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Diversity of small mammal communities of the Tuscan Archipelago: testing the effects of island size, distance from mainland and human density.

Abstract. Barn owl pellets were analyzed for comparing the small mammal fauna among three islands of Tuscan archipelago, Italy. Rarefaction analyses and diversity analyses were used on barn owl pellet samples at a total of 18 sites. Apodemus sylvaticus appeared the most abundant species in Elba and Capraia, and Mus musculus in Pianosa. Diversity profiles showed that Pianosa and Elba were very similar in terms of dominance and evenness profiles, whereas Capraia had higher dominance and lower evenness. Using our original data and literature review, we found that 5 species are found in Pianosa, 1 in Giannutri, 0 in Montecristo, 5 in Giglio, 7 in Elba, 6 in Capraia, and 1 in Gorgona. There was no effect of island area, human population size, or linear distance from mainland on species richness.

Key words: mammals, faunal composition, Mediterranean, owl pellets, Italy

Introduction

The knowledge of the small mammals of the Tuscan Archipelago (Tyrrhenian Sea, central Italy) is still fragmentary and anyway not available for all islands. Comparatively, the bulk of information comes from the island of Elba, such as Uttendorfer (1952) on the diets of the Griffon vulture and little owl, Vesmanis & Hutterer (1980), Vesmanis & Vesmanis (1980), and Contoli et al. (1988) on the diets of nocturnal bird of prey, up to Szpunar et al. (2008) on the same subject. For the other islands, only sporadic data are available, for a summary of sightings, see Angelici et al. (2009). The aim of this paper is to provide an updated overview of the knowledge of small mammals (Soricomorpha and Rodentia) in the islands of the Tuscan Archipelago (for two of these islands, i.e. Pianosa and Capraia, there were no data on mammals species before this study), and to use barn owl pellets for comparisons of the small mammal fauna among the islands of Tuscan Archipelago. More explicitly, we test whether there was any effect of (1) island size, (2) its distance from mainland, and (3) human population size, on species richness and diversity indices. These factors are certainly important in the light of (i) island biogeography theory (factors (1) and (2); see McArthur & Wilson 1963), and in the light of the deep human intervention that has characterized the islands of the Tuscan Archipelago, and the Mediterranean islands in general, since centuries (factor (3); see Masseti 2009a, b).

Material and Methods

The field study was carried out in three islands of the Tuscan Archipelago, that is a chain of islands between the Ligurian Sea and Tyrrhenian Sea, west of Tuscany, Italy (Fig. 1). The archipelago contains the islands of Gorgona, Capraia, Elba (the largest island of the group), Pianosa, Montecristo, Giglio, and Giannutri. All of these islands are protected as part of the Tuscan Archipelago National Park. Our field data were collected at several roosts of Elba, Capraia, and Pianosa, whereas surveys at Giglio did not provide any data because no barn owl roost was detected. More specifically, barn owl pellets were collected at one site of Pianosa, one site of Capraia and three sites of Elba islands during multiple samples (data collected by one of us, RP) (see Table 1).

[FIGURE 1 OMITTED]

For the analysis of the pellets, we used the dry method: pellets were opened along the lines of fracture and skulls (or their fragments) and jaws were taken. The determination of the bone material was carried out through observation of the material to the stereobinocular microscope, using the keys proposed by Chaline et al. (1974), Toschi (1965) and Amori et al. (2008).

For each species of small mammals, separate counts were taken for right and left maxillary and mandibular hemiarchs. For enumeration of each species, we considered only the largest group (or mandibles or jaws), and we used as the final value the highest number between the two sides.

In order to make a comprehensive checklist of small mammal composition for each island, we complemented the original data surveys with available literature.

