a Sardinian population probably related to the
Iberian subspecies. It must be noted that only a deep molecular analysis on a greater number of
specimens will be able to absolutely confirm it but, since the shape of male genitalia proved to be a
good taxonomical character to separate geographical populations, these specimens can be therefore
identified as Sphodromantis viridis vischeri (Werner, 1933).
Fig. 1. Male genitalia of S. viridis vischeri (Werner, 1933). 1) left epiphallus and hypophallus of the specimen
from Santa Giusta in dorsal view; 2) pseudophallus of the specimen from Spargi; 3) pseudophallus of the
specimen from Santa Giusta; 4) pseudophalli of the specimens from Cordoba – Spain (from LA GRECA, 1966).
Scale bar: 1 mm.
Onychium, 13: 25-30
27
Fig. 2. Males habitus of S. viridis vischeri (Werner, 1933): specimen from Spargi on the left and specimen from
Santa Giusta on the right. Scale bar: 10 mm.
Discussion
The two Sardinian specimens, despite their numerical scarcity that excludes any statistical
consideration, contextualized with the previous data collected in the Mediterranean, allow a first
critical analysis of the process of spreading of this species in the Mediterranean islands. According to
La Greca’s biogeographical analysis of the morphology of the subspecies he described, this species
began its spreading north of the Sahara probably in the Pleistocene, during the warming of the
Mediterranean climate, colonizing new territories thanks to the new conditions favourable for its
biology (LA GRECA, 1966). The isolation of the northern populations has presumably occurred during
the post-glacial period and considering the Iberian climatological history (MARTINEZ-CORTIZAS et al.,
1999) we can hypothesise that the arrival of this mantis has been relatively recent in Europe, perhaps
during the Medieval age characterized by a significant rise of the temperatures, and definitely before
the XX century. BOLIVAR (1898) indeed mentions this species as well spread in the Spanish territories
of Andalucia: Malaga, Seville and Cadiz. Its presence in Portugal is reported only later in 2008 and in
the southern areas near the Spanish border (MARABUTO et al., 2014). The same authors however make
evident a historic lack of mantidological records for that country that may have kept the presence of
this species hidden in previous years. From Southern Spain this species seems to have reached only
very recently the insular Spain: it is absent in the detailed investigation of GANGWERE & LLORENTE
(1992) on the ortopteroids of the Balearic, and the first record for this islands is dated 2004 on the
island of Mallorca (CANYELLES & ALOMAR, 2006).
Onychium, 13: 25-30
28
Presumably from here, in even more recent times, the specimens (or their ancestors) found in Spargi
in 2011 (RUZZANTE & LEO, 2012) and in the Sardinian hinterland in 2016 may have arrived. The
orthopteroid fauna of Spargi is quite well known after the entomological surveys of BACCETTI (1996),
where the species is not reported and it seems unlikely that, by its size and appearance, this species
was not collected. On the contrary, its arrival time in the Sardinian hinterland of Santa Giusta, less
known in its entomologic fauna, appears less evident.
In the meanwhile S. viridis also colonized some islands in the eastern part of its habitat: a specimen
was collected in Cyprus in 1977 (EHRMANN, 2011) and its presence in this island has been confirmed
later with a reproductive couple observed by the authors in 2011 (Battiston, unpublished data).
Known for the Tunisian hinterland, this species was found in the island of Djerba in 2005 (BATTISTON
et al., 2010) with another reproductive couple of individuals and some oothecae (Fig. 3). If the arrival
Fig. 3. Male individuals of S. viridis (Forskål, 1775): a) specimen from Santa Giusta, ph. R. Perra; b) specimen
from Cyprus ph. P. Zanolin; c) specimen from Djerba with an ootheca, ph. R. Battison.
