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Distribution of natural populations of the killifishAphanius fasciatus (Valenciennes, 1821) (Teleostei:Cyprinodontidae) in Italy: past and current status, andfuture trendsS. Valdesalicia, J. Langeneckb, M. Barbierib, A. Castellib & F. Maltagliatiba Associazione Italiana Killifish, Viano, Italyb Dipartimento di Biologia, Università di Pisa, Pisa, ItalyPublished online: 28 Jan 2015.

To cite this article: S. Valdesalici, J. Langeneck, M. Barbieri, A. Castelli & F. Maltagliati (2015): Distribution of naturalpopulations of the killifish Aphanius fasciatus (Valenciennes, 1821) (Teleostei: Cyprinodontidae) in Italy: past and currentstatus, and future trends, Italian Journal of Zoology, DOI: 10.1080/11250003.2014.1003418

To link to this article: http://dx.doi.org/10.1080/11250003.2014.1003418

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Distribution of natural populations of the killifish Aphanius fasciatus(Valenciennes, 1821) (Teleostei: Cyprinodontidae) in Italy: past andcurrent status, and future trends

S. VALDESALICI1, J. LANGENECK2*, M. BARBIERI2, A. CASTELLI2, & F. MALTAGLIATI2

1Associazione Italiana Killifish, Viano, Italy, and 2Dipartimento di Biologia, Università di Pisa, Pisa, Italy

(Received 14 May 2014; accepted 18 December 2014)

AbstractThe census of natural populations provides relevant information for both conservation purposes and environmentalmonitoring. In the present work, natural populations of the killifish Aphanius fasciatus (Teleostei: Cyprinodontidae)were censused in the Italian territory. The historical distribution of the species was reconstructed with the aid ofbibliographic material and museal samples, in order to evaluate the extent of changes in its distribution. Three periods(pre-1990s, 1990s, post-1990s) and five habitat types (lagoons/coastal ponds, saltworks, canals, river mouths, inland waterbodies) were considered. Overall, we recorded the presence of A. fasciatus in 82 localities, the majority of which arelagoons/ponds. The species showed a widespread decline and sometimes local population extinction, due to habitatdegradation and competition with the introduced poeciliid Gambusia holbrooki. With a few exceptions, A. fasciatus iscurrently extinct in inland waters, being mainly present in coastal brackish-water habitats. The occurrence of stablepopulations along the central Tyrrhenian, northern Adriatic, Sardinian and Apulian coastal biotopes is confirmed. Thepopulations of Sicilian inland waters, considered extinct, showed recovery capabilities. In northern Tuscany, where thespecies was also considered extinct, there are also recent records. From a temporal perspective, the present work showsthat A. fasciatus experienced population reductions throughout its range in the 1990s, when the species appeared locallyextinct, or strongly rarefied, in several localities. Since then, the species recovered in the majority of the native range.However, careful monitoring is needed in order to control the effects of habitat alterations and competition with G.holbrooki on natural populations of A. fasciatus. Given the importance of brackish-water habitats as both biodiversityhotspots and buffer areas against rainfalls and floods, A. fasciatus may be raised to the status of “umbrella species” withregard to these habitats.

Keywords: Aphanius fasciatus, population census, brackish-water habitats, conservation, Italy

Introduction

The Mediterranean killifish, Aphanius fasciatus(Valenciennes, 1821), is a cyprinodontid fish whosenative range includes coastal brackish-water biotopesaround the whole Mediterranean basin, with the excep-tion of the westernmost and easternmost parts, where itis replaced by the congeneric Aphanius iberus(Valenciennes, 1846) and Aphanius dispar (Rüppell,1829), respectively. Interestingly, in some northEgyptian lagoons, A. fasciatus can be found in sympatrywith A. dispar, with which it can produce hybrid off-spring (Villwock 1985, 1987; Lotan&BenTuvia 1996).Recently, a closely related species, Aphanius almiriensis(Kottelat, Barbieri & Stormboudi, 2007), has been

described from two sites in the Peloponnese (Greece;Kottelat et al. 2007).A. fasciatus inhabits shallow brack-ish-water environments where it is usually found inlarge-sized populations; however, it has also beenobserved in inland waters, such as Sicilian rivers(Ferrito & Tigano 1996) or north African oases(Kraiem 1983). A. fasciatusmay tolerate harsh environ-mental conditions, including high temperature andsalinity and low oxygen concentration. Despite its resis-tance towardswide variations of chemical-physical para-meters, a number of biological traits give vulnerability tothe species. For instance, the within-population compo-nent of genetic variation is much smaller than theamong-population counterpart (Maltagliati 1999) and

*Correspondence: J. Langeneck, Dipartimento di Biologia, Università di Pisa, via Derna 1, 56126 Pisa, Italy. Tel: +39 050 2211447. Fax: +39 050 2211410.Email: jlangeneck@biologia.unipi.it

Italian Journal of Zoology, 2015, 1–12http://dx.doi.org/10.1080/11250003.2014.1003418

© 2015 Unione Zoologica Italiana

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this feature decreases the evolutionary potential of nat-ural populations (Toro & Caballero 2005). At thisregard, several genetic studies revealed that A. fasciatuspopulations are highly divergent, mainly because ofthe virtually complete isolation of brackish-waterhabitats and the highly selective environmental features(Comparini et al. 1984;Maltagliati 1998a, 1998b, 1999;Cimmaruta et al. 2003; Maltagliati et al. 2003; Tiganoet al. 2006; Ferrito et al. 2007, 2013; Rocco et al. 2007;Triantafyllidis et al. 2007).

