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Home , Agathonisi, Agios Efstratios, Alexandroupoli, Alimia, Alonnisos, Ananes

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Acknowledgements Introduction

General of

Greece is a relatively small country, and with a surface area of 132,000 km2 it is only half as big as the UK. Encompassed, however, in this modest area, is a great diversity of habitats, exceeding many European countries of much larger size. For example, one can encounter in alpine areas complete with lush conifer forests, dramatic peaks and extensive snowfields that physiographically resemble Switzerland. On the other hand, some regions of the southern Aegean are closer to Africa than to , and their climate and habitats reflect this proximity. Southeastern for example, con- tains one of the few true European deserts, an area closely resembling certain hamma- da regions of the Middle East. Greece is a country of mountains and . The Pindos range, an extension of the Dinaric Alps, forms the backbone of peninsular Greece. A number of smaller mountains originate as spurs from this block, although some, including , the highest mountain in Greece (2,917 m elevation) arise in relative isola- tion. A second major mountain block, the Rhodopes, located in , runs in a roughly east-west direction separating Greece and . The , a small- er peninsula in the south, is as mountainous as the mainland and encompasses several peaks exceeding 2,000 m in elevation. With the exception of a few large flat regions located mostly in and Thrace, the country lacks extensive plains. Typically the mountains drop rather steeply into the sea and are generally flanked only by narrow coastal plains. While this has his- torically translated into meager agricultural outputs, it also means that large areas of the country are, by virtue of their steep and mountainous terrain, relatively protected from human activities. To visitors, Greece conjures up images of islands and there is a good reason for this. The country encompasses close to 10,000 islands and islets, the vast majority of which are found in the . However, due to difficult transportation and the harsh conditions, only about 200 of them are inhabited. These islands vary significantly in physical and biological characteristics so that visiting a number of them may be neces- sary to obtain a good understanding of the region.

Climate

The climate in Greece is broadly classified as Mediterranean, with mild, rainy winters and long, dry summers. Typically the bulk of the annual precipitation, in the form of 12 Introduction Introduction 13

