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The avifaunas of the isolated Mediterranean islands during the Middle and Late Pleistocene

Article · January 1992

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Josep Antoni Alcover Cécile Mourer-Chauviré Mediterranean Institute for Advanced Studies (IMEDEA) French National Centre for Scientific Research

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The user has requested enhancement of the downloaded file. THE AVI FAUNAS OF TH E ISOLATED MEDITERRANEAN ISLANDS OURING THE MIDOLE ANO LATE PLEISTOCENE ]. A. Alcover, F. Florit, C. Mourer-Chauviré, and P. o. M. Weesie

AB STRACT. The paleurn itholugical record of rhe middlc and uppc r cnJemlques, les aus de presJ no endcmiqlles de talla corpo",1 gr.ln, l'kislocenc of the Medittrranc,1n isla!lrls Sardin ia, , Tavolara, algunes especies de córvids i PuffimH puffinus. Mallorca, "-knorc", Ej,-issa, Malta, e rete, Karpathos, Armarhia, " Eld T ylos is presenleJ. Our kuowledgl' of fossil avifaunas fro m islamls su(h Key worJs: PkislOccne, Medilcrrancan, Jl'ifallna, cvol ulion, biogeo­ as Corsica, Crele, Sardini", and Mallorca is rclativcly good bu! srill graphy, fossil birds. incomplete. The fos,il r<~cord for all of Ihe isla nds incl ll des 160 faXJ. So rn e fa unís!ic, "\lalulionary, and biogeographic considerat iOll s are dis­ cus>ed. Tho: fo ss il Med iter",,,ean islands avifallllas han' a diffcrcm spe­ cies association in (Ompa ri ~on wilh Ihe modem avifaunas on rhe 5ame INT RODUCTlON ¡stands and a grcarer numbcr of endem;c fomls than rod ay. The fossil T he M edilerranean Sea is a dosed basi n of 3,081,880 km' boro isbnd avifaunas lack species of the G ~ llifo r rnes (wit h th" "xc"ption of dered by rhe conrinenrs of F. urope, Afri ca, and Asia (Fig. l), Ir Columix wlumix), l ~rid'l<", and Passer. A ruunber of bi rd specics re(orded as fossil, do not no\\" occur in the MedÍlerranean avifallnas. conrains more !han 2,0

  • ~ da de Horno sapicns. S'han id ent ifica t laxa i nsu l ~rs and Paleolilhic rnan 10 mo ve o nto the island. tndrmics als ordres Accipiltifo nnl"S , Snigiformes, Anseriformes i I'as­ OU T presenl kno wledge of Ihe vertebral e faun<1s, and espe­ serifonnes, i la l vo lra lam bé emrc els Gruiformes i ds Columbiformes. Els canvis en In la ll a corpor.Jl deis rapinyaires Sl'mblen estar relacionals cia ll ,. o f the a vifaunas Iha' inhabited ,he Mediterranean is lands mnb I'ofena de preses nlicrott'riolOgiqu t s qUl' tOS donava a les dift'rt ms befo re Ihe arrival o f HOlllo sapicIIs Linnaeus 1758, is very in­ ilks. S'han rt-gi ~lrat extineions rcccnts i descomparcixcnccs loca ls d'al· complete. The present study presents a SUI1UTlary of the paleo r­ gunes espCde~ d·ocdls. Aq uestes ha n afectar, essencia lm en t, les especies nitho logical record of the middle nnd llPper Pleistocene on rhe ,ipiques d .. climts frtJs, les especies .. n d ~miqlJes grans, I.. s anS de pres.l 11 isolated M editerranean islnnds known to dMe to prod uce EUROPE

    ~~ ~/'/ f ~J1\""""o,",""" ~ .. -

    AFRICA

    Figure 1. .~1ap of ¡he Medllerrancan islands prodlldng ¡he paleoavifaunas wnsidcred In rhlS papeL