Populations of A. fasciatus are interesting models forthe study of morphological polymorphism (Kiener &Schachter 1974; Tigano & Ferrito 1984), genetic diver-sity (Maltagliati 1998a, 1998b, 1999; Ferrito et al. 2007;Triantafyllidis et al. 2007; Pappalardo et al. 2008), rela-tionships between phenotype and genotype (Ferritoet al. 2003; Maltagliati et al. 2003), macroevolutionaryprocesses (Villwock 1982; Hrbek & Meyer 2003),evolution of physiological adaptations (Crivellatoet al. 2006), reproductive behaviour and sexual selection(Malavasi et al. 2010; Cavraro et al. 2013a, 2013b) andcan be also employed as effective “sentinels” to detectgenotoxic pollutants in coastal lagoon ecosystems(Mosesso et al. 2012).

In recent years, A. fasciatus assumed relevance forconservation due to the documented decline and, insome instances, even extinction, of many naturalpopulations. Causes should be found in alterationsand deterioration of natural biotopes and habitat lossdue to the increasing human activities in coastalzones. In addition, the competition with the alienspecies Gambusia holbrooki Girard, 1859, a poeciliidfish introduced in southern Europe with the aim tocontrol the proliferation of malaria-carrying mosqui-toes, caused a strong reduction of A. fasciatus range.Therefore, the species has been included in lists ofprotected species, such as those of the BerneConvention (Bern/Berne, 19.IX.1979) and FaunaFlora Habitat European Directive (ASPIM 92/43/CEE).

The present study is aimed at providing a censusof natural populations of A. fasciatus in the Italianterritory. Moreover, the current picture will be com-pared to the available historical data, in order (1) toassess the conservation status of this species in Italy,(2) to make an attempt to reconstruct temporalchanges of its distribution and (3) to make predic-tions of future trends and possible interventions forconservation purposes.

Materials and methods

The census of natural populations of A. fasciatus inItalian territory was carried out by direct surveys ofsuitable environments. Samplings were carried out by

means of hand nets and minnow traps and specimenswere released in compliance with national andEuropean laws. Additional data, about both the pre-sent and the past distribution of A. fasciatus, were alsoobtained through scientific literature and personalcommunications with biologists and naturalists.Moreover, historical data were also obtained frommuseum collections, in particular from the MuséeNational d’Histoire Naturelle (MNHN, Paris),Natural History Museum of Milan (MSNM, Milan),Natural History Museum of Venice (MSNVE,Venice), National Museum of Natural History(USNM, Washington, DC), NaturhistorischesMuseum (NHM, Wien) and Museo Nacional deCiencias Naturales (MNCN, Madrid). We checkedthe collection databases of many other natural historymuseums in Italy and Europe, but we did not findrecords of A. fasciatus from the Italian territory.In the distributional map of A. fasciatus given by

Bianco (1993) for Italy, five discrete macroareas canbe identified, namely: (1) northern and centralTyrrhenian Sea, (2) western Sardinia, (3) southernSicily, (4) southern Adriatic Sea (Apulia) and (5)northern Adriatic Sea. We used this map as a start-ing point, in order to compare Bianco’s (1993) the-oretical distribution (Figure 1A) with data gatheredin the present work. In particular, the evolution ofA. fasciatus distribution was partitioned into the pre-1990s, 1990s and post-1990s, in order to test thehypothesis of range reduction in the 1990s withsuccessive possible recovery. Censused localitieswere categorised into the following habitat types:(1) lagoons/coastal ponds, (2) saltworks, (3) canals,(4) river mouths and (5) inland water bodies, takingthe cue from Carrada and Fresi (1988) andTagliapietra et al. (2009), in order to examinealso the distribution in ecologically differentenvironments.

Results

The total number of censused localities was 82(Table I). Stable populations of A. fasciatus aremainly found in brackish-water environments, suchas lagoons, saltworks, coastal lakes, ponds andcanals. No recent records were reported fromCampania and Liguria (in the latter region the spe-cies has been reported only on the basis of specimenscollected offshore; Torchio 1967), and no recordswere reported from Marche (Bianco 1993),Abruzzo (Bianco 1997), Molise, Basilicata orCalabria. Distributional maps regarding the pre-1990s, 1990s and post-1990s situations have beenproduced (Figure 1B–D).

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The largest part (53) of the 82 censused localitiesbelong to the “lagoon/coastal pond” category,whereas “inland water bodies”, “saltworks”, “canals”and “river mouths” are represented by 10, eight,seven and four sites, respectively. From a temporalperspective, in the pre-1990s, 1990s and post-1990speriods, A. fasciatus is reported from 26, 35 and 66localities, respectively. Within each period, the mostrepresented habitat type is the “lagoon/coastal pond”.Habitat types do not show changes in their relativeabundance, with the exception of the “inland waterbody”. In fact, this habitat shows a reduction from sixsites in the pre-1990s, to three and four in the 1990sand post-1990s, respectively.

