Paris Pavlakis International Institute for Human Evo l u t i o n a r y Researc h , Po rt l a n d 1

The Messinian events and the Greek fossil mammal re c o rd

Pavlakis, P., 1999 - The Messinian events and the Greek fossil mammal record - in: Reumer, J.W. F. & De Vos, J. (eds.) - EL E P H A N T S H AV E A S N O R K E L! PA P E R S I N H O N O U R O F PA U L Y. SO N D A A R - D E I N S E A 7: 233-251 [ISSN 0923-9308]. Published 10 December 1999

Greek Neogene mammal faunas, being centrally located to the stage of the Messinian play, hold clues to the understanding of this phenomenon. 23 Greek fossil mammal faunas containing 398 taxa, and spanning the period of MN Units 10 to MN15 are analysed, in order to investigate effects the Messinian Events had on Greek Neogene. Updated faunal lists are subjected to cluster statistical analyses for a quantitative estimation of chronological segregation patterns of the faunal assemblages. The Simpson's Faunal Resemblance Index was used as the variable for the faunal correlation. K-Means cluster analy- sis for two groups distinguished the 3 Samos and the 2 Pikermi faunas as the group with less variation within, than between the two groups in which the 23 faunas were separated. Join cluster analysis' basic pattern did separate the pre- from the post-Messinian faunas, but sampling error due to small faunal size, bias due to skewed sampling of the faunal diversity at the population level, and geographic proxi- m i t y, could not be factored out. Biochronological (MN Unit) range analysis of 196 species in the fau- nas showed that the majority of mammals lived in MN12-13 times, and that there was considerable reduction in numbers of large mammal species present in the faunal assemblages after the Messinian. This can be caused not only by the alleged low turn out of (large) Pliocene mammals, but also by samp- ling error due to insufficient amount of paleontological research done in this time period. Micromammals around the Messinian Stage/Age indicate the presence of variable habitats in the area, dry and wet, forested and open country before, during and after the Messinian. Greek Neogene mam- mal faunas seem to support the recent understanding that the Messinian 'Event' was lengthy, cyclic in tempo, and diverse in mode.

Correspondence: Paris Pavlakis, International Institute for Human Evolutionary Research, Central Oregon University, Portland, Oregon, USA; 1present address: Pythagora 15, Holargos, GR-15562 Athens, Greece, e-mail [email protected]

Keywords: Greece, Neogene, Mammalia, faunas, Messinian.

I N T RO D U C T I O N For over 30 years earth scientists are trying to Event. Recent data show that the Messinian describe the events that took place in the lasted for a long period of time, close to 1.9 Mediterranean Basin during the formation of My (Hilgen et al. 1997), and that it was typi- the Messinian Stage, and to study their clima- fied by cyclicity of conditions. The geologi- tic and biological effects. The international cal age for the onset of the Messinian Stage conference on biotic and climatic effects of has been calibrated to 7.26 - 7.12 My the Messinian Event in the Mediterranean, ( B e rggren et al. 1995, Vai et al. 1993). T h e o rganised in 1995 by the International beginning of the inflow from the Atlantic to Institute for Human Evolutionary Research the Mediterranean is dated to c. 5.8 My. and the Department of Earth Sciences of the Reflooding of the basin dates to 5.0 - 5.3 My University of Garyounis, held in Benghazi (Hodell et al. 1994). During this time period, Libya, reviewed the data on the Messinian 7 depositional cycles are known to have