[a{ak Kraljevo SERBIA BOSNIA-HERZEGOVINA a Priπtina Kruπevak BOSNIA-H. MONTENEGRO YUGOSLAVIA D r n 1913 S k≤ i Bartin Troglav SARAJEVO * Montana Is nLovech YUGOSLAVIA KOSOVO Vratsa s Pirot t a J. a Gabrovo BLACK SEA Zonguldak Ard a YUGOSLAVIAM a l2376 Botev CROAT-MUSLIM FED. Gora≠de r P E o a YUGOSLAVIA v european part Niπ * r r a o s 2228 s ∏ibenik SERBIA v Karabük BULGARIA u l Plo‹no a SOFIYA Kazanl≤k Podgorica r SERB REP. OF Novi Pazar Leskovac o m B s p Split BOSNIA-H. T s Mostar a (Sofia) u Bo ra BULGARIA r CROATIA a 2386 2522 o Echinos Magli‹ Durmitor p SEA OF 2155 s 1511 Plovdiv Dimitrovgrad Tykhero CROATIABra{ t o Kato Pogled 1827Iasmos Pazardzhik Kirklareli Düzce Nevrokopi MARMARA B Skopje estos i a Edirne Bolu N Hvar * Kyustendil Asenovgrad Khaskovo * Xylagani a MONTENEGROPriπtina 2925 2378 Sidirokastro 2484 Pe‹ Musala M F.Y.R.O.M 2212 2232 Podgorica Sliven Agios * a Kor{ula Komovi Köro‘lu Da‘i Charalampos 2656 KOSOVO Blagoevgrad r Lüleburgaz Izmit * Nea Karvali l i Adapazari ANATOLIKI MAKEDONIA KAI THRAKI R t s Former Yugoslav Republic ]aravica K≤rdzhali s Yambol Skoutari s k h a e n s a Feres Rodolivos of o o Nea Peramos r v Dubrovnik 2693 d Istanbul E Prizren S o Tekirda‘ Yalova Xylopoli Mikropoli 2914 p LRodopoli Dojran L Kerkinitis rdanell Jezercë Kumanovo t e Stara Zagora s r Vikhren o a u Aridaia Stri D mo 1956 g m SEA OF nas a Tetovo SKOPJE a Bursa A l Kukës V Ke≈an Bilecik x Ulcinj Mudanya Edessa i a Pernik o Nigrita 1127 e 2748 r Iznit Lake ADRIATIC s Samothraki da MARMARA Arnissa 2029 Mavrothalassa Thasos Shkodër Titov r Komotini Sariyer Polatli 1963 P a Platy 1091 L Volvi Palagru≠a 2764 a Bozüyükkary Polykastro KENTRIKI MAKEDONIA 1611 i s Sa Naousa L Koronia Asprovalta Limenaria o Croatia Lake Kourab Xanthi SEA Amyntaio Kolpos n le Karacabey Nea Santa Lagkadas Scutari Drama l Eski≈ehir Sivrihisar Stavros k Peshkopi o Alexandroupoli e Lake Vergina Nea Orfanou Lezhë n L Vegoritis Giannitsa KalamariaEvosmo i (Former Yugoslav Republic of Macedonia) a Lake Ptolemaïda Prespa Gerakarou d d Korinos 1165 k Serres Laker Ku≈ Maniakoi Veroia 2524 Prilep Kavala a Ulubat 2543 PolygyrosThessaloniki(Salonica) ADRIATIC F.Y.R.O.M Eceabat Argos Orestiko a e ÇanakkaleD Uluda‘ NeaAGIONOROS Roda r Galatini 2334 (Mont Athos) L KastoriasVelvendos c Samothraki Siatista Servia Kosani Leptokarya h 430 SEA TIRANË (Tirana) Bitola Sfendami A K Samothraki Platamonas g ol Akra T 2052 s p a a uOro Akrathos n Sikeas Durrës Thasos L Aliakmonas Koufaliao Kütahya m Portoou Karras Akra k K s 2030 Kalamaria Edremit a K o Deskati li a l MourtzeflosAthos Elbasan 2194 s po M A s s Veroia Thermaïkos an Alexandreia dr Arnaia Akra Agia Lake Ohrid Polygyros as Foggia LakeKorçë Ak≈ehir Verdikoussa Kolpos Eirinis (Salonica) Balikesir 2111 Vasilika Galatista Gökçeada 2520 Kalampaka Aiginio Akra Drepano Limnos Plaka Kosani Prespa 2030 Mesimeri Akra Katerini Athos S Nikaia Akra 2917 Paliouri S t r Farkadhon Kassandras Bari t Larisa a Olympos r Vlorë i DYTIKI MAKEDONIA Nea Moudania Moudros 968 t Neos Katsikas s TURKEY a Rigaio Agios Myrina o IoanninaPramanta 2637 Nea Anchialos Efstratios i ALBANIA Liapades TrikalaKarditsa Kassandreia e t Kula f Smolikas Koufos Akra 1615 d 800 Limnos Dinar O Kallifoni 680 o Ak≈ehir Ao 1978 Paliouri os Neo MonastiriC a ) t h a s i a r Giouras f Lake Kerkyra r 2497 e Akra Brindisi 2917 Perdika Thesprotiko Larisa o 792 Lefkimmi a 1054 d Maleas Mt Olympus Kanallaki P Pelasgia Pyrgetos p a Mytilini Matera O 2637 Afyon n S r Potenza Lecce 2485 Filippiada Petas o Lesvos Smolikas i p t Papingut Volos Northern (Mytilène) t io S Sigri Igoumenitsa o e rn () r U≈ak Burdur Agia 1551 or e P Izmir Nazilli Ereikoussa Krikellos V th Akra Kartsino a s STEREA ELLAS or Akra E‘ridirlake eio Kerkyra z i Pin (N Taranto i TURKEY is Vamvakas n Mytilini Vonitsa Amfilochia ed Lake Bey≈ehir Lake m i G Mathraki a Kato TithoreaTHESSALIA Pagasitikos VOREIO AIGAIO Çe≈me Denizli y t Lesbos h T A StefanovikiKolpos Manisa Lefkada Palairos r o a Davlia n Chios c n h Levadeia () t A Kandila h c Skyros Izmir ITALY o h Chalkida Akra Sarakiniko Thermo