    fossil birds. Larer, we descnbe sorne characrcrislics of the Plcis· T here are references 10 the foss il birds of Mall orca andj or rocene island avifaunas. Menorca in the works of Ba te (1928 ), Lam brechr (1933), Villalta (1964), Ballmann and Adrover 11969), Adrover (1972), Pons and THE RECORD Roca (1974), M ourer-Chauviré et al. (1975, 1977, 1980), A1cüve r This srudy is confincd ro the avian species known from the er al. (1981 ), Reu1ll CT (1982), Nonhcote and Mourer-Chauviré middle and upper P1c isroccne, excluding those that have bcen (1988), and Flori¡ and A1cot'er (1987). fo un d in assocíatíon with Ho mo saplens on vanous Mcditcr­ The paleoavifaulla of Ei vissa was fi rst dcalt wi¡h by Moya et ranean lslands (Mallorca: Uerpmann, 1971; LevanlO: Cassoli an<] al . (1984), Alcove r and Muntallcr (1985 ), and, more extensively, Tagl iacozlO, 1982; Malta : various da ta from Lambrecht, 1933; Florit et al. (1989). Sllriano (1980) and Weesie (1982, 1987a, Sicíly: Regalía, 1907). The following insular paleoavifaunas fro m 1987h) studied ¡he palcoavi fauna of Crete; ¡he lalter also studied rhe Mcditcrrancan are, rhercfore, excluded: ¡hose of lhe Plio­ rhar of Karpath os (Weesie, 1984). ccne-Quaternary boundary of Mallorca and Menorca (Mourer­ The fossil bi rds of Corsica bave been studie d by Lydckkcr Chauviré el al., 1977; Alcover et al., 1981), a Pliocene fauna of (l891b), Newton (1921), and Mourer-Chauviré and Wecsie Eiv issa (Akover er al., 1981; Ako\'t"r and Agusri, 1985; and ne ..... (1986). The latter also worked on the fossil birds of , unpublishcd fi ndillgs), the pa1eofaunas o f rhe Pliocenc islands as did Wagner (1833), Giebe! (1847), De la MaTl1lOra (1857), of Gargano (Ballmann, 1973, 1976), and rhe pa leofaunas of Ihe Malaresla and Sc ttepassi (1954), Malares!a and Suriano (1970), Miocenc islands of Tuscany (Ponis, 1884, 1889). and Mourer-Chauviré (in Ulzeg.1 and Our, 1980). Also, ¡his study is conlined to paleoavifaunas of those isla nds The fossi l birds o f Malta ha ve been studied by Parker (18 65, thar were not connected 10 the cominenl during Ihe 1lliddle 1869), Ad ams (1877), Lydekker (1890, 189b ), Bare (1916a, 1920), andj or upper PleislOcene. This inclunc (for 1964) have bcc>n provisionaJly accepted [Pu/finus <'yermani Shu­ more informatiOIl aboUl Ihe island of Pianosa, see Azwroli, fddt 1895 becomes synonymous with Caloncctris diomedra 1978 ). Also, Ihe fossil bird bOlles from Palmaria are losl (Delle (Scopoli 1769); Cygnus cquitum Bate 1916 is prol'isio nal1y trans­ Cavt" el al., 1984), whic h makes il impossible to verify Iheir fered 10 the genus Anser Brison 1760; and Strix mclitensis Ly­ idemilicalion. The paleoavifallna of Sicily is excl uded because dek ker 1891 is rr:msfered ro the genllS Tyto l~ illberg 1828]. We it was connected 10 the main land dllring the upper P1eistocene. also follow Olson (1976) reg.l rd ing Scolopax ghardalamensis The avifauna of the middl e and upper P1eisroeene on Ihe Fi scher and Srephan 1974, which is synonymiud wi rh Cotumix Meditcrranean islands has bc c>n Ihe subiect of nu merous sludies. coturmx (Linnaeus 1758).

    274 . Akover et al. Nor inc1l1ded in Ih is smdy are rhe fln dmgs of GlI!bel (1847), rhe vici nily of frcsh water. Fun hermore, Ihe large SPCCil'S wirhin Shufc1J ¡ (1896), Newton (1921 ), Wagner (1833), and De La this o rder are nOI very att rnctive prey spceies for most avian Marmora (1857) bceause rhe t3xonomic deTe nnm:lt1ons madI' raplOrs and, Ih us, would nOI have been earried inro calles 10 be by Ihl'SC aUlhurs are in need o f revision. AIso exclllded are rhe calen. T he kat1ó r deposils we slUdied haJ a poor repres ... mation Jubious dClenninalions prescnted by Fischer and Srcphan (1974). of sorne familres of small passeriforl1ls. I:or inslanc ... , the Sy l­ T axonomic dClcnninarions al su praspecific levc1 have been con­ lIiidaC' are \'eT)' poorly represenled in Ihe record. This may be siJ ereJ only if Ihey we re madI' by rhe aUlhors of Ihe presenl because of the extreme fr:lgi lil)' of Ihei r bones. On the orher slUJy. hand, we cncountered a good representalion of medium-sized Exc1udcd l S wclllre rhe observaTlons of Lydckker (1891 b) and large spceics of rhe Passeriformes. The Carvidae, for in­ Wilh rcspccl ro Bubo sp. cL B. africalllts (Tcmm inck 1821) stancc, are \'Idl reprcscmcd in Ollf record. Their robust bones (mcruio ned as BI/bu eL cinerascells Guarin · Mancvill c 1843) and hal'c a good chanee ro fossilizc, and various species of the family ro CoraciilS sp. eL C. abyssillica Hermann 1783. Lydckkcr Corvidae frequcntly brecd and livc in rocky shclrers such as rhe (1891b) b cl i c~ed ¡hal Ihe fossi l faunas o f Ihc Mcditerranean cntranccs of calles. isl,mJ s were of African origin, bUl il has bccome d car Ihac th is Somc species of TurdrlS Linnacus 1751! are also nume ri cally is nOI tlle cas~' (e.g., Dc rmilZakis and Sondaar, 1979). We Ih ink wcll rcpresented. The)' also h,.we robust bones, and they are, Ihal in lile fi r.;¡ case Ihe fossils may represcnt Ihe recentl)· de­ IOgclher wilh sorne species of the faml1)' COI'\'ldae, regular prey scribed endcmic species Buba in,u/aris Mourcr-Chauyiré and species oí birds of prey that may have broughl thrm into the Weesie 1986, whercas in Ihe SIXond case Ihe foss ils may rep­ caves. resenl Corruillsgllrm/us Linnaeus 1758. L)'dekkt'f(l891b) like­ The orJers Aecipiuifonlll'S and Slrigifurmcs have a goO

    Medilcrrancan bland Avifaunas.Z7,5 Tahlc 1. Thc (¡(eunence of avian spcdes in lbc p;i l eo fauna ~ of the Meditcrra nea n isla nds. The followinll desillnarions llave bec n used. (al If the fossil material mar belonll 10 more Ihan ooe spe<:~s,t h c nama of these spa:ia are s.cpn .... IW with a slash (J). lb) Sorne new. undC$Cribcd forms have ,,"o ioduded. ThC$ol.' are idemified al lhe generic or Ihe familr 1C"\·cl. (e) Thc designalion "d." indiCllllcs Ihat the chara(leri,lk. obscrvcd in the fl})s ib ooi ncide wi lh IboK'" 01 Ibe lalton 10 .... bich thcy are attributw, hui Ihal ,hese charaelenslÍ() do nOl supply all Ibe inforrnalíon necesnry for pos i li~' c idcmi6cation. (d) Tbe designalion " aft." indicate5 that nOI all of Ihe characleristics 01 Ih e fl})si l bona roincide with tbose 01 Ihe laxon 10 wh ic h Ih~y are amibutcd. Tbis designalion in dicalcs only th at the lossil has sorne liitnilarily 10 Ibe narrn::d 'pccies.

