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The Evolution Dynamics of the Strigiformes in the Mediterranean Islands with the Description of Aegolius Martae N. Sp

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The Evolution Dynamics of the Strigiformes in the Mediterranean Islands with the Description of Aegolius Martae N. Sp View metadata, citation and similar papers at core.ac.uk brought to you by CORE provided by Institutional Research Information System University of Turin ARTICLE IN PRESS Quaternary International 182 (2008) 80–89 The evolution dynamics of the Strigiformes in the Mediterranean islands with the description of Aegolius martae n. sp. (Aves, Strigidae) à Marco Pavia Dipartimento di Scienze della Terra, Museo di Geologia e Paleontologia, Via Valperga Caluso 35, I-10125 Torino, Italy Available online 14 June 2007 Abstract Living and fossil owls (Aves, Strigiformes) constitute an important group for understanding the evolutionary dynamics of birds in island environments. After their different trends in island evolution, the Strigiformes can be seen as a representative of insular adaptations of birds as a whole. In fact they respond quickly to isolation with deep changes in body size, including dwarfism and gigantism, and allometric variations, such as reduction of wings, lengthening of hindlimbs and strengthening of digits and claws. The only exception is the loss of the ability to fly, which has never been recorded in Strigiformes. In this paper I report on all the endemic owls found in Mediterranean Islands, both living and fossil, in order to emphasize trends in insular evolution and the relationships between the different species sharing a certain island. The description of Aegolius martae n. sp. completes the guild of endemic Strigiformes of the early Middle Pleistocene of Sicily and allows to use Sicily as the best example of a biogeographical island type with intermediate characteristics between the oceanic and the continental ones, with the presence of some non-flying mammals, but the lack of terrestrial carnivores. r 2007 Elsevier Ltd and INQUA. All rights reserved. 1. Introduction islands are inhabited only by a subset of continental species, those with the highest vagility; they tend to lack After the works of Darwin (1859) and Wallace (1881) the non-flying mammals, amphibians and freshwater fishes. study of islands became a powerful tool in many biological Continental islands still lie close to continents, to which studies, as the study of the fossil of past faunas gives an many were connected as recently as the last glaciations, historical dimension to the insular biogeography and 10,000–12,000 years ago, when sea-levels dropped. Due to ecology. Typical insular communities are characterized by their proximity to the mainland, or to the presence of land their disharmony, the so-called ‘‘insular poverty’’, their bridges during glaciations, the plants and animals of these high degree of endemism and an organisation of ecological islands are more or less representative subsets of mainland relationships among species that are different from those species, even if some new species can eventually evolve. on continents. The number of terrestrial bird species found on a Birds are a key group in understanding the biological particular island is normally related to the size of the changes in island environments. According to their island and its degree of isolation. These two parameters geological origin, islands can be divided into two different lead to the equilibrium theory of MacArthur and Wilson groups: oceanic islands and continental ones. The former (1967), according to which the rate at which a species are volcanic islands of fairly recent origin, less than 10 reaches oceanic island declines with the increasing remote- million years, which have emerged from the ocean floor ness of the island, while the rate of extinction of an and have been never connected to any continent. Con- established species declines as the size of the island tinental islands were originally parts of a continent, which increases. Certain case studies show that the colonisation have since separated from the main land mass. Oceanic of an island can be rapid, such as for Krakatoa, in Indonesia, where a century after the complete destruction à Tel.: +39 0116705197; fax: +39 0116705339. of the island following a volcanic eruption, more than 40 E-mail address: [email protected] bird species have been reported (Thornton et al., 1993) 1040-6182/$ - see front matter r 2007 Elsevier Ltd and INQUA. All rights reserved. doi:10.1016/j.quaint.2007.05.018 ARTICLE IN PRESS M. Pavia / Quaternary International 182 (2008) 80–89 81 along with a great variety of plants and insects. Addition- cassowary (Casuarius)(Parker, 1984; Rich et al., 1988) ally, modifications such as the evolution of flightlessness and gigantism in the extinct Passeriformes Emberiza and the loss of bright colours, especially the full colour alcoveri of