233 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

occurred in the margins, and about 25 cycles desert environments were not widespread in the center of the basin near Sicily (Butler before 2.8 My, as indicated by deep-sea et al. 1995). This is evidenced by the sea- records of wind-blown dust off the We s t floor seismic reflectors and by the deep-sea Coast of Africa (DeMenocal & Bloemendal cores (Stanley & Wezel 1985). These cycles 1995). The Messinian was apparently not a consisted of changes in water depth and che- period of radical vegetation change, but rather mistry brought on by evaporation, influx of a time of more widespread open vegetation, new water from the Atlantic over the moderated by climatic oscillations including G i b r a l t a r, as well as by additional influx of monsoonal rains. fresh water from rivers draining into the Mediterranean Basin (Hodell et al. 1 9 8 6 ) . Zoogeography provides another clue to biotic These large rivers cut deep canyons in the conditions during the Messinian in the cir- circumference of the Mediterranean Basin, as cum-Mediterranean region (see Janis 1993). the level of the sea dropped. Geophysical The taxonomic composition of paleofloras data from northern Sahara confirm the pre- north of the basin clearly shows strong con- sence of large channels which had cut deep nections with Asia. On the other hand, the into Tortonian beds before entering the paleo- macroflora of Sahabi, Libya shows aff i n i t i e s Sirt Gulf. Some faunal elements indicate a with tropical Africa (Dechamps 1987). T h e tropical source for these rivers (Gaudant north-south latitudinal gradients implied by 1987). the paleofloral pattern, are also shown by the vertebrate faunas (see Hill 1995). Rodent fau- Global paleoclimate at the end of the nas from Asia show affinities with northern Miocene cooled, as indicated by both deep and central Spain (Berry 1995). Messinian sea and terrestrial oxygen isotopic records rodent faunas in the south, however, include (see Potts 1998). Carbon isotope data indicate the presence of dry-country gerbil species that between 8 and 6 My there was a global migrating from Africa. In Italy, there are increase of plant mass using C4 photosynthe- African primates, artiodactyls and sis, such as grassland vegetation (Cerling e t Stegotetrabelodon lybicus of Messinian age a l . 1997). Data from vertebrate paleontologi- in sites in southern Italy, while similarly aged cal sites in Greece, Libya (Sahabi), Spain and sites in northern Italy lack African elements. Abu Dhabi seem to support that the Greek Upper Neogene large mammal faunas Mio/Pliocene time was a period of greater show open vegetation environments and do a r i d i t y, and that grasslands were widespread. not indicate major faunal interchange with Yet, paleofloral records especially in the north- Africa. Rather they show connections with ern circum-Mediterranean regions indicate contemporaneous sites in Asia, such as the continued presence of warm-temperate Maraghe (Bernor et al. 1987, 1996b). T h e and high humidity vegetation well into the Abu Dhabi fauna on the Arabian Peninsula Pliocene (Velitzelos 1995). This apparent (Whybrow et al. 1999) also shows this latitu- contradiction in paleoclimate indicators can dinal correlation. It shares many faunal ele- be explained by the possible presence of alti- ments with Sahabi, but lacks close connec- tudinal and/or latitudinal gradients. Higher tions with similar age sites in the north. For elevations could maintain more humid vege- example, the anthracothere M e r y c o p o t a m u s, tation during arid phases, due to the presence the most common element in the Sahabi of cloud moisture. Similarly, northern fauna (Gaziry 1987), is present in other late European sites could have had more forest Neogene North African sites, such as in cover than sites located more southerly. Chad, but is absent from Abu Dhabi. Although the oldest paleofloral indications of desert conditions date to the Late Miocene, Floral and faunal paleobiogeography during

234 PAVLAKIS: Greek Messinian mammal record

the Messinian is an active research area, stu- taxa. They were selected based on their chro- dying such topics as how clear-cut were the nological proximity to the time period of the apparent latitudinal patterns of vegetation and deposition of Messinian Stage beds i.e., faunal provinciality during the Messinian. faunas of Late Turolian to Early Ruscinian Paleogeography of the Mediterranean basin Mammal Age, or MN11 to MN15. Second during the Messinian is also a major research criterion was the sample size. An effort was area. The land bridge connecting North made to compile the largest possible mamma- Africa and Spain during the Messinian inclu- lian faunal samples. Faunal samples with a ded dry open country and sandy habitats, as size of less than 5 taxa were excluded from indicated by the presence of African gerbils the study, for reasons of approximation to the north of Gibraltar (Moya-Sola & Agusti 1989). populations and for statistical confidence of But more vegetated conditions were probably the analyses. An effort was made to use the present as well. African ostriches, monkeys, most recently taxonomically updated faunal proboscideans among other species, reached lists. Cf., aff., question-marks (?) and quotes southern Europe while other species, such as (‘’) in taxa names were omitted. Genera were micromammals and bears migrated the other used without referring to sp. nov. , gen. (et w a y, southwards, inhabiting North Africa. sp.) nov. or names in quotes. The compiled faunal assemblages at the species level and F i n a l l y, the age of the Messinian, c. 7 to 5 the estimated age range in terms of MN Units My coincides with the estimated Ape - used in this study are listed below in alphabe- Hominid split. The effect of the Messinian tical order. ‘ E v e n t ’ on the origin and evolution of early Hominids has passed the stage of being a Ano Metochi 2,3 source of speculation (Brain 1981), and MN 13-14 (partially after Van der Meulen & reached the stage of active scientific research Van Kolfschoten 1986). (Hill 1995; WoldeGabriel et al. 1 9 9 4 ) . Amblycoptus s p ., Apodemus dominans, Paleoanthropological field research is coming Apodemus gudrunae, Hipparion s p. , close to the discovery of definitive hominid M i c romys bendai, Myomimus maritsensis, fossils in this time period. Occitanomys adroveri, Pliopetaurista dehne - li, Pliospalax s p ., Prolagus michauxi, The Greek Neogene faunal localities, being Pseudomeriones abbreviatus, Rhagapodemus centrally located to the stage of the Messinian hautimagnensis, Schizogalerix s p., Tamias s p . p l a y, hold clues to the understanding of this (de Bruijn 1989, Doukas 1989, Steffens et al. phenomenon. Specifically, to the climatologi- 1979). cal and environmental effects it had to the area, as well as, to the relation of the A p o l l a k i a Messinian to faunal and floral turnover and MN 15 (Benda et al. 1 9 7 7 ) evolution in the circum-Mediterranean region Apodemus dominans, Blarinella s p ., Castor and beyond. The major mammal faunas on f i b e r, Cerv u s a ff . philisi, Cro c i d u r a s p. , the Greek mainland and islands of Late Dolomys s p ., Episoriculus gibbero d o n , Miocene to Early Pliocene age will be stu- G a z e l l a s p., Hipparion crassum, Hyaena died here, in order to investigate aspects of p y renaica, Mimomys occitanus, Ochotonoides the effects the Messinian had on land mam- sp. (Van de Weerd et al. 1982, Benda et al. m a l s . 1977). Hipparion crassum is referred to ‘ ?P l e s i o h i p p a r i o n ’ c r a s s u m by Bernor et al. M ATERIAL AND METHODS 1996a. A total of 23 Neogene mammal faunas were used in this study consisting of a total of 398