We used rarefaction analysis to compare all diversity indices to the same number of individuals sampled, and based on this analysis, it was clear that a plateau phase was reached as for the whole archipelago, despite the samples of the various single sites were not equally reliable (Fig. 2A, B). The following indices of prey species diversity were calculated: 1) species richness (Taxa S), that is the total number of taxa recorded during either the wet or the dry season; 2) evenness, calculated by Pielou formula; 3) Simpson index; 4) Shannon index; and 5) Chao 1 bias corrected index, and 6) Dominance index (see Magurran 1988 for the formulas). Statistical differences among islands in terms of diversity indices were calculated using bootstrapping procedures. More precisely, upper and lower confidence limits of the estimates for the above-mentioned indices were calculated by 10000 bootstraps, with all random samples being produced with the same total number of individuals as in each original sample. The random samples were taken from the total, pooled data set (i.e. with dry and wet season counts cumulated). For each individual in the random sample, the taxon was chosen with probabilities according to the original abundances calculated on the pooled data set. A 95 % confidence interval was then calculated. Comparisons between frequencies of occurrence of the various prey species from original field surveys and literature were made by contingency

[FIGURE 2 OMITTED]

table [chi square] test. A General Linear Model (GLM) was designed to assess the effects of island area ([km.sup.2]), human population size (number of inhabitants as in 2012), and linear distance from the mainland (km) on the species richness per island. The intercept term was included in the model. The GLM was designed assuming that the overall sample was accurate (as showed by the curve plateau in Fig. 2B).

All statistical analyses, including rarefaction analyses, were performed with a PAST statistical software, apart GLM which was performed wit PASW 18.0 statistical software. In all cases, tests being two-tailed and alpha set at 5 %.

Results

Original barn owl pellet data

The synoptic survey of collected data is given in Table 1. The relative percent frequency of individual species across surveyed islands is given in Fig. 3. The frequencies of occurrence of the various species differed significantly across islands ([chi square] = 132.1, P < 0.005). Apodemus sylvaticus was by far the most abundant species in Elba and Capraia, whereas Mus musculus dominated the sample in Pianosa (Fig. 3). A dendrogram analysis revealed that three clusters of species are found (Fig. 4). Diversity profiles showed that Pianosa and Elba were very similar in terms of dominance and evenness confidence intervals profiles, whereas Capraia, despite having the same species richness as Elba, showed a significantly higher dominance (P < 0.05) and a significantly lower evenness (P < 0.01) when comparing the estimates (Table 2).

[FIGURE 3 OMITTED]

[FIGURE 4 OMITTED]

[FIGURE 5 OMITTED]

Comparisons between original and published datasets A quantitative comparison between our original data and literature data can be done only for Elba, using Contoli et al. (1988). Overall, the relative frequencies of occurrence of the various species did not differ significantly between Contoli et al. (1988) and our samples ([chi square] = 2.54, df = 6, P = 0.864). Nonetheless, one species was recorded only in our samples (S. etruscus) and one only in Contoli et al. (1988) samples (R. norvegicus) (Fig. 5).

Comparisons of faunistic assemblages across islands Updated information on the presence/absence of small mammals in all islands of the archipelago revealed that five species are found in Pianosa (i.e. S. etruscus, A. sylvaticus, R. rattus, R. norvegicus, M. musculus), one in Giannutri (R. rattus), no in Montecristo (after the recent eradication of R. rattus, finished in 2012), five in Giglio (C. suaveolens, A. sylvaticus, R. rattus, R. norvegicus, M. musculus), seven in Elba (C. suaveolens, S. etruscus, G. glis, A. sylvaticus, R. rattus, R. norvegicus, M. musculus), six in Capraia (S. etruscus, C. suaveolens, G. glis, A. sylvaticus, R. rattus, M. musculus), and one in Gorgona (R. rattus). A GLM design showed that there was no effect of either island area, human population size, or linear distance from the coast on the species richness per island (Table 3).