Discussion

Past and current distribution of A. fasciatus in the Italianterritory

A detailed census of historically known populationsof A. fasciatus in the Italian territory allowed us todetermine the current species’ distribution and makean attempt to evaluate whether it was subject topossible range contraction. The pre-1990s distribu-tional map (Figure 1B) can be assumed to be theoldest data on the species’ historical range; along thecoast of the peninsula, this map identifies the widestknown range for the species, which seems to havebeen reduced and partially recovered in the last three

Figure 1. Past and current distribution of Aphanius fasciatus in the Italian territory. A, reference map (from Bianco 1993, modified); B, pre-1990s; C, 1990s; D, post-1990s distributions reconstructed on the basis of data reported in Table I. Blank maps were obtained from sitehttp://d-maps.com.

Census of Aphanius fasciatus in Italy 3

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Table I. List of censused localities along the Italian coastlines, with geographical coordinates, year and references. Legend: abs. = the censusgave a negative result; PC = personal communication; PO = personal observation; habitat types: L = lagoon/coastal pond; S = saltworks;C = canals; M = river mouth; I = inland water body. MNCN: Museo Nacional de Ciencias Naturales (Madrid, Spain); MNHN: MuséeNational d’Histoire Naturelle (Paris, France); MSNM: Natural History Museum of Milan (Milan, Italy); MSNVE: Natural HistoryMuseum of Venice (Venice, Italy); NHM: Naturhistorisches Museum (Wien, Austria); USNM: National Museum of Natural History(Washington, DC, USA).

Locality Lat.; Long. Habitattype

Year (reference)

Isonzo River mouth 45°47ʹN;13°27ʹE

M 1996 (Tempesta & Zuppa 2000); 2001 (Castellarin PC)

Grado Lagoon 45°41ʹN;13°24ʹE

L 1961 (Cavicchioli 1962); 1982 (Tigano & Ferrito 1984; Ferrito et al. 2007); 2006 (Ferritoet al. 2013)

Marano Lagoon 45°41ʹN;13°04ʹE

L 1950 (Bandella PC); 1982–2006 (Comisso PC); 2007 (Valdesalici PO; Comisso PC)

Treviso town 45°40ʹN;12°12ʹE

I 1860 (Ninni 1863)

Venice lagoon 45°26 N;12°17ʹE

L 1822–1827 (Nardo 1827); 1858 (Bellotti 1858); 1860 (Ninni 1863); 1888 (MNHN); 1899(MNHN); 1947; 1956 (MSNVE); 1961 (Cavicchioli 1962); 1970–1980 (Cappon PC);1981 (Marconato 1982); 1982 (Comparini et al. 1984); 2000 (Cappon PC); 2002(Franco et al. 2009; Cappon PC); 2002–2011 (Cavraro et al. 2011); 2004 (Cappon PC);2007 (Malavasi et al. 2010); 2010 (Cavraro et al. 2013b; MSNVE); 2012 (Cavraroet al. 2013a)

Chioggia 45°13ʹN;12°14ʹE

L 1943–46 (Peloso 1946); 1970 (Cellere PC); 1981 (Comparini et al. 1984)

San Michele diRavenna

44°24ʹN;12°06ʹE

I 1965–1970 (Magnani PC); 2012 (abs. Magnani PC)

Marina Romea,Marina di Ravenna

44°31ʹN;12°15ʹE

C 1999 (Valdesalici PO); 2000 (Cuni PC); 2004 (Eberl PC)

Francolino 44°54ʹN;11°40ʹE

I 1984 (Parisi PC); 2004 (abs. Valdesalici PO)

Saline di Cervia 44°15ʹN;12°22ʹE

S 1999 (Valdesalici PO); 2000 (Cazzulani PC); 2003 (Montanari PC); 2006 (Mordentiet al. 2007); 2007 (Mordenti et al. 2008; Valdesalici PO); 2010 (Mordenti et al. 2012)

Comacchio lagoon 44°41ʹN;12°11ʹE

L 1962 (Cavicchioli 1962); 1970s (Kiener & Schachter 1974); 1994; 1996 (Maltagliati PO);2004 (Montanari PC)

Lido di Spina 44°07ʹN;12°15ʹE

C 2006 (Kärst PC.)