235 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

D y t i ko 1,2,3 Megalo Emvolon - Karaburun MN 13 (Koufos 1989) MN 14-15 (Benda et al. 1979, de Bruijn A d c rocuta eximia, Anomalomys c f . ru d a b a n y - 1984, Steffens et al. 1 9 7 9 ) . ensis, Bohlinia attica, Choerolophodon pente - Dolichopithecus ruscinensis, Gazella licus, Cremohipparion matthewi, bailloudi, Nyctereutes s p ., Oryctolagus C remohipparion mediterr a n e u m , laynensis, Parabos makedonicus, C remohipparion periafricanum, Plesiohipparion longipes, Spalax odessanus, D o rcatherium puyhauberti, Gazella s p ., Sus minor, Trischizolagus maritsae ( K o u f o s H i s p a n o d o rcas orientalis, Hystrix primigenia, & Pavlides 1988, Steffens et al. 1979, De Mesopithecus pentelicus, Palaeoreas linder - Bonis et al. 1988, Bernor et al. 1 9 9 6 a ) mayeri, Palaeotragus roueni, Plesiogulo crassa, Pliocervus pentelici, Plioviverro p s K a rdia-Ptolemais 1 orbignyi, Pro s t re p s i c e ros s p., Pro t i c t i t h e r i u m MN14-15 (NOW 1995, Steininger et al. crassum, Protragelaphus theodori, 1 9 9 6 ) Tr a g o p o rta x gaudryi (de Bonis et al. 1 9 8 8 , Apodemus dominans, Castor fiber, Micro m y s Koufos 1989, NOW 1995, Andrews et al. steffensi, M. bendai, Mimomys davakosi, 1996, Sen 1996). Mesopithecus pentelicus i s Occitanomys brailloni,’?Plesiohipparion’ referred to M. pentelicus and/or M. monspes - crassum, Prolagus michauxi, Pro m i m o m y s s u l a n u s by Andrews et al. 1996. i n s u l i f e rus, Pro p o t a m o c h o e rus pro v i n c i a l i s , Rhagapodemus hautimagnensis (Koufos e t H a l my ro p o t a m o s a l ., 1988, Van der Meulen & Van Kolfschoten MN 11 (NOW 1 9 9 5 ) 1986, Bernor et al. 1996a, Fejfar & Heinrich A d c rocuta eximia, Ancylotherium pentelicum, 1 9 8 9 ) . C remohipparion mediterr a n e u m , Deinotherium giganteum, Dicero rh i n u s K a s t e l l i o s , 1 , 2 , 3 ‘orientalis’, Gazella gaudryi, Helladotherium MN 10 (Steininger et al. 1 9 9 6 ) duvernoyi, Hipparion ‘koenigswaldi’, Hystrix C r i c e t u l o d o n c f . sabadellensis, Dorc a t h e r i u m primigenia, Machairodus giganteus, Mammut s p ., Hipparion s p., Muscardinus s p ., borsoni, Metailurus major, Metailurus par - P rogonomys woelferi, P. cathalai, vulus, Microstonyx major, Palaeoreas linder - Spermophilinus c f . b re d a i (De Bruijn & mayeri, Palaeoryx pallasi, Pliocervus penteli - Zachariasse 1979). ci, Pliohyrax graecus, Pro s t re p s i c e rus ‘wood - w a rdi’, Protragelaphus skouzesi, Simocyon Limni 1,3,4 primigenius, Tr a g o p o rta x amalthea, Ursavus MN 14-15 (Van der Meulen & Va n e h re n b e rgi (Melentis 1969, Koufos 1989, Kolfschoten 1986) N O W 1 9 9 5 ) . Apodemus dominans ( 1 , 3 , 4 ) , Mimomys occi - tanus ( 3 , 4 ), Myomimus maritsensis ( 3 , 4 ) K a l l i t h i e s Occitanomys neutrum ( 1 , 3 , 4 ) , Pro m i m o m y s MN 12 (partially after Sen et al. 1 9 7 8 , i n s u l i f e ru s (1) (Van der Meulen & Va n Sondaar et al. 1 9 8 6 ) Kolfschoten 1986). Byzantinia s p ., Gazella deperdita, Gerbilus s p ., Hipparion s p ., Ictitherium orbignyi, M a r a m e n a Kowalskia s p., Machairodus aphanistus, MN13 (Fahlbush 1996, De Bruijn et al. 1 9 9 6 , Occitanomys neutrum, Paleoreas lindermaye - De Bruijn 1989) ri, Paleotragus s p ., Tr a g o c e r us amaltheus A d c rocuta eximia, Alilepus turo l e n s i s , (Sondaar et al. 1986). Apodemus gudrunae, Choerolophodon pente - licus, Hypsocricetus s p ., Dibolia dekkersi, D i c e ros heumayri, Helladotherium duvernoyi,