Discussion

Our study provided quantitative information on the small mammal communities of three islands of the Tuscan Archipelago, whereas it was not possible to obtain original data for the islands Gorgona, Giannutri and Giglio. It is suggested that it would be appropriate to carry out additional well-focused field research in Giglio because of its relatively large surface (24 [km.sup.2]). A surprising result of our own field research was in that Capraia, despite its small size (19 [km.sup.2]), showed a community of small mammals (six species) similar to that of Elba (224 [km.sup.2], seven species). Equally surprising was the discovery of dormouse (Glis glis) among the prey of barn owl in Capraia. We suggest that both the above-mentioned findings deserve further confirmation through direct field sightings on the local presence of these species (and particularly G. glis). Only dormouse specimen found in the present study may have been preyed upon by a barn owl in Elba island or in Corsica (respectively located at 33 and 27 km distance from Capraia), the same being true of course also for some of the other species. Indeed, it has been demonstrated that the barn owl can make long distance to go to catch prey (Bunn et al. 1982, Guerra et al. 2014), and the distances between the islands of the Tuscan Archipelago (range 13-75 km) are possibly compatible with this hypothesis. Accidental (passive) human introduction can be also another explanation for our record of G. glis from Capraia. In addition, the high dominance of commensal species (Mus musculus and Rattus rattus, overall accounting for 86 % of the total sample) could be explained by the fact that these samples were collected around a cheese factory. However, the small size of the island might mean that all owls in this island may have access to every habitat present in the island.

Interestingly, dominance index was very high in Capraia, which also showed a low evenness of barn owl dietary niche as compared to the other studied islands. It is likely that the high dominance index value for Capraia depended on the fact that the small mammal community is characterized by a preponderance of Apodemus sylvaticus in the maquis habitats of this island. Unfortunately, however, no field trapping surveys of small mammals are available to date to confirm this hypothesis.

The non-effects on species richness of island area, human population size, and linear distance from mainland could be explained by the history of human colonization and agricultural practices of the various islands (Sara 1998). Therefore, the eventual biological effects predictable on the basis of island colonization theories (e.g. McArthur & Wilson 1963, Simberloff 1976, Ricklefs 1982) could not be confirmed in the case under study. Interestingly, previous studies on birds (Massa 1985), amphibians and reptiles (Massa & Di Palma 1988, Parlanti et al. 1988), and mammals (Sara 1998) also showed that MacArthur & Wilson's (1963) theory fails in continental islands in terms of species-area relationships. Alternatively, it is also possible that the observed insular patterns were shaped by the hypothesis of Sara & Morand (2002), who suggested that mammals with a high density compared to the density/mass relationship are the best invaders and/or probably have less chance of going extinct on small islands when population size is small.

Acknowledgements

This study was funded by the Tuscan Archipelago National Park. We are indebted with Dr. Giuliano Milana for drafting figures and useful discussion on barn owl habits, and to two anonymous referees for helpful comments on the submitted draft.

Literature

Amori G., Contoli L. & Nappi A. 2008: Erinaceomorpha, Soricomorpha, Lagomorpha, Rodentia. Fauna d'Italia Mammalia II, vol. XLIV. Edizioni Calderini de Il Sole 24 Ore, Milano.

Angelici F.M., Laurenti A. & Nappi A. 2009: A checklist of the mammals of small Italian islands. Hystrix 20: 3-27.

Bunn D.S., Warburton A.W. & Wilson R.D.S. 1982: The barn owl. Poyser, Calton.

Chaline J., Baudvin H., Jammot D. & Saint Girons M.-C. 1974: Les proies des Rapaces. Doin, Paris.

Contoli L., Aloise G. & Filippucci M.G. 1988: Sulla nicchia trofica di Tyto alba ed Athene noctua nell'isola d'Elba (Toscana) in rapporto all'Italia Tirrenica. B. Ecol. 19: 367-373.

Guerra C., Garcia D. & Alcover J.A. 2014: Unusual foraging patterns of the barn owl, Tyto alba (Strigiformes: Tytonidae), on small islets from the Pityusic archipelago (Western Mediterranean Sea). Folia Zool. 63: 180-187.

Magurran A.E. 1988: Ecological diversity and its measurement. Princeton University Press, Princeton, NJ, U.S.A.

