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MONTESANTO ET AL.: DISTRIBUTION OF TYLOS IN THE MEDITERRANEAN                   53

uate the degree of divergence between the various popula-                                , and P. J. Schembri. 2008. Assessing inter-beach differences in
tions of Tylos in Sicily and the circum-Sicilian islands. Such                     semi-terrestrial arthropod assemblages on Maltese pocket sandy beaches
studies will also highlight any incidence of genetic introgres-                    (Central Mediterranean). Marine Ecology 29(suppl.): 108-117.
sion within boundary areas separating current ranges of T.                      Dyer, K. S. 1979. Estuarine Hydrography and Sedimentation: a Handbook:
europaeus and T. ponticus. This will make necessary a sec-                         i-ix, 1-230. Cambridge University Press, Cambridge.
ond sampling campaign that would focus on the coastal ar-
eas separating the two sets of populations – such zones are                     Fallaci, M., I. Colombini, S. Taiti, and L. Chelazzi. 1996. Environmental
present in Sicily but are completely absent on the circum-                         factors influencing the surface activity and zonation of Tylos europaeus
Sicilian and the Maltese Islands.                                                  (Crustacea: Oniscidea) on a Tyrrhenian sandy beach. Marine Biology
                                                                                   125: 751-763.
   Biological interactions between the two species, namely
in the form of competitive exclusion, could also be operat-                     Giordani Soika, A. 1954. Ecologia, sistematica, biogeografia ed evoluzione
ing, especially since the niches of the two species partially                      del Tylos laterillei Auct. (Isop. Tylidae). Bollettino Museo Civico Storia
overlap. Because the current study concluded that median                           Naturale di Venezia 7: 63-83, pls. I-XI.
grain size cannot explain the allopatric separation of the                               . 1956. Sull’esistenza nelle coste adriatiche di una nuova razza del
two species, niche separation between the two targeted Ty-                         Tylos sardous (Arc.). Bollettino del Museo Civico di Storia Naturale di
los could be attributed to different variables, which might                        Venezia 9: 14-15, pls. 1-3.
include the water and organic content of the sand, the degree
of porosity and penetrability of the same beach sand, and the                   Goncalves, S. V., M. A. Pardal, P. G. Cardoso, S. M. Ferreira, and J. C.
exposure to wave action, and the slope of the beaches.                             Marques. 2005. Biology, population dynamics and secondary production
                                                                                   of Tylos europaeus (Isopoda, Tylidae) onthe western coast of Portugal.
                     ACKNOWLEDGEMENTS                                              Marine Biology 147: 631-641.

The authors thank Prof. Paula B. Araujo, Universidade Federal do Rio            Grebnicki, N. 1874. Materialy dlja fauny Novoressijskogo kraja. Isopoda.
Grande do Sul, Porto Alegre (Brasil), for critical comments on the first            Zapiski novorossi˘ıskago Obshchestva Estestvoispyˆtatele˘ı 2: 250-262,
versions of this paper. We wish to thank Prof. Domenico Caruso for                 pls. I-III. [In Russian].
providing us the opportunity to access the “Caruso Collection” of terrestrial
isopods, housed at the Museum of Zoology (University of Catania,                Hammer, W., M. Smyth, and E. Mulford. 1969. The behavior and life
Italy), and Prof. Richard Muscat from the University of Malta Research             history of a sand-beach isopod, Tylos punctatus. Ecology 50: 442-453.
Committee for funding such a study. The authors would also like to thank
two anonymous referees whose constructive comments greatly improved             Hayes, W. B. 1974. Sand-beach energetics: importance of the isopod Tylos
the value and rigour of the manuscript.                                            punctatus. Ecology 55: 838-847.
                                                                                         . 1977. Factors affecting the distribution of Tylos punctatus
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                                                                                AVAILABLE ONLINE: 13 December 2013.
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