236 PAVLAKIS: Greek Messinian mammal record

Hylopetes macedoniensis, Kowalskia bro w n i , Parapodemus gaudryi, Plesiogulo s p ., Mustela s p ., Myomimus s p ., Occitanomys P l i o c e rvus pentelici, Pliohyrax graecus, n e u t rum, Pliopetaurista dehneli, Pro l a g u s P l i o v i v e rrops orbignyi, Promeles palaeattica, michauxi, Tr a g e l a p h u s s p . P romephitis lartetii, Pro s t re p s i c e ros ro t u n d i - ( D a x n e r-Höck 1992, Van der Meulen & Va n cornis, Pro t o r yx carolinae, Pro t r a g e l a p h u s Kolfschoten 1986, Doukas 1988, De Bruijn skouzesi, Schizogalerix moedlingensis, 1 9 8 9 ) . Simocyon primigenius, Sinictis pentelici, Sporadotragus parvidens, Stegotetrabelodon M a r i t s a grandincisivus, Tr a g o p o rta x amalthea MN14 (De Bruijn et al. 1996, Van der (Bernor et al. 1996b, Symeonidis et al. Meulen & Van Kolfschoten 1986) 1979). Apodemus dominans, A t l a n t o x e rus rh o d i u s , Calomyscus minor, Castillomys cru s a f o n t i , P i ke r m i - C h o m a t e r i Cricetus lophidens, Episoriculus gibbero d o n , MN12 (Bernor et al. 1 9 9 6 c ) Keramidomys carpathicus, Mesocricetus pri - Aceratherium s p ., Alilepus sp., Byzantinia mitivus, Myomimus maritsensis, Paraethomys pikermiensis, Chalicotherium s p ., anomalus, Pelomys europaeus, Pliospalax C h o e rolophodon pentelicus, Cre m o h i p p a r i o n sotirisi, Pseudomeriones rh o d i u s , m e d i t e rraneum, Desmanella dubia, Galerix Rhagepodemus vanderw e e rd i , moedlingensis, G. atticus, Gazella depert i t a , Spermophilinus giganteus, Tr i s c h i z o l a g u s Hystrix primigenia, Kowalskia a ff . l a v o c a t i , maritsae (De Bruijn et al. 1970, Van der Mesopithecus pentelici, Microstonyx major, Meulen & Van Kolfschoten 1986, Fejfar & M u s c a rdinus s p ., Myomimus c f . d e h m i , Heinrich 1989, Agusti 1989). Occitanomys pro v o c a t o r, Parapodemus g a u d ryi, Pliocervus pentelici, Pro l a g u s c f . P i kermi-Megalo Rema c r u s a f o n t i, Tr a g o c e rus amaltheus (Bernor e t MN12 (partially after Bernor et al. 1996b and a l . 1996b, NOW 1995, De Bruijn 1976, Steininger et al. 1996) Lopez Martinez 1976, Rümke 1976, Marinos Aceratherium incisivum, A d c rocuta eximia, & Symeonidis 1972). Ancylotherium pentelicum, Bohlinia attica, Bohlinia speciosa, Ceratotherium neumayri, P rochoma 1 C h o e rolophodon pentelicus, Cre m o h i p p a r i o n M N 11 (Steininger et al. 1 9 9 6 ) m e d i t e rraneum, Cremohipparion a ff . m a t t h e - A d c rocuta eximia, Choerolophodon wi, Deinotherium giganteum, Desmanella pentelicus, Cremohipparion macedonicum, dubia, ‘Dicero rh i n u s ’ s c h l e i e r m a c h e r i , Gazella s p ., Helladotherium duvernoyi, Enhydriodon laticeps, Felis attica, Galerix Hipparion dietrichi, Ictitherium s p ., sp.,Gazella capricornis, Graecoryx valen - M i c rostonyx s p . , Nisidorcas planicornis, ciennesi, Helladotherium duvernoyi, P l i o v i v e rrops orbignyi, Pro s t re p s i c e ros Hipparion prostylum, Hipparion gettyi, zitteli, Tr a g o p o rtax ru g o s i f ro n s ( N O W 1 9 9 5 , ‘ H i p p o t h e r i u m ’ brachypus, Hyaenictis grae - Kontopoulou et al. 1992, Koufos 1989). ca, Hyaenotherium wongii, Ictitherium viver - rinum, Indarctos atticus, Lycyaena chaere t i s , P y r g o s M a c h a i rodus giganteus, Mammut borsoni, MN10/12 (NOW 1995, Andrews et al. 1 9 9 6 ) M a rte s woodwardi, Mesopithecus pentelicus, Bohlinia attica, Cervus sp., Equus cf. stehlini, M e t a i l u rus major, Metailurus parv u l u s , E. stenonis, Felis issiodorensis, Gazellospira M i c rostonyx erymanthius, Miotragoceru s t o rticornis, Graecopithecus fre y b e rg i , monacensis, Oioceros rothii, Palaeoreas lin - Helladotherium duvernoyi, ‘Jordanomys major’ , dermayeri, Palaeoryx pallasi, Palaeotragus Leptobos s p ., Tr a g o p o r tax amalthea ( Van der roueni, Palaeotragus coelophry s , Meulen & Van Kolfschoten 1986, NOW 1995).