    ., , • • :¡¡• o • , , ~ • , ~ u • 'E ..> .:; ~ ] ~ ~ • • ¡;:¡ ..• • E ~ ~" <3 >- ~ ~" ~ 5 ~ < >- Procdlamdae CalolllU:lrú diomedea x' x' X I'U!fiIlUS (p. ) maurclówicus X ¡'U!fillUS (p.) yelkouGn x' X Sulidac Mmu! ban ana X" l'ha].,crQ(O t:lcidac Phala(rQCorax aristotdis X' X X X X

    Ciconiidae CicO/lia cicollia X

    An;lIidae B"ml!a s angus/irostris X' Ayt/'ya {",Iig",/a X

    AccipllridJC Mi/vu5,mlvus X',d.' Mi/vus migra>!, X AcópiuT gCIl/i1is X X X Acópitcr /Ii sus X X d. Buteo lagopus d. Buteo f¡ uteo X X X Hreraetus fascilJtlls X' Aqut/a htliaca d. d. Aqurla chrysaet05 X X X X ?' X A. c. SI/Irurgh X Hallaeetl/$ alhiril/a X X X X Clrcus rIIacrQurus X' C,rcus IIeruginosu5 X CITCU5 eylJ m:IIS d.' CyfJactllS blJrbmu$ XU X At"Bypills monachus XU X Gyps {u11.'1I5 X X X G)"¡IS ."I'I,ll'nSI5 X'" X

    276. A I C<)~cr el al. Table 1. Condnued.

    , ;¡¡• • i ] " Fak·onidae Faleo peregrinus X' x X Falco subbutro X X F~/co dronorae X x X x F~/co naum~nnr X F~/co tinnrmculu$ x X X X X X F~lco sp. X Phasiallidae Coturnix cOlUrnix x X X' X x X' x Gruidat· Crus primigenia X X Crus mel,tens/S X'·' Rallidae Rallus aquaf/Cus x x l'oTz~na porzallQ X X x l'oT~alla parv~ X l'onana pusllla X Crex CTex X d. Gallinu/a ch/oropus X X F'lllCo atril X Glarrolidae G/areola pratmcolaf nordmanni X Ülitidae Ol's tardo. X X' Ot/S tetrax X X' Charadriidae Vanellus vane//us X xolopacidae Ca"dris canutU$ X d. Tringa crylhropus X Tringa fOlanus X úmosa limosa X' X Scolopax m stkola X X x x GaWnago glllli,lo.go X' X GaWnago media X Alcidae FTattTCul~ ~rt . ca x Columba palumbus X x X Columb,'l oell~S X X CoIumba liVla xu X cf.' x x x X X Co/.mlba OC>lasflivra X X' X X X X Slrcptopelra wrtur X' Slreptopelra turtu'f decaolo X Cuculidae Cu(ufu s canorus X Tytonidae Tyto alba X' X' X X Tyto ",elrte,/SI! X'M Ty/o ba/emica

    Mtd;u:, rrall~n Island Avifaunas. 2n T ablc 1. Cominued. ! , • ~ • • • ~ e • j , u .~ • ..> -;; .; E ~ ~ u• ~ • • > .. E • > ~• d ~ ~ ~ w ~ u ~ .;; ~ Slris idnc Bubo "'$ula';5 X X d .' Bubo hubo ,... Asia oIU5 X Asio f/ammeus X X X O/liS seop! X X X X,aIT. X' X X Acgolius {u"creu! X A/I"~"e " octua , ff. X X X A/hN.e cretl!"si! X X" Ketupa uglo"e"sis X Capnmulgldae Caprimulgus (UTO~ uS X X d. Apodld.le Apus apusj pllllidu$ X X X Apus me/ha X X X X Meropldae Merops IlpÍlHter X COl'3clidac CmllcillS garrulu5 X ?' Upupidac Upupa epops X Pici d.1C !ynx IQrquillll X Pleus lIitldis d. De"drocllptJS mlljor X De"drocllptJS klleotos X Alaudid.lc Calandrel/Il brlleh)'dactyla d . ,1,,1e1llllocnrypllll calandra X Gakrida eruta/a X Lul/u/a arborca d' X A/auda arwn515 X Huundlnldae "" ,ulldo rnMstris X X "",ulldo .ustlca d.' X /-/mmdo dal/rlca d. Hmmdo m stlcaj d,wnCIl X' Delicholl urbicll X

    MOI~ cil hda e MotaCllfa .:lIba X Amhus sprnolett.:l X

    l~nud;¡e t.:lIlUU collllrl0 X L"'""$millor X X X Lamlls t XCllb,lo. X TroglodYlidae Troglod)'It'J troglodrtt'J X Pnll1cll idae l'fUllel/a collaris X X X X

    278 . Alcovn CT al. TabJe L Conlinued.