2204 s e AkraValaxa Kymis Agrinio l Itea 2980 o Argalasti Skopelos d Isparta o 906 2469 s L Trichonida 530 2248 Delegöl Da‘i Kerkyra IPEIROS 1297 Gulf of Chalkida d 1726 Akra Meston Mt Pollino u Milâs Antalya (Corfu) Acigöl Burdur 1974 2393 2184 V Akra Masticho I (Corfu) 1924 Taranto s Volissos s Ithaki 1589 o Skyros a p o o r Chios l v Loutra s I 2042 or o Peiraias Paxoi 1158 a Spercheio V eios Akra Kafireas Corfu t Sami t T s AidipsouE 1343 1743 Kymi s Dalaman o Molos vv Aydin re Argostoli Antipaxoi oï Oxylithos M Lamia Chios e Lixouri u ko d M Amvrakikos Kolpos 2315 s Akra Fanari en IONIOI NISOI( s Agios Konstantinos K Ochthonia Akra Kampanos Pyrgi n M 2152 Kifisos Malesinaolp o os t Korinthos ük Oiti Martino 1434 Crotone Agios Dimitrios Büy Bozburun s Korinthos AliveriEvvoia (Euboea) s I 2510 Drosia Krieza Catanzaro i u Giona OrchomenosAkraifnio Samos 2457 IONIAN Almyropotamos Samos Parnassos r Katastari Amfissa 2457 2341 Schimatari Agioi Neo Karlovasi AEGEAN SEA o Parnassos2221 ApostoloiPetaliori Akra Livada a n Zakynthos 1627 Erymanthos Thiva AEGEAN SEA Akra Papas Marathokampos Lefkada Marathonas 1398 1033 Mu‘la u 1748 Erythres

SEA Mataragkaös Leucade Astakos Kapareli 1413 Nea Makri

o Panaitolio Platanistos

l Nafpaktos 1409 e n Pyrgos n (Leucade) Kifisia

a Aitoliko h

c s Agios

o Mandra Kolpos Gavrio 994 Gulf of A n Kamares i r Nea Liosia Andros Christos Kirykos Cosenza DYTIKI ELLAS Mesolongi o 1981 Petalion Andros T 756 Aigio TYRRHENIAN Keffalonia 3086 M Korint Xylokastro Megara Koropi Antalya i hiako SalaminaZografos Ikaria MakronisiAgathonisi 678 sKo Kiato 650 Patra Bey Da‘lari pos Peristerio Keratea ATHINA (Athens) s Loutraki Aigina Vrachnaïka Agioi Theodoroi SEA I o o 1419 Agios Voula a Akra Doukato Kato Achaïa olp Zevgolatio Tinos Patraïkos K Patra GeorgiosATHINALavrio(Athens) ErmoupoliTinos 2374 Agios DimitriosPalaia s GREECED Ithaki Poros Fokaia Gyaros Kyllini Akra Dafnoudi Varda Messina Akra Katakolo Kleitoria SalaminaSofiko n Nemea Chiliomodi AkraVoudi i Loutra l Tripoli o AkraLechaina Araxos Kandila Poseidonia D () Peiraias Gastouni Argos Lygourio Kallithea Mykonos a Kefallonia Poros Kanala Zante Peloponnese Tropaia ( d Levidi Kea s Tripoli Naxos Kalymnos o Platanos Donoussa o I Akra Mounta Paros D im Kefalos n r e Krestena Ydra e 846 N Kranidi Naxos 690 s Trianta Megisti PeloponnisosAstros d P c Rodos () s Akra Skinari 2374 d Zacharo MegalopoliKyllini Kamares o Greece PELOPONNISOS Kamiros Afantou Irakleio Kato Agios Petros Spetses Paros 735 Catania a Pyrgos Figaleia A Akra e Kos s l lfei 1001 Archangelos os 1199 ATTIKI 821 d i A Leonidi Ierax 771 (Rhodes) AkraEmponas Rodos Reggio di Calabria n Zakynthos r Serifos 751 Akra Paraspori Megisti Rhodes a g Kyparissia K o Irakleia k Meligalas o l AmorgosSikinos e 698 Lindos e l ik Ydra s (Zante) Arfara p 1113 Akra Marathia P Sparti o o 566 Vatio n Filiatra s s s Kyparissiakos Geraki a a c 3323 Strait 2404 Milos e GargalianoiKolpos Akra Maleas Anydro d o Chora M t Etna Tavgetos s r KykladesOfidoussa a n Attavyros Kattavia Petalidi of n Skala Molaoi 1215 Thira s T Arna Rodos (Rhodes) i a o Emporeio n i Chalki Ragusa Longa Papadianika MirtoöMonemvasia Ios Makra Messina Akra vTainaro n ) g Zafora s Methoni KoroniKalamata Velopoula e Strofades 1224 e Areopoli o t Evrota Karavonisia Akra KastellouDiafani o s Akra KapelloNeapoli s Pélagos Folegandros Astakida s s Milos Anafi e Proti 2404 ) Chamili Karavas Thira Divounia Akra Vathia Akra Agios Messiniakos Potamos Karpathos Siracusa Kolpos Akra Akritas Akra Spatha Ioannis National Other Avlemonas NOTIO AIGAIO Sapientza (Cyclades) Lakonikos Kythira Capital city Schiza Elafonisos Kolpos 507 Irakleio Antikythira Karpathos over 1,000,000 SEA OF CRETE Nea 1215 Siteia Palaikastro Alikarnassos Limin Chersonisou Kythira Malia Neapoli Zakros , Rodopos over 500 000 Akra Lithino Kastelli 600 Perama Anogeia Kastelli Arkalochori Kalo Chorio 2456 Krousonas Psiloritis over 200,000 Kournas Ano ViannosKasos 2147 National Capital (3,093,000 In 93) 2456 Elos Psiloritis Chania Asimi other main city 1331 2452 Tympaki Moires over 350,000 Samaria Palaiochora other city 1231 over 150,000 Ormos Kolpos Almyrou Crete Chanion over 100, Antikythira 000 other main city 1476 MEDITERRANEAN SEA other city KRITI 0 km 50 100 150 km MEDITERRANEAN SEA 0km 28 56 84 km Kriti (Crete) * Yugoslavia is divided into 2 federated states : Montenegro and Serbia. Vojvodina and Kosovo are autonomous provinces of Serbia. Since June 1999, Kosovo is under protectorate of UNO. Gavdos Photo 1: Southeast Europe. X. XXX Photo 1: Physical map of Greece. X. XXX