Massa B. 1985: Considerazioni sui popolamenti di uccelli terrestri delle isole mediterranee. Biogeographia 11: 261-284.

Massa B. & Di Palma G. 1988: Rettili, anfibi e uccelli terrestri delle isole circum-siciliane. B. Ecol. 19: 225-234.

Masseti M. 2009a: Mammals of the Mediterranean islands: homogenisation and the loss of biodiversity. Mammalia 73: 169-202.

Masseti M. 2009b: The wild goats Capra aegagrus Erxleben, 1777 of the Mediterranean Sea and the Eastern Atlantic Ocean islands. Mammal Rev. 39: 141-157.

McArthur R.H. & Wilson E.O. 1963: An equilibrium theory of insular zoogeography. Evolution 17: 373-387.

Parlanti C., Lanza B., Poggesi M. & Sbordoni V. 1988: Anfibi e rettili delle Isole del Mediterraneo: un test dell'ipotesi dell'equilibrio insulare. B. Ecol. 19: 335-348.

Ricklefs R.E. 1982: Ecology, 2nd ed. Chiron Press, New York.

Sara M. 1998: I mammiferi delle isole del Mediterraneo. L'Epos, Palermo.

Sara M. & Morand S. 2002: Island incidence and mainland population density: mammals from Mediterranean islands. Divers. Distrib. 8: 1-9.

Simberloff D. 1976: Species turnover and equilibrium island biogeography. Science 194: 572-578.

Szpunar G., Aloise G. & Filippucci M.G. 2008: Suncus etruscus (Soricomorpha, Soricidae): a new species for Elba Island (Tuscan Archipelago, Italy). Ital. J. Zool. 75: 445-447.

Toschi A. 1965: Lagomorpha, Rodentia, Carnivora, Ungulata, Cetacea. Fauna d'Italia. Mammalia, Vol. VII. Bologna, Calderini.

Uttendorfer O. 1952: Neue Ergebnisse uber die Ernahrung der Greifvogel und Eulen. Eugen Ulmer, Stuttgart.

Vesmanis I.E. & Hutterer R. 1980: Nachweise von Erinaceus, Crocidura und Microtus fur die Insel Elba, Italien. Z. Saugetierkd. 45: 251-253.

Vesmanis I.E. & Vesmanis A. 1980: Ein Nachweis des Siebenschlafers, Glis glis (Linnaeus, 1766) aus Eulengewollen von der Insel Elba, Italien. Faunistische Abhandlungen 7: 167-170.

Giovanni AMORI (1*), Vincenzo RIZZO PINNA (2), Giampiero SAMMURI (3) and Luca LUISELLI (4,5)

(1) CNR Institute of Ecosystem Studies, Viale dell'Universita 32, 00181 Rome, Italy; e-mail: giovanni.amori@uniroma1.it

(2) Systema Naturae, 5800 Grosseto, Italy

(3) Federparchi, via Nazionale 230, 00184 Rome, Italy

(4) Niger Delta Ecology and Biodiversity Conservation Unit, Department of Applied and Environmental Biology, Rivers State University of Science and Technology, P.M.B. 5080, Port Harcourt, Nigeria

(5) Centre of Environmental Studies DEMETRA, via Olona 7, 00198 Rome, Italy

Received 30 March 2015; Accepted 30 May 2015

(*) Corresponding Author
Table 1. Synoptic table summarizing the original data collected in the
present study.

Island      Precise locality of owl roosts   Dates         S. etruscus

PIANOSA     Cheese factory                   11/07/2013         9
PIANOSA     Cheese factory                   27/04/2013         8
PIANOSA     Prison (lateral building)        18/10/2013
PIANOSA     Prison (Judge's palace)          18/10/2013
PIANOSA     Prison (gymnasium)               17/10/2013
Total                                                          17
ELBA        La Zanca                         28/04/2013
ELBA        Marciana                         28/04/2013
ELBA        Mortigliano                      27/04/2013
ELBA        Mortigliano                      08/07/2013         1
ELBA        Rio nell'Elba                    09/07/2013
ELBA        Poggio cemetery                  09/07/2013
ELBA        Mulinaccio                       08/07/2013
ELBA        Calcinaio S. Stefano             12/12/2013         1
ELBA        Cala Seregola (mines area)       12/12/2013
ELBA        Cala Seregola (mines area)       Sept. 2013         1
Total                                                           3
CAPRAIA     Abandoned Penal Settlement       04/12/2013
CAPRAIA     Cemetery                         19/11/2013
CAPRAIA     Abandoned Penal Settlement       04/12/2013         2
Total                                                           2