237 ELEPHANTS HAVE A SNORKEL! DEINSEA 7 1999

R avin de la Pluie C remohipparion mediterraneum, Gazella MN10 (Steininger et al. 1 9 9 6 ) c a p r i c o r n i s / d e p e rdita, Hipparion gettyi, H. A d c rocuta eximia, Bohlinia attica, c f . p rostylum, ?Microstonyx ery m a n t h i u s , C h o e rolophodon pentelicus, Cre m o h i p p a r i o n Samotherium boissieri, Sporadotragus macedonicum, Decennatherium macedoniae, p a rv i d e n s (Bernor et al. 1 9 9 6 b ) Graecopithecus fre y b e rgi, Hippotherium pri - migenium, Mesembriacerus melentisi, Samos - White Sands Palaeotragus coelophrys, P. ro u e n i , MN12 (Bernor et al. 1 9 9 6 b ) P l i o v i v e rrops orbignyi, Progonomys catalai, A d c rocuta eximia, Bohlinia attica, P ro s t re p s i c e ros vallesiensis, Pro t i c t i t h e r i u m Ceratotherium neumayri, Chilotherium sami - g a i l l a rdi, Samotragus praecursor ( K o u f o s um, Cremohipparion nikosi, Criotherium 1989, De Bonis et al. 1988, NOW 1 9 9 5 , a rgalioides, Gazella capricornis/deperd i t a , Bernor et al. 1 9 9 6 a ) . Hyaenotherium wongii, ‘Hippotherium’ giganteum, Ictitherium viverr i n u m , R avin de Zouaves 1 ? M i c rostonyx erymanthius, Pachytragus cras - MN10 (NOW 1 9 9 5 ) sicornis, P. laticeps, P. quadricornis, A d c rocuta eximia, Choerolophodon penteli - P a l a e o reas lindermayeri, Palaeoryx pallasi, cus, Cremohipparion macedonicum, Palaeotragus coelophrys, Pseudotragus Helladotherium duvernoyi, Ictitherium s p ., capricornis, Samotherium s p ., Sporadotragus M e s e m b r i a c e r us melentisi, Ouzocerus graci - p a rvidens, Tr a g o p o r tax ru g o s i f ro n s ( B e r n o r lis, Samotragus praecursor (Koufos 1989, et al. 1 9 9 6 b ) . Bernor et al. 1 9 9 6 a ) . Samos - Main Bone Beds R avin de Zouaves 5 MN12 (Bernor et al. 1 9 9 6 b ) M N 11 (Steininger et al. 1 9 9 6 ) A d c rocuta eximia, Ancylotherium pentelicum, A d c rocuta eximia, Chasmaporthetes bonisi, Belbus beaumonti, Byzantinia hellenicus, C h o e rolophodon pentelici, Cre m o h i p p a r i o n Ceratotherium neumayri, Chilotherium macedonicum, Cremohipparion pro b o s c i d - samium, Choerolophodon pentelicus, eum, Dicero rhinus orientalis, Gazella s p ., C remohipparion matthewi, C. pro b o s c i d e u m , Helladotherium duvernoyi, Hipparion dietri - Criotherium argalioides, Deinotherium chi, Hyaenotherium wongii, Ictitherium viver - giganteum, ‘Dicero rh i n u s ’ s c h l e i e r m a c h e r i , rinum, Mammut borsoni, Mesopithecus pente - Felis attica, Gazella capricornis/deperd i t a , licus, Microstonyx s p ., Nisidorcas planicor - G r a e c o r yx valenciennesi, Helledotherium nis, Palaeoreas zouavei, Postpotamochoeru s duvernoyi, Hipparion dietrichi, hyotherioides, Pro s t re p s i c e ros ro t u n d i c o r n i s , Hyaenotherium wongii, Hystrix primigenia, P ro s t re p s i c e ros zitteli, Tr a g o p o r tax ru g o s i - Ictitherium viverrinum, Indarctos atticus, f rons, Zygolophodon tapiro i d e s (Andrews e t Lycyaena chaeretis, Machairodus giganteus, a l . 1996, Werdelin & Solounias 1996, Bernor Mammut borsoni, Metailurus parv u l u s , et al. 1996a, Koufos 1989). M i c rostonyx erymanthius, Miotragoceru s monacensis, Muntiacus s p ., Occitanomys Rema Marmara p ro v o c a t o r, Oioceros wegneri, Ory c t e ro p u s MN13/14 (Benda & Meulenkamp 1990) g a u d ryi, Pachytragus crassicornis, P. M i c romys bendai, Occitanomys neutrum, O. laticeps, P. houtumschindleri, Palaeoryx pal - brailloni, Parapodemus gaudryi, Pro l a g u s lasi, Paleotragus rounei, Parataxidea pola - michauxi (de Bruijn 1989) ki/maraghana, Pliocervus pentelici, Pliohyrax graecus, Pliospalax sotirisi, Samos - Old Mill Beds P l i o v i v e rrops orbignyi, Promeles paleatica, M N 11 (Bernor et al. 1 9 9 6 b ) P romephitis lartetii, Pro s t re p s i c e ros