Finale Ligure 44°09ʹN;08°20ʹE

- 1966 (Torchio 1967)

Calambrone 43°36ʹN;10°18ʹE

C 1962 (Cavicchioli 1962); 2013 (Langeneck PO)

Tombolo 43°38ʹN;10°19ʹE

C 1962 (Cavicchioli 1962)

Piombino 42°57ʹN;10°29ʹE

C 1994; 1996 (Maltagliati 1998a, 1999)

Elba 42°48ʹN;10°19ʹE

L 1983 (Castelli PO); 1994–1995 (Maltagliati 1998a, 1998b, 1999); 1996–1997 (Maltagliati& Camilli 2000); 2013 (Langeneck PO)

42°45ʹN;10°23ʹE

L 1983 (Castelli PO); 2013 (Langeneck PO)

Diaccia 42°44ʹN;10°57ʹE

L 1997–1998 (Cognetti et al. 1998); 2000 (Maltagliati 2002)

Ombrone river mouth 42°40ʹN;11°00ʹE

M 2011 (Fastelli et al. 2012)

Orbetello lagoon 42°26ʹN;11°12ʹE

L 1957 (MSNM); 1962 (Cavicchioli 1962); 1960s (USNM); 1980 (Varisco PC); 1982(NHM) 1994; 1996 (Maltagliati 1998a, 1998b, 1999); 1999–2000 (Mancini PC); 2003(Thorup PC); 2004 (Farmer PC; Valdesalici PO; Cellerino PC); 2005 (Valdesalici PO);2011 (Mosesso et al. 2012); 2012 (Langeneck PO)

Caprolace lake 41°21ʹN;12°59ʹE

L 2001 (Mancini PC; Valle PC); 2002 (Montanari PC); 2007–2008 (Manzo 2010)

Sabaudia lake 41°18ʹN;13°02ʹE

L 1994: 1996 (Maltagliati 1998a, 1998b, 1999); 2001 (Mancini 2001)

Tarquinia salterns 42°12ʹN;11°42ʹE

S 1980s (Mastrolia & Gallo 1989); 1998; 2003; 2007 (Cimmaruta et al. 2003; Angelettiet al. 2010; Ferrito et al. 2013); 2011 (Mosesso et al. 2012)

(Continued )

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Table I. (Continued).

Locality Lat.; Long. Habitattype

Year (reference)

Fusaro Lake 40°50ʹN;14°04ʹE

L 1861 (MNHN); 1962 (Cavicchioli 1962)

Mar Piccolo 40°29ʹN;17°18ʹE

L 1800 (Costa 1850); 1962 (Cavicchioli 1962); 2010–2013 (Baldacconi PC)

Pond near S.Isidoro 40°16ʹN;17°53ʹE

L 1994 (Bianchi PC); 2001 (Salvagiani PC); 2002 (Denitto PC)

Margherita di Savoiasaltwork

41°23ʹN;16°08ʹE

L-S 1800 (Costa 1850); 2000 (Bufo PC)

Lesina lake 41°53ʹN;15°22ʹE

L 1982 (Tigano & Ferrito 1984); 1994; 1996 (Maltagliati 1999); 1999 (Ferrito et al. 2007);2001 (Ferrito et al. 2003); 2006 (Ferrito et al. 2013; Cepollaro PC); 2006–2007(Manzo 2010); 2008 (Valdesalici PO)

Varano lake 41°51ʹN;15°46ʹE

L 1800 (Costa 1850); 2002 (Piccolo PC); 2004 (Kärst PC); 2008 (Valdesalici PO)

Acquatina pond 40°25ʹN;18°15ʹE

L 2002 (Denitto PC)

Cesine 40°23ʹN;18°18ʹE

L 2002 (Denitto PC)

Pilo pond 40°51ʹN;08°17ʹE

L 2000 (Maltagliati 2002; Ferrito et al. 2007); 2001 (Ferrito et al. 2003; Kärst PC); 2007(Ferrito et al. 2013); 2011 (Ruozi & Zubani PC); 2012–2013 (Morreale PC)

Mistras 39°55ʹN;08°28ʹE

L 2000 (Maltagliati 2002)

Casaraccio pond 40°54ʹN;08°15ʹE

L 1994; 1996 (Maltagliati 1998a, 1998b, 1999); 2007 (Ferrito et al. 2013)

Stintino saltworks 40°54ʹN;08°14ʹ E

S 2013 (Barbieri PO)

Calich pond 40°54ʹ N;08°13ʹE

L 1982 (Comparini et al. 1984)

Capo Comino 40°30ʹN;09°48ʹE

L 2002 (Parisi PC)

Baia del Sole 40°55ʹN;09°31ʹE

L 2000 (Maltagliati 2002); 2002 (Wischmann PC)

San Teodoro pond 40°47ʹN;09°40ʹE

L 1995; 2002 (Wischmann PC); 2004 (Lo Stimolo PC)

Budoni 40°09ʹN;08°58ʹE

L 1995; 2002 (Wischmann PC)

Girasole 39°57ʹN;09°40ʹE

L 1997; 2002 (Wischmann PC)

Santa Giusta pond 39°53ʹN;08°36ʹE

L 1996 (Wischmann PC); 2001 (Ferrito et al. 2007); 2003 (Pappalardo et al. 2008); 2005(Valdesalici PO)

Porto Taverna 40°55ʹN;09°33ʹE

L 1995; 1997 (Wischmann PC); 2000 (Maltagliati 2002)

Pauli Majori/Figu 39°52ʹN;08°37ʹE

L 2001 (Ferrito et al. 2007); 2003 (Pappalardo et al. 2008)

Marceddì 39°43ʹN;08°30ʹE

L 2001 (Ferrito et al. 2007); 2003 (Pappalardo et al. 2008)

S’Ena Arrubia 39°49ʹN;08°33ʹE

L 2001 (Ferrito et al. 2007); 2003 (Pappalardo et al. 2008)

Cala Gilgolu pond 40°54ʹN;09°38ʹE

L 2000 (Maltagliati 2002)

Stagno Longu 40°50ʹN;09°41ʹE

L 2000 (Maltagliati 2002)

Raza ‘e Juncu pond 41°07ʹN;09°32ʹE

L 2000 (Maltagliati 2002)

Palau saltworks 41°10ʹN;09°24ʹE

S 1995 (Wischmann PC)

Isuledda 41°08ʹN;09°26ʹE

L 2002 (Wischmann PC)

(Continued )

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Table I. (Continued).

