Note on the Origin of the Present Mammalian Fauna from the Balearic and Pityusic Islands

NOTE ON THE ORIGIN OF THE PRESENT MAMMALIAN FAUNA FROM THE BALEARIC AND PITYUSIC ISLANDS

JOSEPANTONI ALCOVER

Alcover, J. A., 1980 (1982). Note on the origin of the present mammalian fauna from the Balearic and Pityusics islands. Misc. Zool., 6: 141-149. Barcelona.

Given the present state of palaeontological knowledge today's mammalogical fauna of the Balearic and Pitysic Islands can be considered as coming from a recent colonization resulting from the activity, voluntary or not, of man. In this paper the number of mammalian species are considered in tems of the theory of island biogeography. The human activity would have brought about a deterministic renewai of the island's mamma- logicai fauna, with a noticeable increase of the number of species and with the substitution of some stenocorous species, subjected to K selection during many years, by euricorous species, which are typicai r strategists.

Josep Antoni Alcover, Dptnt. de Zoologia (Vertebrats), Fac. de Biologia, Universitat de Barcelona, Av. Diagonal 637-647, Barcelona 28.

INTRODUCTION that some components of the present balearic mammalian fauna may have been It is still taken for granted by various established during Messinian. In this work I authors that the colonization of the Balearic will try to demonstrate the shortcomings of and Pityusic Islands by the present mam- these ideas, on the basis of the paleonto- malian fauna is very ancient, and happened logical knowledge which we now possess long before the first arrival of man. Only about the components of the present mam- very few classical mammalogical papers pose malian fauna from the Balearic and Pityusic the question of colonization (THOMAS, Islands. 1901; CABRERA,1914; KOLLER,1941), and UERPMANN(1971), REY & REY (1974) and COLOM (1978) have very re- PHYLOGENETICAL ANALYSIS OF THE MAM- cently expressed the idea of an ancient MALIAN BIOTA FROM THE BALEARICS origin for this fauna as common place. The former (UERPMANN , 197 1) distinguishes in Before beginning the analysis of the phylo- this work those species which were introduced genetic characteristics of the species which into Mallorca by man from other species compose this fauna, it is worth noting that which, in this opinion, should be considered none of. them has ever been found in the as autochtonous and which presumably en- Quaternary of the Balearics. The remains tered in ancient times, through a "Land- indicated by ADROVER (1966) at Mu- briicke". REY & REY (1974) are amazed by leta Cave are aii Holocenic or more re- the fact that the shrew Crocidura suaveolens cent. However, from this lack of findings we is not found at present in Mallorca, while cannot conclude anything about the absence this island "se ha mantenido discontinua- of this fauna in past times, although the mente unida a Menorca (where C. suaveolens abundance of quaternary mammalian re- does now exist) a 10 largo de diversos pe- mains found ui till now makes us think riodos geológicos". In his turn COLOM that we already have a complete record of (1978) suggests, in the modified text of the the Quaternary fauna of terrestrial, non- new edition of his biogeographical book, flying, marnmalian phyla. The fact that the known Quaternary terrestrial mammals the accurate range of geographical distri- (Myotragus, Nesiotites, Hypnomys) present bution of the species. STORCH (1970) some traits which may be well understood and MALEC & STORCH(1972) consider and explained as adaptations to an insular that E. algirus has been very recently intro- environment in the absence of predators and duced by man into Malta from the neigh- in the absence or scarcity of competitors bouring African mainland. At present, the (LEINDERS & SONDAAR,1974; AL- ancestor species of E. algirus is not known; COVER & ROCA,1975; ALCOVER, 1976; in fact, not even fossil or subfossil remains SONDAAR , 1977) is not in itself conclusive; of this species are known. It seems that the nor can we draw any conclusion from the apparition of the genus took place about the knowledge of the average life of Mammalian late Upper Miocene (KURTEN, 1961 a; species, which, according to KURTEN(1961 SABAN,1958) or about Lower Pliocene b) oscillates around 540.000 years (inclu- (THENIUS,1969), so perhaps it did not yet ding, for this estimation, the Chiroptera, an exist during Messinian. The evolutive rate of order which includes very conservative Insectivora is, in addition, relatively high. species, with an average life of 1.600.00 The average life of their species oscillates years). In the orders wich we shall consider around 490.000 years. Furthermore, only in this work, the average life of the species is one species of Erinaceidae which evolved in sit uated around the following figures: insular conditions has ever been studied, 490 .O00 years (Insectivora, Rodentia), Deinogalerix koenigswaldi, from the Neo- 600.000 years (Lagomorpha), 610.000 years gene of Gargano (Italy; FREUDENTHAL, (Carnivora). The morphological plasticity of 1972) and we know that it acquired a phy- the Rodents has been well studied by siognomy which makes it very different BERRY and his collaborators, who re- from any mainland Insectivora. Another corded, under insular conditions, morpho- fossil insular Erinaceidae has been found at logical changes that took place in Muridae in Capo Mannu (Sardinia; PECCORINI,RAGE only 70 years (BERRY,1964). In any case, & THALER,1974). As far as the genus for our purposes we shall analyse the em- Erinaceus