238 PAVLAKIS: Greek Messinian mammal record

rotundicornis, Protragelaphus skouzesi, with the fossil faunal assemblages in this Pseudomeriones pythagorasi, Pseudotragus s t u d y. SRI accomplishes this by dividing the capricornis, Samokeros minotauru s , number of common taxa between two faunas, Samonycteris majori, Samotherium s p ., by the number of the smaller of the two Schizogalerix atticus, Spermophilinus bre d a i , samples, thus maximizing similarities Stegotetrabelodon grandincisivus, between the faunas than differences, as is the Tr a g o p o rtax amalthea, T. curvicornis, T. case with, among others, the Jaccard formula. ru g o s i f rons, Ursavus depereti (Bernor et al. [For further discussion on fossil mammal 1 9 9 6 b ) . faunal correlation coefficients, see Shuey e t a l . (1978) and Bernor & Pavlakis (1987)]. Va t hy l a k ko s The statistical program Systat ver. 7 for MN12/13 (NOW 1995) Windows was used for the calculation and Bohlinia attica, Ceratotherium neumayri, production of the faunal correlation matrix, C h o e rolophodon pentelici, Cre m o h i p p a r i o n as well as for the subsequent cluster analysis. macedonicum, Dorcatherium puyhaubert i , The Join or hierarchical method of cluster Gazella s p ., Hipparion dietrichi, analysis was used, in which the set of faunas Hyaenotherium wongii, Ictitherium viverr i - is partitioned into sets of nested groups, with num, Microstonyx major, Mesopithecus pente - the most similar faunas nested first and closer licus, Nisidorcas planicornis, Palaeoreas lin - (for details see Sokal & Sneath 1963). T h e dermayeri, Plessiogulo crassa, Plioviverro p s output of the hierarchical method is a tree or orbignyi, Pro s t re p s i c e ros zitteli, Samotherium dendrogram. boissieri, Tr a g o p o rta x ru g o s i f ro n s (Bernor e t a l . 1996a, Koufos 1989, Werdelin et al. The pair cross correlation of SRI's was com- 1 9 9 6 ) . puted as ‘distances’ between two faunas. These distances are normalised to allow com- The faunas were subjected to the multivariate parison of clustering across widely diff e r e n t statistics of cluster analysis for detecting SRI values. These pair cross correlation were natural groupings in the data. The primary produced by 4-distance metrics: Eucledean, goal of this test is the estimation of the 1-Gamma coefficient, 1-Pearson correlation degree these Greek Neogene mammal faunas c o e fficient, and Percent disagreement. T h e are separated into a Pre-Messinian and a Euclidean distance uses the square root of the Post-Messinian group. It is a gross quantitati- SRI correlation value of a pair of faunas ve estimation of how different the faunas are (mean for more than one faunas in clusters). before and after the Messinian. First, a sym- The Gamma option uses 1-g, where g is the metrical matrix of correlation coeff i c i e n t s Goodman-Kruskal gamma correlation coeff i - between all 23 fossil mammal faunal assem- cient between two SRI's (pairs of faunas). blages was computed. Simpson's formula for The Pearson distance is calculated as 1-p, Faunal Resemblance Index (SRI, Simpson where p is the Pearson product-moment cor- 1960), was used to produce this input data relation between two SRI values. T h e matrix for the cluster analysis. The SRI for- Percentage produces a distance index, which mula is: C/Nm i n, where C is the number of is the percentage of the difference of two SRI common taxa between two faunas, and Nm i n values. For more information see Hartigan the size of the smaller of the two faunas. (1975) and Sokal & Sneath (1963). Cheetham & Hazel (1969) compared the SRI correlation formula with several others, and For each one of the four-distance metrics, the a rgu ed that it showed greater consistency following linkage clustering methods were when the size of the faunas vary considera- used to compute the distance of a cluster to b l y. Large sample variation is indeed the case a n o t h e r, in order to decide whether the two