SERBIA BLACK SEA Priπtina BOSNIA-H. MONTENEGRO YUGOSLAVIA Edirne YUGOSLAVIA KOSOVO s Istanbul a Ard TURKEY Orestiada E YUGOSLAVIA v european part r o s BULGARIA s Didymoteicho u Podgorica r o Lavara p os CROATIA Soufli B Plovdiv Echinos SEA OF Tykhero Kato Sapes Nevrokopi MARMARA stos Skopje Ne Komotini * Xylagani Sidirokastro F.Y.R.O.M Xanthi Irakleia Chrysoupoli Agios * Charalampos Former Yugoslav Republic ANATOLIKINea Karvali MAKEDONIA KAI THRAKIAlexandroupoli Skoutari Drama Keramoti les Rodolivos s Feres of Macedonia o Nea Peramos r Serres Kavala v Kilkis Xylopoli Mikropoli E anel LRodopoli Dojran L Kerkinitis rd o s a Aridaia Str D imo g nas a A l x Edessa i o Nigrita Thasos e ADRIATIC s Samothraki Arnissa Mavrothalassa Thasos P Platy L Volvi Polykastro KENTRIKI MAKEDONIA Florina Naousa L Koronia Asprovalta Limenaria o SEA Tirana Amyntaio Kolpos Nea Santa Lagkadas Stavros k Vergina Nea Apollonia Lake L Vegoritis Giannitsa Kalamaria Orfanou i Ptolemaïda Prespa Evosmo Gerakarou Kastoria Korinos k Maniakoi Veroia Thessaloniki(Salonica) Argos Orestiko Polygyros a Nea AGIONOROS r Galatini Roda(Mont Athos) L KastoriasVelvendos Karyes Siatista Servia Kosani Katerini Leptokarya h Sfendami A K Samothraki Platamonas gi o Akra T s o lp a uOo Akrathos n Sikeas Grevena L Aliakmonas Koufaliao r Elassona m Portoou Karras Akra ALBANIA k Ko s a K Deskati li a l Mourtzeflos s po Konitsa A s s Tyrnavos Thermaïkos an Alexandreia dr Arnaia Akra Agia as Verdikoussa Kolpos Eirinis Vasilika Galatista Kalampaka Aiginio Akra Drepano Limnos Metsovo Mesimeri Akra Plaka S Nikaia Akra t Paliouri r a Farkadhon Kassandras i DYTIKI MAKEDONIA Nea Moudania Moudros t Katsikas Neos s TURKEY Rigaio Agios Myrina Liapades o Filiates IoanninaPramanta Pyli Trikala Nea Anchialos Kassandreia e Efstratios f Sofades Farsala Volos Koufosd Skyros Akra Sigri O Paramythia Kallifoni Almyros ITALY Igoumenitsa Ao Paliouri os Neo Monastiri a ) t r Giouras Kerkyra r Domokos e Akra Perdika Thesprotiko Larisa o d Mithymna Maleas Lefkimmi a Pyrgetos p Agios Efstratios Mytilini Kanallaki Pelasgia ra n Filippiada Petas S o Piperi Parga Stylida i p Lesvos t Arta o S Sigri o Makrakomi ei n (Lesbos) Agia r er Ereikoussa Krikellos o th Akra Kartsino Eresos Karpenisi s STEREA ELLASV r Chios Polichnitos eio o Akra Preveza Pin (N Alonnisos Othonoi is Vamvakas Plomari Vonitsa Amfilochiam Mathraki a Kato TithoreaTHESSALIA Pagasitikos VOREIO AIGAIO y