is concerned we know that the pirical data supplied by phylogeny and species E. europaeus, phylogenetically rather spatial and temporal paleodistribution of the close to E. algirus, is found occasionally in different taxa involved. deposits of Upper Pleistocene and Holocene The Hedgehog inhabiting the Balearic and from Europe (ALTUNA,1972). Their re- Pityusic Islands, Erinaceus algirus, lives also mains found in wiirm-I1 of Grotte Hortus along a narrow coastal strip of eastem and "sont d'une taille nettement tres supérieure southem Spain, in southern France, as well aux plus grands indvidus actuels" (JUL- as in vast regions of northern Africa and in LIEN, 1972). All these facts, namely, the the islands of Malta and Djerba. The pattern plasticity of this related species, the data on of distribution which this species present the average life of Insectivora species, the made COLOM (1978) think -following tirne of apparition of the genus, the con- perhaps the considerations of CABRERA sideration of the evolution of an Erinaceidae (19 14)- that it was autochtonous, with an in insular conditions, the identical morpho- assigned entrance during Messinian. logy of the Iberian and insular populations KOLLER(1941) also assigned it an ancient of E. algirus (belonging to the same sub- origin, and concluded that it had spread species, vagans), and the fact that this spe- from southern Spain. UERPMANN(1971), cies may be easly transported by man who did not find it at S'Illot, also assumed it (VERICAD& BALCELLS,1065;MALEC & to be autochtonous. KAHMANN& VES- STORCH,1972) make us think that E, al- MANIS (1977) say that they have not con- girus was brought by man to the Balearic sidered human interventions in presenting and Pityusic Islands. As far as the shrews, Cvocidura russula all of these (finds being) Late Pleistocene from Eivissa and C. suaveolens from Me- age ...Even more uncertain are records from norca, are concerned, the present state of the Middle Pleistocene of small forms that knowledge does not allow any conclusion at may belong in the ancestry of its species". all. The fossil fmdings of both species re- In the Middle Pleistocene of Cartagena, the corded in Europe all date from Upper Pleis- present species of rabbit has not been found, tocene. All the findings in the Iberian Pe- but instead a relative, Oryctolagus cf. lacosti ninsula belong to Upper Pleistocene or to (PONS & MOYA, 1978). It appears that Holocene. Numerous remains of another both taxa of Lagomorpha now iiving in the species, C. kornfeldi, have been found in Balearic and Pityusic Islands are of very re- Middle Pleistocene from Cartagena (Murcia: cent origin. UERPMANN (1971) when re- PONS& MOYA, 1978). These data make us ferring to the rabbit and hare, consider them think that both the present species are very as autochtonous, because their "hohe phylo- recent, and thus they would only have genetische Alter" (sic). COLOM (1978) con- been able to colonize Eivissa and Menorca sider also an ancient origin for both Lago- through the action (direct or indirect) of morpha. The ideas expressed by these man. In addition it must be mentioned that authors do not fit present-day knowledge of C. suaveolens has not been found in subfossil their phylogeny. These two species must be state in Menorca, in spite of the nurnerous considered as very recent and therefore we remains of small mammals coming from a set must accept their presence in the Balearic of deposits of the talaiotic age already ana- and Pityusic Islands as due to importation lysed. It is also noticeable that C. russula has by man. not appeared in the early subfossiliferous de- The Rodentia fauna from the Balearic posit which we know in Eivissa (Cova des and Pityusic Islands contains 6 species: Cuieram). It seems clear that C. suaveolens is Eliomys quercinus (F. Giiridae), Apodemus a species that colonized Menorca after 500 sylvaticus, Mus musculus, M. spretus, Rattus years B. C. (date of the later subfossiliferous rattus and R. norvegicus (F. Muridae). deposit studied in Menorca). Unfortunately Among these species, M. musculus, R. rattus the deposit of Cova des Cuieram is very and R. norvegicus are typical comrnensals of poor, and of undetermined age, so it does man. All the authors consider them as not allow us to say anything about the chro- allochtonous. M. musculus may have been nology of colonization of Eivissa by C. rus- imported before talaiotic age (UERPMANN, sula. 1971), while the Rattus species may have The genera of Lagomorpha that live at colonized the islands more recently, R. the present time in the Balearic and Pityusic rattus probably during the Middle Ages, and Islands are not very ancient. According to R. norvegicus provably around the first half THENIUS (1969), "Ihr ersters Nacheweis of the 19 th century (ALCOVER,1979). starnmt aus den alterlen Pleistozan". DE- About M. spretus we can say very little. SACHEAUX (1958) holds the same view, This species has very recently identified and as does WALKER(1968) when he reports typified as an independent taxon (BRIT- the genus Lepus. As far as species are con- TON, PASTEUR & THALER, 1976), and cerned KURTEN(1961 a) states that "some now it will be necessary to revise all the records of the modern rabbit evidently back fossil material considered up till now as M. as far as the D-Holstein (e. g. Lunel-Viel in musculus. In any case the fossils gathered sothern France), but more definitive identifi- under this name in Europe are only kriown cations of this species refer to the F-Eemian from Middle and Upper