    ," u

    Musckapidal' Aaocepbalus paludiwla x Sy/via alnearil/a X Syl/lia sp. x Muscicapa s/riata d. Saxlwla rube/,a x Oman/he hispanica x Oman/he sp. x Mun/ieola solilarms P/1

    (1) Lydekker, 1890; (2) Lydekker, 1891b; (3) L yd~kker, 189101; (4) B.:m, 19 1601; (5) MJ)"lud and Schaub, 1950; (6) Mabtesla and $ctlepassi, 1954; (7) Ballmann and Adrovcr, 1969; (8) Mablcsls and Suriano. 1970; (9) Fisch"r and Slq>nan, 1974; (10) Suriano. 1980; (n ) Pil-per, 1984; H pf/.."Scncc inferred.

    Mcditcrranean Island Avifaunas. 279 Tht: abst:l1ct;" of sp t;" cies of All.'(lori~ Kaup 1829 frorn [he pa­ Ihc dis[ribution of spccics [har !Oda)! are considcrcd more north­ leoavifa unas may be similarly explaint:d. Thc 6m records of ern Europtan spccics. The spccies ¡hat brcd in rhe high moun­ 0 11 Mediterranean islands coincide wilh ¡he appca r· [ai ns of ¡lit Mcdi¡erranean proba bly undcrwcnr a vertical dlS­ ance of Homo sQpicns (e.g., Cassoli ano Tagliaco"LZo, 1982). h plact:lIlem of [heir brecding zoncs (Moreau, 1955) during the is thcrdore highly probable [ffi"t¡ Ihe- preSCIl¡ diSl ribution o f glacial periods. The Medilcrrancan basin ma)· have scrved as a parlridges on Ihe Mediterranean islands is rda[t'd [O tilO: preso:nce n·ruge: fo r ¡hose spc:cies dcprivcd of suitablc habitars farther of m:lO. Recenl · o f Alrctoris have been rec- north o r in Ihe moumains. The wi lhdrawal of glacial ice from 0 " I Eur.. ""." A. I\f' ..· .. ..I""~.~·;;:;0~~·;:~~~~;~~:!~~~'::~ place in only one species. Furthennore, it has nar always been lidae. So far, fossil endemie toxa of the fami ly Corvidae havt' ¡he same genus from which rhe cndemic fonn has o riginared . been found on the islands of Mol1 orea (Co rvidae n. sp.; Alcover, Amold (1976), when swdying cases of gigantism in insular sauri a, in prep.), possi bly Eiviss.:!. [Corvus alltecorax Mourer-Chouviré suggested rhar on small islands ¡he development of one giant 1975 (n. ssp.~ ) ; Florit et al., in pressJ, and possibly Cret(' [Carru/us form would preven! rhe emergence of anorher gian! species. The glandariu5 (LinnJells 1785) (n . 551')); We('sie, 1987bJ. Of the genus 10 which the giant fo nn bc10ngs is probably dctcrmined famil ies Sylviid3e and Sinidae, no fOS$il (' nd('mie fornlS are kn own. by historical factoTS; i.e., the firs r species of samia 10 coloni"e Nevenheless, Reeem Sitta whiteheadi Sharpe 1884 from Cor-sica an island could be the one 10 monopolize rhe resources and 10 is an endemie species Ol dapred 10 li ving in rhe /'Íllu5nigra Arnold devr:lop gigantismo Something similar may have happened to the 1785 forests. The genus Sy/via Seopoli 1769 has radiared in lhe species of . Once an endemic fOflu is established on an Mediterranean area, and some of its Recent species are exclu , island and exploits i!s specific resources, it would be difficult sively inSlllar, e.g., Sylvia sarda Temminck 1820 (on Ihe is lanJs for another species to develop si mila r adaptations and oceupy of the westem Mediterranean) ond Sylvia melanolhorax Tri· th(' same niche. For this reaSOIl, endemie species of Athene Boie slam 1871 (on Cyprus). A recem find of a very robust bill from 1822 mol' have evolved on some is lands, ond endemie speeies the upper Pleistocene of Ihe island of Mallorca (wi¡h Menorca ofTyto and Buho on other isla nds. Only 011 larger is la nds might the mosl isolated island of the Medirerranean) has revealed ¡he ir be possible for vorious specie<> o f the Strigifonnes to evo ke presence of a new fringillid species, lhe only record of an en­ by eirher increasing or decreasing rh eir body si7. e as an adaptotion dcmic species of ¡he family Fring illidae from lhe Mediterranean ro feed ing on different resourees. The limired rong(' of resources islands. ayailab1c on smal1cr islands would probably only oll ow rhe evo­ lution of a single endemic strigiform species. It is imponant 10 emphasize that a panicular slrigiform species EXTINCTIONS ANO does not always change its size in the same way. FOf examp1c, DISAPPEA RA NCES FRO M T H E in ¡he genus Trio there are gia nt forms on many islands in the MEDITERRANEAN ISLA NOS world (e.g., Cuba, Gargano, Mall orca, Menorca), bUl there ore Tbe extinetions ond immigrat ions