Onychium, 13: 25-30
29
of this presence in Tunisian island just a few kilometres far from the inland, was easily conceivable,
less clear is the situation in Cyprus since its presence in neighbouring Turkey remained uncertain
until 1996 (EHRMANN, 2011). Its presence on the Greek islands is at the moment uncertain, mostly
because of the recent spreading in that area of Hierodula transcaucasica Brunner von Wattenwyl,
1878 (Battiston, unpublished data), a closely related mantis from the same Tribe but with an Asian
distribution, with whom can easily be confused. While it is clear that the insular distribution of this
species is generally very recent, the question of the naturalness of this trend remains open. The size
and wing length of this mantid (BATTISTON et al., 2010) makes it unsuitable for long-range flights.
The passive transport of the ootheca, in this species with a particularly thick protective cover (Fig.
3c), seems the most likely hypothesis, but the naturalness of this process, in relation to this
extraordinarily recent expansion, seems not plausible. The life cycle of this insects is annual and the
permanence of the larvae in the ootheca can be measured, depending on the latitude, from a few
weeks to a few months. The timing of a passive drift of marine larva from the Spanish coast to the
Balearic platform has been measured in about one month (OLIVER, 1991) and the shortest distance
from Mallorca to Sardinia is greater by about two times but the surface circulation of course follows
much longer routes (ROBINSON et al., 2001). This means several months or years of travel, which is
not compatible with the biology of this insect. On the contrary, in almost all locations where this
species has been observed in nature by the authors in recent years (Morocco, Tunisia, Sardinia,
Cyprus), it has always shown not to dislike well anthropized habitats, and was often found in city
parks or gardens, car parking lots and roadsides verges. Significantly different however is the case of
Spargi, a small island, uninhabited and with occasional, well-regulated tourist accesses. Even the
arrival of this species on the port area of Calvià, in the island of Mallorca from the near Spanish coast
was considered as a probable artificial introduction (CANYELLES & ALOMAR, 2006). The introduction
of alien species of mantids in Europe has already been reported for Portugal (MARABUTO, 2014), even
if related to Afrotropical and Middle East species as Miomantis caffra Saussure, 1871 and M.
paykullii Stål, 1871, completely foreign to Western Europe. In this case the arrival of S. viridis in the
Mediterranean islands by natural oothecae transportation can not be excluded (and the specimen of
Spargi may actually fall into this dynamic), although generally this situation is unlikely to appear.
More likely is the artificial transportation of oothecae laid on manufactured goods, timber or other
material transported quickly on the well frequented shipping routes of the Mediterranean. Even if the
morphological homogeneity of the two individuals collected supports the hypothesis of a wide spread
Sardinian population, we cannot exclude that they may have been also introduced directly and in
separate occasions, i. e. attracted by artificial lights on ferries or boats, as already hypothesized for
Lethocerus patruelis (Stål, 1855) (CIANFERONI & NARDI, 2013). At present, the presence of this
mantid in Italy is still a questionable case. However the collected evidences: the spread of this species
in the Mediterranean islands occurred in recent times, together with the increasing of marine vessel
traffic, and the well documented attraction of mantids for artificial light and surfaces, to lay the
oothecae (LOPEZ, 1998), tend to consider this a potential alien species for many Mediterranean
countries. Local and specific studies are however encouraged to check the individual cases and the
impact that this new guest can have on already delicate balance of the Mediterranean insular
environments.
Acknowledgements
We would like to thank Piero Leo for his help in dissecting and analyzing the specimen from Spargi, and Renzo
Perra and Paola Zanolin for sharing their beautiful photos of the living individuals. A special thank to Damiano
Andria who made possible the collecting of the specimen from Santa Giusta and to the two anonymous reviewers
that significantly improved the quality of this manuscript.
References
BACCETTI B., 1996. Gli Ortotteroidei delle piccole isole che circondano la Sardegna (Notulae Orthopterologicae.
53). Biogeographia, 18 (1995): 385-390.
BATTISTON R., PICCIAU L., FONTANA P. & MARSHALL J., 2010. Mantids of the Euro-Mediterranean Area. World
Biodiversity Association, Verona, 239 pp.
Onychium, 13: 25-30
30
BOLIVAR I., 1898. Catalogo sinóptico de los Ortópteros de la fauna ibérica. Anales de Ciencias Naturales, 4: 105-
232.