Pleistocene (KUR- deposits (e. g. the Pinar Cave, GranadaT... TEN, 1961a) and not from early times. For "Fossil finds in Europe have occasionally this reason, we must assume that this referred to this species (Lepus capensis) ,..., species, is of recent importation. Remains of Apodemus sylvaticus are MANN (1971) and PALACIOS,CASTRO- furnished by all the subfossiliferous deposits VIEJO & GARZON(in RODRIGUEZ DE from the Balearic and Pityusic Islands, and LA FUENTE,1975) developed rather ima- UERPMANN (1371) considers it as autoch- ginative theories in order to explain the tonous, because it is a very early taxon. The origin of the present day distribution of sub- presence of the genus Apodemus in the species groups quercinus (with the underpart European fauna goes back to the end of of the tail completely white) and lusitanicus Turolian (MICHAUX& PASQUIER,1974; (with a black ring on the under part of the CHALINE& MEIN, 1979). Nevertheless, it tail). In Central Europe, remains of E. quer- must be taken into account that during Plio- cinus from Lower and Middle Pleistocene cene this genus was represented by two have been found. Previusly, other species of extinct species, A. primaevus and A. do- the genus are found, which are probably its minans. The oldest remains of A. sylvaticus ancestors, E. intermedius from Upper Plio- come from the early Middle Pleistocene. In cene and E. truci from Middle and Lower different deposits of Lower Pleistocene from Pliocene (STORCH,1978). It is clear then, the Iberian Peninsula and southern France that the present species could not have reach- an Apodemus sp. has been found that may ed the Balearic and Pityusic Islands during perhaps belong to the evolutive line of A. the Messinian, because it did not yet exist. sylvaticus (MICHAUX& PASQUIER,1974). Furthermore, it must be pointet out that the In addition, it is known that A. sylvaticus is morphological plasticity of Gliridae is very morphologically very plastic, with more than high. Some endemic genera are known from 30 subspecies described in Europe the Plio-Pleistocene of certain Mediterranean (NIETHAMMER,1978) and at least 3 in Islands, Hypnomys (Mallorca and Menorca), northen Africa (SAINTGIRONS,1972). Leithia (Sicily and Malta) and Thywenoglis Besides, we know that during the Quater- (Sardinia). It seems reasonable to assume nary giant species of Apodemus lived in that if any species of Eliomys had colonized Sardinia (A. mannu; PECCORINI, RACE & the Balearics or Pityusics during the Mes- THALER, 1974) and in Sicily (A. major; sinian, it would have evolved in a quite dif- THALER , 1972). A very ciose genus, Rhaga- ferent way from those on the mainland, podemus, colonized Sardinia and originated without becoming E. quercinus. Thus, we the endemic line of Rhagamys (R. minor and must agree with THALER(in MASCOMA, R. orthodon; BR ANDY , 1978). This mor- 1978) that its present occurrence in the Ba- phological plasticity of Apodemus and rela- learic and Pitysic Islands is only explainable ted genera facilitates rapid morphological if we accept that it has been irnported by changes when it is put under insular condi- man. tions. Thus we would expect that if an an- Regarding the wild Carnivora from the cestor of A. sylvaticus had colonized the Balearic and Pityusic Islands, it must be in- Balearic and Pirysic Islands during Messinian, dicated that the Genet, Genetta genetta, is some endemic species, or even a genus, probably an African element which probably would have originated. This has not been the invaded Europe after Wiirm (KURTEN, case, and therefore we must consider it as a 1961a), but of which no fossil remins are very recent colo~izationdue, beyond any known. The Genetta sp. material mentioned doubt, to importation by man. by ADROVER(1966) from Muleta Cave Among Rodents, only E. quercinus re- was wrongly identified, and is surely very mains to be dealt with. This species, found recent. The Weasel, Mustela nivalis, is a in all the subfossiliferous deposits of Ma- species of recent apparition, which arose lorca, Menorca and Eivissa, has been con- from another, M. praenivalis, which existed sidered as autochtonous by various authors. up to the earlier Middle Pleistocene (KUR- On the basis of this consideration UERP- TEN,1961a; THENIUS,1969). Both species time time Fig. 1. Colonization curves of mammalian fauna from Balearic and Pityusic Islands. A. After Colom (1978); B. Model proposed in this paper. m. Messinian; h. arrival of man; n. present-day species number. of Marten which occur now in the Balearic Balearic and Pityusic Islands depends. Fig. and Pityusic Islands are likewise of relatively 1A shows the classical conception of the recent origin. They arose from Martes vetus colonization process of these islands by (ANDERSON, 1970), a species found in mammalian fauna. It has been drawn on the Lower Pleistocene from Europe. The oldest commentaries of COLOM(1978), who states remains of M. martes come from Middle that "desde el punto de vista de sus ma- Pleistocene, while those of M. foina come miferos las tres islas se caracterizan por po- from Upper Pleistocene. All these data can seer un reducido conjunt0 de especies, con- only be related to a colonization of the junto que a través de 10s tiempos cuater- Balearic and Pityusic Islands due to importa- narios y en la fase antigua de 10s actuales ha tion by man. ido empobreciéndose cada vez mis debido a Considering all we have said, we can con- la misma acción del medio insular", and far- clude that the present terrestrial (non-flying) ther on, delaing