o( bird species in ¡he Medi­ also dwarf forms on other is lands (e.g" the Canar)' Islands, the renanean islands in the middlc ond upper I'leistoeene ","ere non­ Galápagos), random evems, ond extinetion did not offt'et all lhe endemies in the 5.1me way. \V irh regard 10 the faunos thot eoneern US, the Anseriformes coses of extinction andj or disappeOlronces that have token place haye affected Ihe follo\Ving groups. In the ordcr Anserifonnes, there is a species, C)"gnus (aleonen· 1. T he occllrrence and dis.:!.ppearanee of a eertain number of I'arker 1865, that has abandoned rhe aqualie habits oí its ances­ species from ¡he region was probably direerly tied ro eh3nges rors fo r a semi-terrestrial way o í li fe . I[s allolOmical adaplalions in climate. ro th is way of life have been described by Northcote (1981- 2. Large, endemic, fl ighdess species are panicularly suscep­ 1983, 1982b). It wos a large swan (approximately 16 kg). prob­ tible 10 ex¡inction after human colonization of islands (Martin ably flight!e5s, wilh a very robust hind limb and partieularly short and Klein, 1984; Olson and James, 1982, 1984). In lhe case of fcer. The remains of Amer equitum (BOle 1916) fro m Mal ta Mediterranean avifaunos, only one flight1ess taxon is known­ should be reexamined. [xc Northcote, 1988; this volume.­ Cygllus (alcolleri of lhe lI pper PleislOeene of Malta. \,(fe havc Ed.! no evidence for Ihe causes of ils extincti on, bUl human colo­ nization of the island may hove played a role. Gru iform es 3. Mosr endemie birds of pr('y probobly dt'pended on en­ Grus melitensis Lydekker 1890 wos described from Malta. This demie mammals for prey. The disl ppear:l.I1ce of th e lalter as a is a of the same size as Grus antigone (Linnaeus 1758) result of the activities of mOln andjor 10 the fauno he brought and G. primigenia Milne-Edwards 1869, and morphologieally with him mar have meant the concomitant disoppeoT:J.nce of il varies from the hmer only in a fe ...... details. Studies aTe currently raptorial birds. AIso, direct hun ting of rh e endemie speei es may under woy regarding the speeifie identity of all these forms have played a role. This is cen oin for Ihe following group. (Mourer-Chauyiré et al., 1975; Horrison and Cowles, 19n; 4. Large, non-endemic birds of prcy disappearcd from several Northcote 1982.0, 1984). 1I seems improboble [har an endemie, is la nds, e.g" lhe disappearanee of Ha/iaeetu$ a/bicilla and Ae­ vobnt sp('cies of Grus lived only on Malta. gypiU5 rnonachus (Linnaeus 1758) from Crete and Aquila chry' asd05 from Mallorca). Of the non"endemic raplOTS, ir was most­ Ral1iformes Iy the largc species ¡hat disappcarcd. According 10 Blonde! (1979), Ihe el'ological niches of birds of prcy on islands are oecupied Although endemic insular species of ral1ifonns are found in by smaller species than on rhe Illainland. This m3y be (fUe {or numerous isbnds of the world, they are unknown frOIll the the present, bU! rhis does not appear 10 apply 10 the upper Mediterranean. Usual1y, insular species of the bmily Rallidoe Plcistoeene faunos. The insular paleo fallllos of the upper P1cis­ are flightless forms, odopted 10 o mo re terrestrial way of life rocene comoined no! only speeies of birds of prey of idt'ntiul than their continent.11 coumerports. They are generally found size 3S rhose of continental fou nas, but they also contained even on mamm.1l -free isbnds al great distancc from Ihe continenl. la rger speeies, e.g., A. c, simurgh o n Crete. The reJson for lhe absence of endcmic insular species of ra ils 5. The extinction or disappearance of speeies of the bmily from [he Mediterranean moy lie in the migratory habits of mos! Corvidae hos occurred in Mallorca, Menorca, Eivissa, and Sar­ members of Ihis order. dini.1. The di5."lppeaT:J.nce of Pyrrhocorax gracu/us (Linnaeus 1766) from Malloreo and Sordinia may be attributable ro climali, I'asseriformes ehanges bee Ol use this speeies prefers o eold climate. Bec3use the Endemie fOTlns hove origi noted in at least fOUT families of the modern-day disrribution of 1'. pyrrhocorax covers o large cli­ order PaS$eriform es: Corvidae, Sylvi idae, SinidOle, ond Fringil- matie range, Ihe disappcoranee of 1'. pyrrlJOcorax from Mallo r-