CANYELLES X. & ALOMAR G., 2006. Sobre la presència de Sphodromantis viridis (Forskål, 1775) (Dictyoptera,
Mantoidea) a Mallorca. Bolletí de la Societat d’Història Natural de les Balears, 49: 83-88.
CIANFERONI F. & NARDI G., 2013. Lethocerus patruelis (Stål, 1855) in Italy: a recent introduction or a natural
westward spread? (Hemiptera: Heteroptera: Nepomorpha: Belostomatidae). Zootaxa, 3664 (1): 78-84.
EHRMANN R., 2011. Mantodea from Turkey and Cyprus (Dictyoptera: Mantodea). Articulata, 26 (1): 1-42.
GANGWERE S.K. & LLORENTE V., 1992. Distribution and habits of the Orthoptera (sens. lat.) of the Balearic
Islands (Spain). Eos, 68 (1): 84.
LA GRECA M., 1966. Sulle Sphodromantis del gruppo viridis dell’Africa settentrionale ed occidentale. Eos, 42 (3-
4): 493-516.
LA GRECA M. & LOMBARDO F., 1987. Revisione delle Sphodromantis Ståhl, 1871 (Insecta Mantodea) dell’Africa
orientale. Monitore zoologico italiano, n. s., suppl., 22 (13): 193-234.
LOPEZ A.I., 1998. Un insecto de leyenda en Leon: Mantis religiosa. Universidad de León, León, 148 pp.
MARABUTO E., 2014. The Afrotropical Miomantis caffra Saussure 1871 and M. paykullii Stål 1871: first records of
alien mantid species in Portugal and Europe, with an updated checklist of Mantodea in Portugal (Insecta:
Mantodea). Biodiversity Data Journal, 2: e4117. doi:10.3897/BDJ.2.e4117
MARABUTO E., RODRIGUES I. & HENRIQUES S., 2014. Sphodromantis viridis (Forskål, 1775): New for Portugal
and new records of the rare and small mantids Apteromantis aptera (Fuente, 1894) and Perlamantis allibertii
Guérin-Méneville, 1843 in the country (Mantodea: Mantidae and Amorphoscelidae). Biodiversity Data
Journal, 2: e1037. doi:10.3897/BDJ.2.e1037
MARTINEZ-CORTIZAS A., PONTEVEDRA-POMBAL X., GARCIA-RODEJA E., NOVOA-MUÑOZ J.C. & SHOTYK W.,
1999. Mercury in a Spanish peat bog: archive of climate change and atmospheric metal deposition. Science,
284: 939-942.
OLIVER P., 1991. Dinámica de la población de la merluza (Merluccius merluccius, L.) de Mallorca
(Reclutamiento, Crecimiento y Mortalidad). Tesis doctoral. Universidad de las Illes Balears, 392 pp.
ROBINSON A.R., LESLIE W.G., THEOCHARIS A. & LASCARATOS A., 2001. Mediterranean sea circulation. Ocean
currents: a derivative of the Encyclopedia of Ocean Sciences, 1689-1705.
ROY R., 2010. Mises au point sur le genre Sphodromantis Ståhl, 1871 (Mantodea, Mantidae). Bulletin de la Société
entomologique de France, 115 (3): 345-366.
RUZZANTE G. & LEO P., 2012. 116. Sphodromantis viridis (Forskål, 1775) (Mantodea Mantidae). Quaderno di
Studi e Notizie di Storia Naturale della Romagna, 35: 173-174.
WERNER F., 1933. Neue Orthoptera im Naturhistorischen Museum Basel. Über einige von Herrn H. Hediger auf
Neuguinea gesammelten Mantodeen. Verhandlungen der Naturforschenden Gesellschaft in Basel, 43: 1-3.
Received 1 December 2016
Accepted 20 February 2017
© 2017 Battiston et al. This is an o pen access work distributed under the terms of the Creative Commons Attribution 4.0 International License (CC BY),
which permits unrestricted use, distribution, an d reproduction in any medium, provided the original author and source are credited. To view a copy of
this license, visit http://creativecommons.org/licenses/by/4.0/