Locality Lat.; Long. Habitattype

Year (reference)

Muravera 39°25ʹN;09°35ʹE

L 1996; 1997 (Wischmann PC); 2000 (Maltagliati 2002)

Colostrai pond 39°21ʹN;09°35ʹE

L 1996; 1997 (Wischmann PC); 2000 (Maltagliati 2002); 2008 (Onnis PC)

Quartu pond 39°15ʹN;09°11ʹE

L 1822–23 (MNHN); 1827 (MNHN); 2000 (Maltagliati 2002); 2005 (MNHN)

Molentargius pond 39°13ʹN;09°10ʹE

L 1999 (Onnis, PC)

Macchiareddu 39°13ʹN;09°02ʹE

L 2001 (Demontis PC)

Cagliari Pond 39°14ʹN;09°01ʹE

L 1996; 1997; 2002 (Wischmann PC); 2004 (Pappalardo et al. 2008): 2005–2006(Valdesalici PO; de Rham PC)

Cagliari saltworks 39°12ʹN;09°02ʹE

S 1901 (Mazza 1901); 1920s (Binaghi 1929); 1929–30 (Musio 1930); 1962(Cavicchioli 1962); 2001 (Demontis PC)

Capoterra pond 39°11ʹN;09°01ʹE

L 2005 (Valdesalici et al. PO)

Carloforte 39°08ʹN;08°18ʹE

L 2001 (Demontis PC); 2005 (Valdesalici PO); 2007 (Barone PC)

S.Antioco 39°03ʹN;08°27ʹE

C 1997 (Wischmann PC); 2001 (Demontis PC); 2005 (Valdesalici PO)

Nora 38°57ʹN;08°36ʹE

L 2002 (Spiga & Mattiello PC); 2005 (Valdesalici et al. PO)

Porto Pino 38°57ʹN;08°36ʹE

L 2002 (Spiga & Mattiello PC); 2005 (Valdesalici et al. PO); 2006 (de Rham PC)

Orosei 40°22ʹN;09°43ʹE

L 2010 (Spiga PC)

Porto Pollo 41°11ʹN;09°17ʹE

L 2012 (Gluszkiewicz PC)

La Maddalena 41°15ʹN;09°24ʹE

L 2013 (Valdesalici PO)

Platani river 37°34ʹN;13°41ʹE

I 2001 (Naselli-Flores PC)

Gallo d’Oro river 37°26ʹN;13°50ʹE

I 2005 (Grillo PC)

Fiume Fiumicello 37°33ʹN;13°48ʹE

I 1991 (Vitturi et al. 1995)

Marina di Modica 36°42ʹN;14°47ʹE

M 1994 (Ferrito & Tigano 1996); 1995 (Ferrito et al. 2007); 2001 (Ferrito et al. 2003); 2002;2006 (Ferrito et al. 2013)

Ganzirri pond 38°15ʹN;15°37ʹE

L 1992; 1994 (Ferrito & Tigano 1996); 2000–2001 (Minniti et al. 2003; Ferrito et al. 2007);2003; 2007 (Ferrito et al. 2013)

Marcellino Rivermouth

37°15ʹN;15°12ʹE

M 2001 (Ferrito et al. 2007); 2005 (Pappalardo et al. 2008)

Canale di Brucoli 37°17ʹN;15°11ʹE

C 1985; 1990 (Fruciano PC); 2002; 2004 (abs. Fruciano PC)

Simeto River 37°29ʹN;14°53ʹE

I 1977; 1981; 1982; 1986 (Ferrito & Tigano 1996); 1992; 1994 (abs., Ferrito &Tigano 1996)

Imera River 37°42ʹN;14°09ʹE

I 1982 (Tigano & Ferrito 1984); 1986 (Ferrito & Tigano 1996); 1992; 1994 (abs., Ferrito &Tigano 1996)

Salso river 37°34ʹN;14°13ʹE

I 1978 (MNCN); 1982 (Tigano & Ferrito 1984); 1983 (Ferrito et al. 2007); 1986 (Ferrito &Tigano 1996); 1991–1992 (Vitturi et al. 1995); 1992; 1994 (abs. Ferrito &Tigano 1996); 2005 (Schneider PC); 2007–2009 (Lo Duca & Marrone 2009)

Pantani di Vandicari 36°48ʹN;15°06ʹE

L 1996 (Di Giorgio PC); 2001 (Ferrito et al. 2007); 2004 (Marrone PC); 2005 (Duchi 2006;Pappalardo et al. 2008); 2006; 2008 (Ferrito et al. 2013)