    Meditcrranca n Island Avi faunas. 281 ca, M enorca, and Eivissa is probably not relatcd 10 ehanges in giformes Ihrough Acdeiformes). Bulletll1 of the Florida State Mu· Ihe climare. On Ei vissa, [he youngesr rema ins o f Corvus ante­ $("111 , 8i% glcal Sáences 7: 179-293. corax have been found in a laycr JUSI below rhe layers thar ---o 1964. Catalogue of fossil birds: Pan 2 (Anse ri fonnes tnrough produeed Ihe firs¡ record of mano Therefore, we Ih in k Ihat Calliformes). Bulletm of the f lorida State Mu,eum. Biologlcal ceologieal ehangcs eauscd by Illanllla)' exp lain Ihe disappearanee Sciences 8: 195-335. of P. pyrrhowrax frolll l he previously Illemioned islands and Caswli, P. F., and A. Tagliacozzo. 1982. La Fauna dclla Crona di Cala also ¡he extinetion o f C. antecorax on Eivissa. Genovesi a Levanw. RiVl sta di Sáenza f'reistorica 37:48-58. 6. Among marine bird s, a major ehange is rhe disappcaranee C.'l1 eri ni, F_ 191 1 Resli fossi li di uedli ddb breccia osslfera del Mome of lhe brct"ding .olonies of Puffinu~ puffinu5 rnauretrmicus from Argemario e del deposil o qU31ernario ddl'lsola di Pi anos:.. AUI ¡he island of Eivissa and o f lhe nes¡ing po pulalions of P. p. del/a Sacieta' Toscana dcgli Scicll ze Na/urali 30,75-80. ye1kor.an from Crele. The d isappearnnee o f Ihese po pulalions Cherry, J.F. ]98] Panero al1d proeess in Ihe earliesl coloniz3I ion of is probably relared ro Ihe inrroduction of differen¡ speeies o f rh e Medilerrnnean. rroceedillgs of Ihe r rehistoric Societ}" 47:4]- nummals 10 Ihese islands. Ir is wdl known rha! rats and eals 68. can h:1.\"e a disasrrOllS inA lI enee on breeding eo lonies of ¡hese De La Manll0r<1, A. ]857. Vo}'age e/I Sardaigue 0 14 descriptirm sta· species. At present, Ihe Iargesl breeding eo lo nies of P. pufflnu s ti,tjque, physique el politique de Celle i1e avce des rechcrches Sur (Brünnich 1764) in rh e M edirerranea n area are found in islc!s $t~ productions "ature/les el $(5 ,mliquit,rs, 2 .'ols. Turin: Fréres tha! are free o f mammals. Rocea, 310 pp. Delle Cave, 1.., A. Simonet1a, and A. AzzJroli. 1984. A skull of n fossi1 (Gavia) from Ihe Pliocene of Central ¡Ialy. l'alrolltogra phia LITERAro RE CIT EO Italiea, 11.5. 43:86- 93. Adams, LA. ] 87i. On gigantic 1and tortoises and a freshwa tCt spccies Dcnnitzaki s, M.O., and I'.Y. Sondaar. 1979. Tne Imponance of fossil fmm Ihe or.siferous caverns of Malu, togerher with a list of Iheir ln ammJh in rcconstrucring palcogcography wlth speclal rderenee fossil fauna and a nore on Chelonian remain s fmm Ih e rock·caviries 10 Ihe Plei510ccne Al'gcan Archipelago. A>llIales Géologlqlles des on Gibrnltar.]ournal of the Geological Society of LOll dOlr 33:177- Pa)'s He/liniques 29:808- 40. 91. DesPOII, C. 1927. Repon on the working of the Mu,eum Depanment Adrover, R. ]971. Predadores de la fauna plcistocénica de Mallorca. for /916-17. Malta. London: Bnllsh Museum (Nalural HISlOry). Boletfu de la SOCIedad de Histona Natural de Baleares 17:5- 20. Fischer, K_, and B_Slephan. 1974. Eine PleislOzanc AI·ifallna aus dcr Aleove r, J.A., and J. Agustí. ]985. Ebomys (E/IIiss;a ) umarreÍ<>ns;s n. Ghar Dabm Hshle, Malta. Zcjt,,¡"ift für Geolo8,,,he Wm:en­ gen., n. sp., nou glirid dd P1cisIOC¡' d., la Cova d~ Ca Na Reia schafte'l, Brr/ú¡ 2:5]5-23. ( Pi t iü s~). Endiu, 10-11:57-62. Florit, F., and J. A. Alcover. ]987. EIs Ocel1s del PI ~islOá, Supe ri or Aleover, j .A., and J. M umant r. 1985. El , Vertebral5 de 1"" lIIes Pi ­ de 1" Co,"" Nova (Ca pdepera, Mallorca). 