the Canary Islands (Rando et al., 1999). The plumage of males in dimorphic species (e.g. Anatidae: coexistence of both gigantism and dwarfism in the genus Newton, 2003), can be very rapid in insular environments. Cygnus during the Pleistocene, with the giant Cygnus Moreover the insular endemic forms tend to be more ‘‘K- falconeri and the dwarf Cygnus equitum (Northcote, 1992), selected’’ than their mainland relatives, with lower repro- both endemic to the Sicilian–Maltese archipelago, although ductive and mortality rates, and longer life spans. probably not coeval, as demonstrated by recent analyses of The study of fossil associations changed island biogeo- Sicilian fossil bird assemblages is worth mentioning graphy paradigms based only on the study of extant faunas (Bonfiglio et al., 2002). Apart from overall size variations, (Segui and Alcover, 1999). The fossil record reveals that the a significant number of insular bird species are affected by data obtainable from extant taxa alone are artificial. allometric variations. In this case, generic trends can be Natural extinctions are caused mostly by two geological detected in many bird orders, such as Falconiformes, events, volcanic eruptions and tsunamis, and less often by Psittaciformes, Strigiformes and Passeriformes, in which variations in sea levels. These factors are normally felt main variations concern bill, wing and tarsus measure- more strongly on small oceanic islands. In addition to ments (Grant, 1965; Segui, 1998; Rando et al., 1999; Suarez natural extinctions, many island birds have disappeared in and Olson, 2001; Louchart, 2005). The modifications the last 1500 years because of human actions. Even if affecting wings normally lead to a reduction or loss in human impact is not always clearly demonstrated, in each the ability to fly, as recorded in at least 14 bird orders case the extinctions followed soon after human colonisa- (Louchart, 2001). Trends in allometric variation can be tion (Newton, 2003). Human influences are usually summarized as an increase in the relative size of the bill and indirect, such as the introduction of predators or compe- the relative length of the tarsus, and a decrease in the titors and landscape modifications, but direct over-hunting relative size or length of the wing. These trends are usually is sometimes an important factor (James and Olson, 1991; explained by more terrestrial habits, specialised and Olson and James, 1991; Milberg and Tyrberg, 1993; sedentarity (Louchart, 2005). Steadman, 1995; Mourer-Chauvire´et al., 1999; Blackburn As recently stated by Louchart (2005) the Strigiformes et al., 2004). The lack of recently extinct taxa in present- are a key group in understanding the evolutionary trends day insular associations, even if still endemic, make them of birds in insular environments. In this paper I deal with unnatural. The only way to approximate natural commu- the Strigiformes inhabiting Mediterranean islands during nities and complete taxon lists is to integrate both living the Neogene, as they show all the types of modifications faunas and recently extinct taxa (Louchart, 2005). This is that occur in birds. The description of Aegolius martae n. especially true for birds, where the number of extinct sp. also completes the Strigiformes guild of the early species in the Late Pleistocene and Holocene is very high Middle Pleistocene of Sicily, presently composed of two (Milberg and Tyrberg, 1993). The study of island fossil other extinct species, Tyto mourerchauvireae and Athene birds also reveals that extinction processes are often trinacriae (Pavia and Mourer-Chauvire´, 2002; Pavia, 2004). selective; in fact selective impacts are responsible for eradicating larger species along with the ones resulting from extreme evolutionary pathways. 2. The endemic Strigiformes On the basis of the study of many fossil island bird associations (see Louchart, 2005 for a list of references), The Strigiformes constitute one of the best groups for general trends of modifications occurring in endemic island studying insular bird taxa, both living and fossil. Indeed, bird species can be summarized, if any exist. It is not clear about 150 of the 611 recognized extant taxa are insular whether general trends are towards dwarfism, gigantism or endemics, while extinct endemics recorded so far include other allometric variations, although it seems that gigant- only 36 taxa from the Miocene up to the present. ism, along with the reduction or loss of the ability to fly, is Furthermore, they are widespread and frequently exhibit more frequent. There have been few studies so far, most of size and allometrical modifications; moreover, the extinct them restricted to limited groups of species, based on taxa have an abundant fossil record on islands, and are extant forms and often considered only at a low taxonomic also relatively easy to identify osteologically, as even the level (Grant,
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