239 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

should be merged, in a given step: fauna) and the three Samos faunas (Main O Single: the distance between the two Bone Beds, Old Mill Beds and White Sands). closest members in separate clusters is The other group contains the rest of the fau- taken as the distance between the two nas. This first result brings forward the clusters. It produces long, stringy clusters. Pikermi and Samos faunas as the reference O Complete: it uses the most distant pair of Greek Neogene fossil mammal faunas. Main faunas in two clusters to compute reason for this is the large size of the faunal between-cluster distances. It tends to samples. The Pikermi fauna contains 57 taxa, produce compact clusters. and the Samos (Main Bone Beds) 58, while O Centroid: it uses the average distance the next numerous faunal sample has only 23 value of all faunas in a cluster as a taxa (Halmyropotamos). An additional reason reference point for distances to other for this separation is related to taxonomic c l u s t e r s . identification. Both faunas are the best stu- O Average: it averages all distances between died and the most recently taxonomically pairs in two clusters to decide how far updated. Adequate sample size, which apart these clusters are. approximates with statistical confidence the O Median: it calculates the median of all extinct animal population, and updated taxo- distances between pairs in two clusters to nomic identification of the faunal sample, are decide how far apart these clusters are. prime requirements for reconstruction of the O F i n a l l y, the merging of clusters is decided paleontological record, biochronological or by the Ward method (Ward 1963), i.e., by paleoecological, including that of the Greek producing clusters with minimum N e o g e n e . v a r i a n c e . The Join cluster analysis produced 26 den- The 23 faunas were also subjected to the K- drograms, as described in the methods. In 21 means clustering method. It assigned faunas of these diagrams three major factors were to non-overlapping clusters. Two groups of detected to have been responsible for the pro- faunas were selected for this study. It split the duction of clusters: faunal diversity and size, faunas into two groups by maximizing geography and age of the faunas. Only two between clusters variation, relative to the main faunal groups were almost constantly variation within the two clusters (for details produced. The main pattern is shown by the see Hartigan 1975). The faunal assemblages dendrogram of Figure 1. It is produced by the were not standardised or ranked in both Join Euclidean distance and the minimum variance and K-Means statistics. 50 iterations were ( Ward) clustering method. This clustering done for the separation into the two clusters. pattern will be described, as the basis for dis- The 23 mammal Neogene faunas were further cussing three variations produced by the analyzed in biochronological and paleoecolo- other dendrograms. One major grouping con- gical context. Specifically, the biochronologi- tains three nested clusters. Ravin, Va t h y l a k o s cal range of selected taxa was tabulated to and Prochoma faunas comprise the one clus- show possible faunal turnover around t e r, and Samos and Pikermi faunas belong to Messinian times. another cluster. Dytiko and Halmyropotamos are also part of this group. The sister group R E S U LTS AND DISCUSSION consists of the rest of the faunas in the study. The K-means cluster analysis (less SRI varia- The faunas of Ano Metochi, Ptolemais and tion within clusters than between clusters), Rema Marmara are nested first, and next produced the following two faunal groups: cluster with Limni and Maritsa, with the the first one consists of the Pikermi fauna Apollakia fauna outlayer. A d d i t i o n a l l y, from Megalo Rema (the main, classical Megalo Emvolo, Maramena, Pyrg o s ,

240 PAVLAKIS: Greek Messinian mammal record

Figure 1 Cluster analysis tree diagram of Greek Neogene faunas studied. Distance metric is Euclidean distance.Ward minimu m distance method.

Kastellios and Kallithies cluster together with- ring. Only Dytiko is younger than the other in this group. faunas. It belongs to zone MN11 - 1 2 .

The former of the two major clusters is the The second group of faunas is also solid, in solid set of Pikermi-Samos-Axios Va l l e y, terms that, with the exception of a couple of l a rge size, taxonomically diverse, Late faunas, all cluster runs produce it. This grou- Miocene, Upper Tortonian-Messinian Stage, ping is the result of the high proportion of Late Turolian Age, MN 11-12, Greek main- micromammals in the faunal samples. Time is land Neogene faunas. The following factors also a decisive clustering factor. The faunas were employed, in order of importance: high which belong to this cluster belong mostly to faunal diversity and large size of the faunal zone MN13 or younger. Only the set of sample are decisive factors. They make the Kastellios and Kallithies faunas are older. sample approximate the population and make Geographic proximity is the next clustering the statistical analysis reliable. Time and factor (Strimon Basin - Ano Metochi, geographical proximity are additional factors Maramena, Rema Marmara; Ptolemais; that played key role in producing this cluste- Thessaloniki - Megalo Emvolo; Rhodos).

241 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Figure 2 Cluster analysis tree diagram of Greek Neogene faunas studied. Distance metric is 1-Gamma coeffi c i e n t . Ave rage linkage m e t h o d .

P y rgos interchanges groups as outlayer in the the two pre- and post-Messinian basic faunal other runs. It is not an adequately sampled groupings are still distinct, the problematic and studied faunal assemblage. In Figure 2 faunas of the three previous patterns are out- we have the same division into two groups, layers to both major groupings. These five with the Pyrgos fauna clustering with the faunas are the oldest Kastellios (MN10), the M N 11/12 faunas. In this dendrogram, P y rgos fauna of uncertain age (MN10-12), Maramena fauna is an outlayer to the early and the Maramena, Kallithies and Megalo age group. In Figure 3 we have the clearest Emvolo faunas. In Figure 4, therefore, only separation of the faunas into a group dating the Dytiko fauna violates the clear-cut divi- less than or equal to MN 13 (pre-Messinian) sion of the 23 (or 18) faunas into a pre- and a group younger than MN13 (post- Messinian and a post-Messinian group. Messinian). The only exceptions are the Geographic proximity with the rest of the Kastellios fauna, which clusters with the Axios Valley faunas played the decisive role young faunas, and the Dytiko fauna, which in the Dytiko clustering. clusters with the older. Finally, in Figure 4 we have a slightly different cluster arrange- Biased sampling of the true animal diversity ment, and perhaps the most realistic. W h i l e in the extinct population, is a possible cause

242 PAVLAKIS: Greek Messinian mammal record

Figure 3 Cluster analysis tree diagram of Greek Neogene faunas studied. Distance metric is 1-Pe a r son correlation coeffi c i e n t . Complete linkage method.