1 & 11 , Bollelt de la Socicla/ tiüs ~ . Estudl' Baleanes 16: 105-16. d'Historia Nawral de les Balears 3]:8- 44, !\lcover, J.A., S. MOj·a, and J. Pon,. 1981. L"" Q uimeres del Passal. Fl ori¡, F., C. MOllrer·ChauI·iré, Jnd j.A. Akover. ]989. Els ocell, Els Vertebral5 (Ossi1, del Plio·Quatunari de les Balears i Pi li üses. pleislOcenics d'Es Pouas, Ei\"issa. NOla preliminar. Bolletí de la MOIlOgrafies Cie,ui{iques, Editorial MolI 1: 1-260. ¡uSlll uciú C atalana d"H lstona Natural 55 :35- 40. Amold, E. N. ]976. Fossi1 reptiles frolll AldrabJ Atoll, Indlan OCeJn. Ciebd, CC. ]847. V¿;gel und Amphibltl1 de, Vor",elt mI{ steter Benck· Bulletin of (he Brirish Museu,," (Natural Hislory) 29:85-] 16. 51chtlgung der lebendr Vogel und Amphlbien. Lelpzig: F.U. Brock· Azzamli, A. 1978. Possil rnammals from tne Island Planosa in nOrtnCrn halls, 217 pp. Mcdilerr.mcan Sea. BoIletillo del/a Societa' I'aleonlologica ltali· Harrison, c.J.O., and CS Cowles. 1977. Thc cxtincr large cranes of a/la ]7:] 5-27. ¡hc nonh· ..... csl l'alcarell(. jouTnal of AnthropologlCal S"ence 4: Ballmann,l'. ]973. Foss ile Vogcl aus dcm Ncogen der Halbi nsd Caro 25-27. gano (1Ialien). Scnpta Gevlogica 17:1 -75. Hsii, K.J., W ,B. Ryan, and M.B. Cn¡a. 1973, La¡e Mloccnc desiccar ion ---o 1976. Fossilc Vi>gcl aus dem Neogen der Halbmsd Cargano of Ih" Mcdilrrranl'an Sea. Natu, O' (London) 242,240-44. (l ulien), zwel1er T ell . Scripta Geologlca 38:1 - 59. Lambrechl. K. 1931. Die Fossik Omiso Spef¿jolog/S cher MO>logra· BaJlmann, 1'., and R. Adro~er . 1969. Yacimiento paleontológico de la phie", Wiell 7-8,7-9_ Cueva de Son Baus:a (Mallorca). Acta Geolúgica Hispa'liea 5:58- ]9 33. Ha"dbuch der Pa/iiornith%g,e. Ber1in: Cebrüdcr 62. Bornlrnegcr, 1024 pp. Ba le, D.M.A. 19 16a Bird bon"" from ca.'es of Malta. Geological Lydekker, R. 1890. On ¡he ~mains of some b.rge extinct birds from MilgaÚne ]9]6:332-33. ¡he cavern deposirs of ",hha. Zoological Sociel)' of Londo", p",­ 19]6b. On a slI1a ll eolleeli on of vertebrale Telllains from tne uedings 28:403- 11. Har Dabm Cavem, Malla, with a note on a new speeies of tne ]89la, Catalogue of the f055i1 bird5 i" (be British Museum genus C)"gn us. Zoological Societ}' of umdon, l'raceedhlgs19] f): (Nalllral History). London: Brirish Museum (Natuml History), 368 421 -30. pp. 1920. NOl e on a new vole and other temams from ¡hc Char ---o ]89Ib. On a Plcis!Oeene bird-Remains from the Sardinian Da lam Cavem. GeoJogical Magazine 1920:208-1l. and Corsican lslands. 20010gi,al Society of LcmdOlI, l'roceedillg5 1928. EXC3vJ¡ion of a Mousleri an Rockshclter al Devi]"s 3:467-76. Tower, Cibraltar. The animal rcmains. joumal of Ihe Royal An­ Malaresm, A., and F. Senepassi. 1954. Risultali del n lcvamcmo del thmpologlc I,lstltute 58:104-09. foglio 192 (Alghl'ro, hola di Sardcgna). Bulletino del ServiCIO Gro­ Blande! , J. ] 979. Biogeographre et á vlogie. I' ans: Masson Edi¡orial, logico I/aliano 76:33- 39. 17] pp. Ma!a¡cs¡a, A" and F. Suriano. 1970. Avifauna plelslOcenica di Alghcro --- o ] 985. H¡slorieal and ecological evidence of Ihe dt'"l·clopmem (Sardegna). Bolll'lillo del St'Tvicio G<,ologlco Ilaliauo 91:149-58. of Ine Mednerrancan avifaunas. Acta X VIlI CongreS5 Tllterna­ Marti n, 1' .5., and R.G, Kkin. 1984. Quatemary <,xtmellOns. A pre­ !lonal of Omrthology, Mosco"" 373- 86. historie revo/ution. TlIC'SOJ1' The Universi{y of Arizona Pre", 892 Brodkorb, 1'. 1963. Catalogue of fossi1 birds: Pan 1 (Ar,haeoptery- pp.