Siracusa saltworks 37°03ʹN;15°16ʹE

S 1982; 1986 (Ferrito & Tigano 1996); 1991–1992 (Vitturi et al. 1995); 1992; 1994 (abs.,Ferrito & Tigano 1996)

Ribellino 37°05ʹN;15°18ʹE

I 1981–82 (USNM); 1992 (Ferrito & Tigano 1996)

Trapani salt work 38°01ʹN;12°31ʹE

S 1991–1992 (Vitturi et al. 1995); 2001 (Tigano et al. 2006); 2003 (Ferrito et al. 2013); 2005(Randazzo PC) 2006 (Pappalardo et al. 2008; Ferrito et al. 2013); 2010 (ValdesaliciPO); 2013 (Grillo PC)

Marsala pond 37°50ʹN;12°29ʹE

L 1994; 1996 (Maltagliati 1999); 2001 (Tigano et al. 2006; Marrone PC); 2003; 2006(Pappalardo et al. 2008); 2007 (Sardo, PC)

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decades. On the other hand, along the coasts ofSardinia and Sicily, the historical distributionappears severely narrower with respect to the currentdistribution. This is due to the scarce number ofstudies on A. fasciatus carried out before the 1990s,as several suitable environments for this species arepresent along the coastlines of both islands. Theapparently narrower range in Sardinia and Sicily istherefore the result of a lack of accurate sampling.This historical reconstruction is highly consistentwith the continuous distributional map given byBianco (1993) (Figure 1A), as all recorded pointsfall within the identified areas.

The comparison of Bianco’s (1993) map with thedistribution of A. fasciatus relative to the 1990s(Figure 1C) is consistent with the hypothesis of thespecies’ range reduction, at least in some parts of theItalian territory. At the same time, the distribution ofA. fasciatus in Sardinia and Sicily appears broadened;as already stated, this is an artefact due to the recentgreater research efforts made throughout the Italiancoasts. In Sicily, A. fasciatus appears severely rarefiedin inland water bodies, whereas its presence is quitestable in coastal environments (Ferrito &Tigano 1996). Although its absence from several envir-onments along the Italian coasts was not accuratelyproved, we can speculate that a similar reductionoccurred both along the Tyrrhenian and the southernAdriatic coasts. The species was not reported fromnorthern Tuscany, Campania (from the Fusaro Lake,where it was surely present at least until 1960s) andfrom several areas along the Adriatic coast of Apulia.Populations from Sardinia and north Adriatic regionappear unchanged from a distributional point of view,even though there are no records in the VeniceLagoon, due to the lack of studies in that area.

After the 1990s, A. fasciatus showed signals ofrecovery in many areas where it was scarcelyreported before that period. In particular, it reap-peared in Sicilian inland waters, in northernTuscany, and it was also reported from severallagoons and coastal ponds along the Ionian andAdriatic Apulian coasts. The distributional mapobtained through the post-1990s records of the spe-cies (Figure 1D) can be regarded as the currentdistribution of A. fasciatus in the Italian territory.This distribution differs in some parts from the refer-ence map provided by Bianco (1993) (Figure 1A).Along the Tyrrhenian coast, the species has beenrecorded from the low basin of the Arno River(Tuscany) to the Cape Circeo (southern Latium),with quite scattered areas of presence. The stronganthropogenic impact and the consequent relevantalteration or loss of wetlands along the coast ofLatium and Campania may account for the absence

of records south to Cape Circeo. The distribution ofA. fasciatus along the Tyrrhenian coast is thereforesignificantly narrower than in the reference map. Inthe northern Adriatic region, the species is present inthree lagoon systems: the Grado and MaranoLagoon, Venice Lagoon, and the Comacchio andPialasse Lagoon system with related canalisations.The observed distribution matches the referencemap, even though there are no species records inthe areas between the lagoon systems. These threesystems appear discrete and well defined; theirseparation, due to quite large rivers (the Piave andTagliamento rivers between the Grado-Marano andthe Venice lagoons, the estuary of Po river betweenthe Venice lagoon and the Comacchio-Ravennasystem) supports the hypothesis of three discreteareas.In Apulia, the presence of two discrete areas (one

in the north, another in the south) is confirmed, eventhough, as regards southern Apulian biotopes, popu-lation connectivity between Ionian and Adriaticareas is restricted by the presence of long stretchesof unsuitable habitat. Low population connectivitycan be also assumed between northern and southernApulian populations along the Adriatic coastline.Therefore, the more scattered distribution obtainedin the present study is probably more representativeof the actual species distribution.As regards the Italian peninsula, the discrepancies

between the reference and current distributional mapare very few and mainly due to changes in the distribu-tion of A. fasciatus, and probably an over-simplificationof a naturally scattered distribution. Such discrepanciesbecome larger when considering Sicily and Sardinia,where, in recent decades, a high number of investiga-tions on brackish-water environments were carried out.Bianco (1993) reportedA. fasciatus only in the southernand eastern parts of Sicily; our data show that the speciesis present also in the Ganzirri Lagoon (northeasternSicily) and in the system constituted by Trapani salt-works and Marsala Lagoon (western Sicily). InSardinia, records of A. fasciatus are densely distributedalong the whole island coastline, whereas in the refer-ence map the species is considered present only in thewestern part. Given the presence ofmany suitable envir-onments along the Sardinian coasts, which, in general,are much less human-impacted than the peninsulacounterparts, the observed distribution of A. fasciatusin the area is not surprising.