of sampling error; an ever-present problem in and not a species FAD and LAD in the fossil p a l e o b i o l o g y. Sampling error cannot be record. Filling-in was used if a taxon was excluded in the paleofaunas used in this clus- present in the one MN Unit before and the ter analysis. Beyond this factor, geographic one after the unit where the taxon was mis- and age proximity are the two factors causing sing. The majority of the bovids in the 23 the two major faunal clusters. In order to far- faunas belong to MN 12. The biochronologi- t her investigate the effects the Messinian had cal distribution of equids and rhinos, as well on Greek mammal faunas, biochronological as that of carnivores, range mostly between ranges of specific species of major mamma- MN 11 and 13. There is an apparent conside- lian orders have been constructed, with the rable number of species reduction at the onset MN Unit biochronological system as a refe- of the Pliocene. That the environment changed rence scale. Figures 5 to 11 tabulate the MN around the Messinian, 7-4 My ago, and that biochronological range of 196 species, pre- this change was caused by a trend for colder sent in the 23 Greek mammal Neogene fau- and dryer climate worldwide, has been suff i- nas. This is a recording of the presence of ciently documented. Potts (1998) provides a each species in the MN Unit range 10 to 15 complete treatment of recent data used in

243 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Figure 4 Cluster analysis tree diagram of Greek Neogene faunas studied. Distance metric is 1-Pe a r son correlation coeffi c i e n t . Median linkage method.

paleoenvironmental reconstruction. F i n a l l y, looking at the micromammals (Figs. Environments, however, always change. A n d , 5-7), their record in the Greek Neogene is however low is the turn-out of mammals in better than that of the large mammals. the Pliocene, certainly there were mammals M o r e o v e r, small mammals are environmental around. No study, to my knowledge, equates indicators. The disadvantage is that they can the effects of the Messinian ‘Event’ with the only tell the environment in a small area. T h e major effects of the bolide impact at the end record of Insectivora is mostly pre- of Cretaceous. The large-mammal distribu- Messinian. B l a r i n e l l a and C ro c i d u r a have a tion is shown in Figures 8-11, with many spe- late presence in Apollakia. Squirrels of all cies not making it after the Messinian, and three types are present (flying, tree and few new ones appearing. In addition to the ground). The flying squirrel H y l o p e t e s is pre- known environmental changes after the sent in Maramena (MN13), and Ta m i a s i s Messinian mentioned above, this is also due present in ? post-Messinian Ano Metochi to sampling error and to poor taxonomic (MN13-14). Most of the Cricetidae and identification for some faunal assemblages Muridae do not generally support a single collected long ago. widespread dry or wet environment after the

244 PAVLAKIS: Greek Messinian mammal record

Figure 5 Biochronological range in MN Units of specific Insectivo ra , L a g o m o rp h a , P rimates and Proboscidea present in the studied Greek Neogene faunas.

Figure 6 Biochronological range in MN Units of specific Cricetidae (Rodentia) present in the studied Greek Neogene faunas.

Messinian. Instead, the rodents tell us that believed. It is now known that it was not a there were trees, as well as open country, dry single and short-lived event. The effects, the- sandy ground, wet, as well as dry environ- refore, to the mammalian faunas must also be ments in the area before, during, and after the far more complex than a direct simple eff e c t Messinian. This is supported also by paleo- on them by a unique, unidirectional trend of botanical studies in the Greek mainland environmental change. More scientific data ( Velitzelos 1995). It is obvious today (due to out of fieldwork and a study of paleontologi- the multidisciplinary research done) that the cal museum collections work will further Messinian effect on the circum-Mediterranean explain the Messinian phenomenon and its area is more complex than was previously effects on the Greek Neogene mammal faunas.

245 ELEPHANTS HAVE A SNORKEL! DEINSEA 7, 1999

Figure 7 Biochronological range in MN Units of specific Muridae (Rodentia) present in the studied Greek Neogene faunas.

Figure 8 Biochronological range in MN Units of specific Carn i vo ra present in the studied Greek Neogene faunas.

Figure 9 Biochronological range in MN Units of specific Pe rissodactyla present in the studied Greek Neogene faunas.

246 PAVLAKIS: Greek Messinian mammal record

Figure 10 Biochronological range in MN Units of specific A r tiodactyla present in the studied Greek Neogene faunas.

Figure 11 Biochronological range in MN Units of specific Bovidae present in the studied Greek Neogene faunas.

AC K N OW L E D G E M E N T S I thank the editors of this volume for inviting Event on the circum Mediterranean. T h e this contribution. I thank dr M. Dermitzakis o rganisers, Dr N. Boaz, International Institute and dr E. Velitzelos, University of A t h e n s , for Human Evolutionary Research, and A. El- Greece, for reviewing the manuscript. Dr N. Arnaouti, University of Garyounis, Benghazi, Symeonidis is thanked for discussions during Libya are thanked for inviting me at this early stages of this study. Dr M. Fortelius, meeting. Pant. Pavlakis provided computer University of Helsinki, is thanked for off e r i n g support for the analysis. Last, but by no me a 1995 version of the NOW database. A means least, I thank Dr Paul Sondaar for his preliminary version of this study was presen- life long interest and contributions to the ted in the International Conference on the paleontology of the Mediterranean region, biotic and climatic effects of the Messinian including Greece.

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