    282 . Aleonc el al. ~byaud , N., and S. xhaub. 1950. u:s puffins 5ubfossiles de Sardaigne. J'lq)(:r, H. 1984. Einr ne .. " Mesocricelus·An: (Mammalia, Cricr l iJa~) Vtrba"d/¡mgm dl."T Na/urf!don. /'rocudmgJ 125:235- Bocagialla 88: 1- 6. 95. Pons,].. and Ll. RocJ. 1974. RCilOS de grolla fOO,I .. n .. 1 "Avene de --o 1966. O n e5lim " ,~s nI ,he pasl members and of .he ave rage N" Corna" (Art.i). Elldi/lS 1:17- 19. longc\lI} of avian 'pccies. 83 :40 3- 15. Pon is. A. 1884. Contnbulloni all a orl1nologia im h.mn. Memorie del/a M O l1 n."r · C h ~u,· ire, c., anrl P.D.M. Wceslc. 1986. Bubo mfu/¡ms n.5J'., rel1l~ Accadem,a ddle Sele ll ~r d, T ur1llll 2: 13- 14. forme enJcmiquc insubire de Gra n·Oue (Aves, Slngl(orm es) du ---o 1889. Gli omnollll dd Valdarno Suprriore e di Jkuna ]ocahta' PlrislOcene de Sarda;gnc e. de Corse, Rc.·uc de Pafrolllol"gie 5: plclslocemchc di TOiCana. Alemoria ddflnstl!uto JllfNrlore d, F, 197-205. rt>J~e, 1889.1:0 pp. Mou .cr·Chauviré, c., R.• ;\drovcr, and J. I'ons. 1975. Prc<;allcc Je G," Rf'g~lia, 1'.. 1893. Sull.l buna dc lla Grona del Colombi (lsola Palmaria, ¡m/igo1le (L.) d.l ns I'A\'cne de Na Coma ii Majorqu .. (Espagne). Spn iJ). Archu'o AmhropoJogiw .. E/hllO/ogleo 2]:257-366. Nnuvdb ArchiFes d'I'¡'sfolre N ll w rel/e L yo>! l3!suppl.):50-55. 1896. Sulla €luna della G rona dl'l Colombi (Isob P3 lmariJ, MOl1rcr·Chauviré. e, S. Moyi, ;lnd R. A d r ov~." r . 1977. le, oiseJux 'pni.l). Archit,o t\mhropolugico e Ellmolug!w 26: 141- 78. des giscments qualcmaircs de Mallorca. N"IIIJ1'I/~ s A'cl!iFcs d' Hi5- ---o1 897. FO$ilh dell'I5Ob d, Tavola ... (S",dC'gna) el di Gri\'e SI. toiTe Naturefl.. Lyol1 1.5(su ppl.¡:61-&4. Alb3n (lsare). A,',eu/a 1:165-66. Mourer·(hauvlre, C., J.A. Akon' T. S. Mo}:i. ~nd J. Pons. 1980. Une ---o 1900. Sulla fauna della "BuGI dcllkrs.'1gierC''' tI sull'era' del nouvdle forme insullm.' d·Effr.IlC granle. T }'W bakanca n.sp. (Aves, dcpomi della '"KlIla "GrO Il~ dc' Colomb,·· (Isola 1'~ lm .lna, S¡x-zla). Smglformcs). du PI ,Q-Pltis,oernn" des Bal¿ares. (]«)blOs 15:803- Ar(hll'O Anlhropo/ugieo c ElI",ologico 30:1:77-8 1. 11. ---o 1907. Sulla f~una drlb Groll~ dd Cmcllo JI TrrTllIllI l mercs~, M o)' ~. S" J. Pons, ).A. Akov.. r, 'Ul d J. Agus ti. 198 4. l3 buna de 1'~lmero. Archwu AII/hmp"logiw ~ Et/lll %guo 37:3 39- 74, \'el'tcbrdos nCQgcno,cllull'nmr ia J~ Ei ' ú sa (l'illuS aS). NOla prr h­ [{ c u 01~r, ).W.F. 1982. Som" rCHurks on ¡he foss11 venebralcs from mmar. Acta Ce%gic" H ,Span;C': \9:33-35. M ~ n orca . Spain. Pmcr<'llings of the Ko"," k Ntdtrl(/l,d Akadem,e dc MUlzon, C. 1981 Les Vért~ b rh f(W;i l ~ de la 1'0rm~t1on PISCO ''l, C¡Vrus. Nature (/.,{mdnlt) 333:554- 57. Prt,(ccdillp 192 1:229- .)2, Sond.l ~r, P.Y. 1987. P I ~ l slOcenc man and eX\IIKl ion\ of island endem· Nonhcme, E.M, 1981 - 1983. Tt'lc giam Malu."St s ..... ~n. /1 Mcrill 22: ics. Mémoire5 de la SnCléré Grologlqr.e dI' Frallcc 1 '0: \ 59- 65 . 1-3. Spoor, C.F .. :ind P.Y. Sondaar. 1986_ H uman fcml ls from ¡he ende mic - --o 1982<1. The eXllnCl M ~ h~ erane Grus m~/¡t~'ws. IbIS 124: ,sbnd fauna of Sardima. ](jurnal 01 Huma/I Evo/Ilrioll 15:399- 76- 80. 40'. ---o 1982b. Siz!.", fOfOl ami habÍl of Ihc cXlinCl M311ese swan Slchhn. H.G. 1918. Übrr .. m.. ~hplcis I 01iin" ~ugCl1erfaun.l von der Cygm/S faleo"/'Ti. ¡!Ji, 124:148- 58. In'l·l PianO!01 . &Ioga~ ~ologi(il e He/L'ttI:U 1: 1:433-69. --o 1984. Cra ne Cru$ fossi ls from the Mahese PlelStOCen/.". PIl­ Su ri:mo, F. 1980. Fossil binis of Simonclh Cave. Qr.adrrm del/e ACIl. laron/o/ug)' 27:719-35. demia Naúo"'lle dd U/Ice! 249:1 23-26. --, 1988. The "xlinCI PlcislOcene " swan·goosc" C)'gn us (Amer)) Ucrpm:",n, H. P. 197 1. Die T ic rknochcnfundc ausdcr Talaloc-Siedltoll g equ,tllm from Maha. l'a/a«mtoJogy 31 :725-40. 'on ,11101 (SI. Ll o ren~, Mallorca). Str.d,e/I liber dIe T/erknochen­ NOrl hcotc. E.M., and C. Mourer-ChaU>·lre. 1988. -rhe eXll net cran .. IlInde aus lbe,ischctl I l albm~f 1::1-111. Grus pr¡''''~nia Milnc·Edwards 10 MalloKa (Sp!/': A,/¡e"e t:rek",is n. sp. ( A~es, S.ngrforrnes) from ls lands: ¡:;\'I d,'ncr for ""holl'",ll .. cxlinc.ion by m:ln before WC$lem Crele. Procudmg f 01 l/N! K,minkli¡ke Nederla"dse Akadem'" "all conlael. Scimu 117:63] - H. Wc/rnschappcn, s,e rj¡,s n 8S:JlJ-36. ---o 198 4. The rol" o( Polynes i.l l1 S in tft e exunwon of ¡he avifauna ---o 1984. On SUl1Ie r l ti~ l oce nf bird fossils from Ihe Som h Aegc dll uf Ih~ H a",,~ i i a n 1.lands. ln Q llilreNJar)' (x tmcf,ulIs,ed. I'.s. Manin lsbnd o ( Ka'lla lhos (G rCtec). Geobws 17:845-49. and R.G . Klcin, 768_80. Tueson: The Uni\'er!iilty o( ATizona Pres5. ---. 198i a. Prd lllllna!)' repon o( Ihe l'leJSloccne blrds from CTC'te. Parker, W.K. 1865 . Prd imin;uy nOles on so rne fossll b,rd. (rom Ihe DOCllmcnts du tD¡'nratoin de Geolog/t d~ L)'O" 99:197-200. Zebbug úvc, Maha. ZooIngicaf SOC /~ly 01 /J)ndon. Proceedillgs --- . 1987b. The QUJ.temary 3\'lfauna of Ü'crc, GTC'ee~. Unpub. 1865:751-53. lish"'] Ph.O. Thesls, Unil'CtS1tl' UIf"ehI. 90 pp. ---o 1869. On SOm" fcmi l bl rds from dIe Zebbug ú¡vc, Maha. Zc>oIogicaf Sode/)' nf V",dcm, T,al1$

    Mcdi.errane.l n b la nd Av i fauna~. 283 PAPERSIN AVIAN PALEONTOLOGY

    HONORING PIERCE BRODKORB

    EDlTED BY KENNETH E. CAMPBELL, JR.

    Incorporating (he Proceedings of (he n Imernational Symposium of rhe Society of Avian Paleontology and Evolution Held at the Natural History Muscum of Los Angeles County 28-30 September 1988

    NO. 36 SCIENCE SERIES NATURAL HISTORY MUSEUM OF Los ANGELES CoUNTY View publication stats