Environmental preferences and distribution in differentbrackish-water environments

The grouping of censused brackish-water biotopes inthe five abovementioned habitat categories was

Census of Aphanius fasciatus in Italy 7

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complicated by the strong alterations carried out byman on the natural environment, in particular afterthe realisation of numerous drainages that occurredmostly in the first decades of the twentieth century.For instance, the Margherita di Savoia saltworks inApulia were formerly the ancient Salpi Lake, whichwas completely drained in the first part of the nine-teenth century and transformed into saltworks (Afande Rivera 1845); therefore, ancient records for thatbiotope should be referred to a coastal pond, whilethose after the 1850s should be referred to saltworks.

Reasonably, the temporal changes of a number ofcensused biotopes registered in most habitat types isaffected by a bias related to differences in researcheffort across the considered periods. Instead, therecorded temporal reduction of “inland waterbody” biotopes is actually due to local extinctionsof A. fasciatus populations that occurred mainly innortheast Italy. In the periods 1990s and post-1990s,all records of A. fasciatus from inland waters dealwith observations carried out in Sicily.

As already observed for similar species (Claveroet al. 2007; Verdiell-Cubedo et al. 2014), A. fasciatusreaches high population densities mainly in environ-ments that are marginal for other species, while inenvironments that support a high fish diversity, it istypically outcompeted and relegated to specificmicrohabitats. Coastal ponds and saltworks are char-acterised by strong variations of main chemical–phy-sical parameters, and are therefore scarcely suitablefor other fish species; as our result show, the speciesis far more widespread in this kind of environment.Probably, the distribution of A. fasciatus in inlandwater bodies was restricted since its origin, as in themajority of Italian inland waters there are not histor-ical records; however, human alteration of this habi-tat type and, in particular, the competition with thealien poeciliid Gambusia holbrooki determined theextinction of a number of A. fasciatus populationsin inland waters of the Italian peninsula. Comparedto the “lagoon/coastal pond” habitat type, “canals”and “river mouths” are less represented in our surveybecause they host quite diverse assemblages of fish,representing unfavourable habitats for A. fasciatus,due to the high predation and competition pressure.

Implications for conservation

Following the International Union for Conservationof Nature (IUCN) Red List categories and criteria(Kottelat & Freyhof 2007), A. fasciatus is includedwithin the “Least Concern” category for its widedistribution, abundance and lack of major wide-spread threats. The results from our study are con-sistent with this assignment, but we want to focus on

particular cases of small-sized populations in unpro-tected areas, such as that present at Elba Island(Maltagliati & Camilli 2000; Maltagliati 2002), orthe other genetically differentiated population livingin a small pool in close proximity to the Pilo pond innorthern Sardinia (Maltagliati et al. 2003).Moreover, freshwater populations of A. fasciatusshould be also considered of high conservation inter-est. In this regard, there are historical records fromnorthern Italy freshwater habitats, as for instancefrom the Treviso town area at the end of 1800(Ninni 1863), Ravenna town around 1960–1970and Francolino (Ferrara town fluvial harbour) untilthe 1980s. The last populations were completelyeradicated by the competition with G. holbrooki (S.Valdesalici pers. obs.; D. Magnani pers. comm.). Atpresent, the only Italian freshwater populations thatshow acceptable conservation conditions are loca-lized in the Sicilian rivers, where A. fasciatus wasconsidered extinct (Ferrito & Tigano 1996; LoDuca & Marrone 2009). It is noteworthy that thesebiotopes show atypical chemical characteristics forfresh water (e.g. high sulphide concentrations andstrong seasonal salinity variations) (Lo Duca &Marrone 2009). We therefore believe that thesepopulations deserve particular conservationattention.Moreover, in some populations of A. fasciatus that

currently show good conservation conditions from ademographic point of view, past traumatic eventsmay have caused bottlenecks with consequent irre-versible genetic loss (Maltagliati 2002; Cimmarutaet al. 2003; Angeletti et al. 2010). Our survey alsorecorded cases of natural population resilience: apartfrom the abovementioned Sicilian inland waterpopulations, which appear completely re-established(Lo Duca & Marrone 2009), we were able to locateA. fasciatus populations in northern Tuscany, whereit was considered extinct. Although A. fasciatus iscommonly included in northern Tuscany specieslists as species of conservation concern, the specieswas not found in the past few decades despitedetailed monitoring in the whole of Versilia and inthe Massaciuccoli Lake (F. Maltagliati pers. obs.).The absence of A. fasciatus north to the Arno Riverhas been typically considered a consequence of com-petition with G. holbrooki; however, to our knowl-edge, there are no data that confirm the historicalpresence of this species in the area (N.E. Baldacciniand P. Ercolini pers. comms) and a recent checklistof fish of the Arno basin did not mention it(Nocita 2007), even if several suitable habitats arepresent. In November 2013, we observed A. fasciatusin the Imperiale canal, close to Livorno town, morethan 50 years after Cavicchioli’s (1962) record.