also with Mammals, he mammalian fauna from the Balearic and speaks of the "...empobrecimiento paulatino Pityusic Islands has all been imported by de las biotas isleñas por la acción directa del man. The source areas have varied, and hombre". Fig. 1B supports the ideas that further data is needed in order to determine have been presented in this paper. Both them accurately; for this reason, they will be graphs are schematic, as they simply try to subject to future researchs. It must also be summarize each theory, without making an considered that the colonization process has accurate analysis of the colonization process. been complex, and some islands have pro- According to our view, the presence of bably acted as stepping stones in the colo- man would certainly not have implied the nization of others. gradual impoverishment of the insular mammalian fauna, at least as for as the number of species is concerned. On the contrary, it may be seen that there has been an increase of DISCUSSION about four times in the number of species in the Balearic Islands and probably a higher The conclusion to which this work comes increase in the Pityusic Islands. modifies noticeably some of the current It seems that the approach of COLOM concepts on which the present-day biogeo- (1978), according to which "las plantas y 10s graphical conception of the biotas from the animales que ahora se desenvuelven en las islas son 10s descendientes directos de 10s these islands continues), higher than the que estuvieron en posesión de facilidades former, S. From the similarity of the mam- innatas para una supervivencia en las pecu- malian faunas presently inhabiting the Ba- liares condiciones restrictivas de 10s monóto- learic and Pityusic Islands together with the nos hiotopos insulares", is not entirely relatively large size of the species pool in suitable to explain the present-day mam- source mainland areas, it may be inferred malian fauna. It seems more proper to con- that the new immigration curve must be very sider the present fauna not merely as a re- concave (see MAC ARTHUR & WILSON, sidue of an ancient fauna, but as the result 1967). So we may consider that the action of a balance between the immigration and of man on the Balearic and Pityusic mam- extinction of species, as the theory of island malian fauna has had the effect of bringing biogeography postulates (MAC ARTHUR & the islands nearer, as it were, to mainland WILSON, 1967). in this sense, it must be source areas, rather than that of reducing the considered that before the arrival of man, island areas. If this action has been similar either the mammalian saturation is not for all the Mediterranean islands, it must be reached, or an equilibrium is reached with a expected that, except for special complica- very low number of species (9 = 3 species of tions (as for instance the establishrnent of terrestrial mammals in Mallorca and Me- unions between the islands during the glacia- norca), due to the fact that the immigration tions), its effect, measured as d log Sh/d A, rate (it must not be forgotten that the Ba- will be slower in larger islands, because "the learic Islands are the most isolated islands in logarithm of the number of species decreases the whole of the Mediterranean, and that the with the distance more rapidly in the ar- dispersa1 capacity of non-flying mammals is chipelago with small islands, althouogh the very limited) and the extinction rate (once absolute number does not". This is a predic- they succeeded in their colonization, the few tion drawn directly from the theory of surviving species would be unlikely to be- island biogeography, which may readily be come extinct, thanks to the great availability tested. of empty habitats and to the lack of pre- The new number of mammalian species in dators and competitors), both being func- equilibrium, Sh, is deterrnined by a new tions of the number of species present in the balance between the immigration and extinc- insular area at any moment, were both very tion curves. We must emphasize that both low (see fig. 2A). The fact that at another processes are not stochastic, but determi- moment of the biogeographic history of the nistic. In fact, the extinguished taxa are, Balearic Islands an insular mammalian fauna among others, those which lived during of apparently only four species is found Quaternary, while those imported by man (ADROVER, 1977) leads us to think that are a selective sample from the species pool the pleistocenic mammalian fauna was pro- in the mainland source areas, including ba- bably saturated o nearly so. sically "good colonists". In this sense, we Therefore, the arrival of man implied not must speak about competitive replacement only a rise in the curve of extinctions, as of the ancient fauna of stenochorial mam- inferred from the work of COLOM(1978), mals by the present-day fauna of eurychorial but also, and to a greater extent, a rise in the mammals. Probably the same process is curve of immigration of new species; thus, found in all the Mediterranean Islands, with given that a perfect balance can never be the arrival of man and also in at least some reached between immigration and extinc- islands during previous periods of faunistic tion, because it would be approached expo- turnover (see MAYHEW, 1977). nentially, the situation tends towards a new The mammalian fauna of the Balearic and number of species "in equilibrium", Sh (in Pityusic Islands, with the action of man, equilibrium so long as the hurnan presence in probably passed from a total lack of har- A B