Island      C. suaveolens   G. glis      A. sylvaticus     M. musculus

PIANOSA                                                       18
PIANOSA                                                       63
PIANOSA                                        2               1
PIANOSA                                        1
PIANOSA                                                        8
Total           0             0                3              90
ELBA                                           3
ELBA                                          13
ELBA                                          32
ELBA            2                             40               3
ELBA            1                             34              23
ELBA            2             1              180
ELBA            1                             53              15
ELBA            3                             55               2
ELBA            3                             48
ELBA            1                             31
Total          13             1              489              43
CAPRAIA         3                             32              14
CAPRAIA         3             1              458
CAPRAIA         2                             25              12
Total           8             1              515              26

Island        R. rattus

PIANOSA          4
PIANOSA         18
PIANOSA          3
PIANOSA          2
PIANOSA          2
Total           29
ELBA             4
ELBA
ELBA            20
ELBA             9
ELBA            18
ELBA            46
ELBA            43
ELBA            57
ELBA            31
ELBA            22
Total          250
CAPRAIA         14
CAPRAIA         51
CAPRAIA         22
Total           87

Table 2. Diversity indices (including upper and lower 95 % confidence
intervals) for the three studied islands, based on the frequency of
occurrences of the small mammal species in the barn owl pellets.

                  Pianosa                          Elba
                  estimate    lower     upper      estimate

Taxa_S            4           4         5          6
Dominance_D       0.4775      0.3062    0.4164     0.4763
Simpson_1-D       0.5225      0.5832    0.6934     0.5237
Shannon_H         0.95        1.041     1.29       0.9162
Evenness_e^H/S    0.6465      0.6014    0.8633     0.4166
Chao-1            4           4         5          6

                                        Capraia
                  lower       upper     estimate   lower     upper

Taxa_S            4           7         6          4         7
Dominance_D       0.3046      0.4118    0.6684     0.306     0.4109
Simpson_1-D       0.5868      0.6952    0.3316     0.5883    0.6936
Shannon_H         1.038       1.293     0.6632     1.041     1.297
Evenness_e^H/S    0.6044      0.8643    0.3235     0.6018    0.8636
Chao-1            4           5         6          4         6

Table 3. Results of the General Linear Model on the effects of
human population size, island size ([km.sup.2]), and distance (km) from
the mainland on the species richness of small mammals in the various
island of the Tuscan Archipelago. Species richness was the dependent
variable.

Source                        Type III Sum of Squares

Corrected Model                     23.469 (a)
Intercept                           13.493
Human population                     9.312
Distance from mainland               7.474
Island area ([km.sup.2])            10.201
Error                               17.388
Total                              161.000
Corrected Total                     40.857

Source                       df  Mean Square      F         P

Corrected Model              3      7.823       1.350     0.406
Intercept                    1     13.493       2.328     0.224
Human population             1      9.312       1.607     0.294
Distance from mainland       1      7.474       1.290     0.339
Island area ([km.sup.2])     1     10.201       1.760     0.277
Error                        3      5.796
Total                        7
Corrected Total              6

(a) [R.sup.2] = 0.574 (Adjusted [R.sup.2] = 0.149).
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Author:Amori, Giovanni; Pinna, Vincenzo Rizzo; Sammuri, Giampiero; Luiselli, Luca
Publication:Folia Zoologica
Article Type:Report
Geographic Code:4EUIT
Date:Aug 1, 2015
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