8 S. Valdesalici et al.

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Research on the recovering status of this populationis in progress. Some characters of the study area arenevertheless quite encouraging because the coaststretching from the Scolmatore Canal to theMassaciuccoli Lake is under protection within theMigliarino, San Rossore and MassaciuccoliRegional Park, and the coastline houses several tran-sitory water bodies that may provide suitable habitatfor A. fasciatus. The reasons for this populationrecovery are currently unknown. We can speculatethat the absence of the competitor G. holbrooki in themajority of the surveyed Sicilian environments hasbeen of paramount importance for the recovery of A.fasciatus populations, whose decline was probablydue to other factors. On the other hand, G. holbrookiis widespread in the environments of northernTuscany. In this case, the reappearance of temporarywetlands, beforehand removed by land reclamationand construction of coastal buildings, may have fos-tered the species’ recovery in this area.

On the basis of the data collected in the presentwork, we can speculate that A. fasciatus experi-enced population reductions throughout its rangein the 1990s, when the species appeared locallyextinct, or strongly rarefied, in several localitieswhere it was found one decade before. Sincethen, the species has recovered in the majority ofthe native range, with records of stable populationsalso in areas where it was considered extinct.However, competition with G. holbrooki remains acritical hurdle to the re-conquering of freshwaterand oligohaline environments and, reasonably, thespecies’ genetic diversity has been altered by thepast contractions, in terms of genetic loss.Therefore, even if, generally, A. fasciatus showsclear signals of recovery, it would be erroneous toassume that its situation in Italian waters hasreverted to the pristine condition.

The overall situation of A. fasciatus in Italy can beconsidered quite good, at least if compared to thesituation of the other European killifish (Doadrioet al. 1996, 2002; Gómez 1997; Kärst 1998; Planelles& Risueño 1999; Kottelat & Freyhof 2007). However,continued population monitoring and protection is acompelling task. The present research represents thefirst step toward the desirable completion of the dis-tributional map of A. fasciatus in the wholeMediterranean region.

Since wetlands are protected at national, Europeanand international levels and many of the known local-ities are natural reserves, Sites of CommunitaryImportance (SCIs) and Special Protection Areas(SPAs), it is likely that many A. fasciatus populationswill not be affected by future improper human activ-ities. Because of its life history features,A. fasciatus can

be considered an environmental indicator of an impor-tant habitat type, whose ecological relevance has beenrecognised only recently and that appears to be inregression along the majority of the Italian coasts. Infact, many coastal wetlands and their typical assem-blages have been eradicated from the Italian coastlineand in several regions they survive mainly in protectedareas. Along with A. fasciatus, a number of indigenousspecies typically associated with coastal wetlands arethreatened and in strong regression across the wholeItalian territory. In the framework of an awarenesscampaign about the importance of coastal wetlands,as both biodiversity hotspots and buffer areas againstrainfalls and floods, A. fasciatus may be raised to thestatus of “umbrella species” (Simberloff 1998;Roberge & Angelstam 2004) with regard to this habitattype.

Acknowledgements

We wish to thank R. Froese for his first goad, F.Fontana, D. Montanari, A. Varriale, G. Paonessa (forthe first idea) and the Migliarino, San Rossore andMassaciuccoli Regional Park for giving us permissionto sample in its territory. This work would have beenimpossible without the contributions of many biolo-gists, naturalists or simply amateurs; therefore wewould like to thank also F. Andreone, N.E.Baldaccini, R. Baldacconi, S. Bandella, G. Bardelli,C. Barone, R. Bufo, A. Cappon, M. Capula, G.Casamenti, C. Castellarin, M. Casu, R. Cazzulani, R.Cellere, A. Cellerino, F. Cepollaro, G. Comisso, D.Cuni, A. Demontis, F. Denitto, P. de Rahm, E. DiGiorgio, W. Eberl, P. Ercolini, S. Esposito, E. Farmer,S. Fick, F. Fontana, S. Frosini, C. Fruciano, E.Gargiulo, T. Gluszkiewicz, A. Grillo, H. Kärst, F.Liverani, D. Lo Stimolo, D. Magnani, G. Mancini,F. Marrone, P. Mattiello, L. Morreale, L. Naselli-Flores, P. Nicolosi, C. Nocchi, C. Onnis, M. Parisi,A. Piccolo, M. Prono, M. Randazzo, A. Ricci, S.Ruozi, P. Salvagiani, A. Sardo, M. Schneider, A.Sensoli, M. Silvestri, A. Solenne, G. Solís Fraile, A.Spiga, M. Tavano, J. Thorup, J. Valdesalici, C. Valle,S. Vanni, A. Varisco, C. Vianello, H. Wischmann, E.Zubani and M.A.L. Zuffi for providing data, M. DalBello for technical support and two anonymous refer-ees for their valuable comments and corrections.

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