al al L .w ,m E

I s s, Fig. 2. Equiiibrium model of the mammalian biota of the Balearic Islands proposed in this paper. A. Equiiibrium model before arrival of man; B. New equilibrium reached in presence of man. It can be seen that a grater change takes place in the immigration curve that in the extinction curve; E. curve of rate of extinction ("extinction curve"); I. curve of rate of immigration ("immigration curve"). mony(') to a certain degree of harmony seems that is different degrees a similar and from a very low number of species in phenomenous have taken place in the herpe- equilibrium to another higher. Regarding the thological, batracological and terrestrial ma- discrete taxa affected by this process, we lacological faunas in the Balearic and Pit- may say that the action of man results in the yusic Islands and also in other, if not all, replacement of a set of taxa that are, islands. Future research is required in order without doubt, very adapted for a high to obtain an accurate knowledge of the way biological efficiency in an insular environ- these phenomena occur. ment by other opportunistic ones; of some species subject during many years to a K-se- ACKNOWLEDGEMENTS lection, so-called K-strategists (for their characteristics MARGALEF, 1974) by The author is indebtd to Llct. Francesc Avellh, Mr. Guiem Daviu, Lct. Salvador Moyh, Mr. Joan Pons others subject in varying degrees to an r-se- and very especially to Robin Aird. lection. The conclusions of this paper, although they refer strictly to terrestrial -non-flying- RESUM mammals from the Balearic and Pit~usic D'acord amb els coneixements paieontoibgics que Islands, may have a wider application. It es tenen, cal considerar la mastofauna actual de les

(1) An insular biota is considered harmonic if it contains all of the basic adaptative types found in other ecologically comparable regions.

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