h

T A Stefanoviki Lefkada Palairos r Kolpos a Davlia Skiathos c Skopelos h Levadeia (Corfu) t A Kandila h c Kyra Panagia Skyros Izmir Thermoo h Chalkida Akra SarakinikoPatitiri

s e Aliartos AkraValaxa Kymis Agrinio l Itea o Argalasti Skopelos o Peristera s L Trichonida Kerkyra IPEIROS Skantzoura (Corfu) Antipsara Akra Meston Marmaro Istiaia Akra Masticho s Volissos Ithaki o p Psara o Lamia r Chios

v Loutra I or Paxoi a Spercheio V eios Mantoudi Akra Kafireas Sami T s AidipsouE Kymi Molos vv Argostoli Antipaxoi oï Oxylithos Lixouri ko Poros Amvrakikos Kolpos s Akra Fanari IONIOI NISOI( Agios Konstantinos K Psachna Ochthonia Akra Kampanos Pyrgi Kifisos Malesinaolp Nea Artaki o os Korinthos Martino AliveriEvvoia (Euboea) I Drosia Krieza OrchomenosAkraifnio Eretria Samos IONIAN Almyropotamos Samos Katastari Amfissa Schimatari Agioi Neo Karlovasi o ApostoloiPetaliori n Zakynthos Thiva AEGEAN SEAAkra Livada Akra Papas Avlida Marathokampos Lefkada Erythres Marathonas

SEA Mataragkaös Astakos Kapareli Nea Makri

o Panaitolio Platanistos

l Nafpaktos

n (Leucade) e Kifisia Karystos Aitoliko h

c s Gyaros Agios

o Mandra Kolpos Gavrio A n Kamares i r Nea Liosia Andros Christos Kirykos DYTIKI ELLAS Mesolongi o Aigio Petalion Andros M Korin i thiakoXylokastro Megara SalaminaZografosKoropi Ikaria MakronisiAgathonisi sKo Kiato Rineia Arkoi pos Peristerio Keratea Vrachnaïkas Loutraki AgioiAigina Theodoroi o o Agios Voula a Akra Doukato Kato Achaïa olp Kalavryta Zevgolatio Tinos Patraïkos K Patra GeorgiosATHINALavrio(Athens) ErmoupoliTinos Leipsoi Agios DimitriosPalaia Kea Mykonos Ithaki Poros Fokaia Gyaros Patmos Akra Dafnoudi Varda Akra Katakolo Kleitoria SalaminaSofiko n Nemea Chiliomodi AkraVoudi i Loutra AkraLechaina Araxos Kandila (Corinth) Poseidonia D Kos Argos Peiraias Kallithea Mykonos Leros Kalymnos Kefallonia Gastouni Lygourio Kanala Vartholomio Poros Syros ( Amaliada Tropaia Levitha s LevidiTripoli Nafplio Kea Naxos Kardamaina Platanos Spetses o Kalymnos o Donoussa m I Akra Mounta Paros D i Kefalos Krestena Ermioni Serifos r Ydra Kythnos e N Astros Kranidi Naxos s Trianta s Akra Skinari Kinaros d P Zacharo Megalopoli Kamares o Gyali PELOPONNISOS Symi Kamiros Afantou Kato Agios Petros Spetses Paros l Pyrgos Figaleia Alf Akra e Kos Nisyros Archangelos eios ATTIKI Keros d i A Leonidi Ierax (Rhodes) AkraEmponas Prasonisi Rodos Zakynthos r Alimia g Serifos Ios Akra Paraspori Megisti a K Antiparos Irakleia k o Kyparissia Meligalas o Amorgos li Dokos Ydra Sikinos e Lindos s (Zante) Arfara lp k Akra Marathia Sparti o o Sifnos Tilos Vatio n Filiatra s s Milos Astypalaia Kyparissiakos Geraki Folegandros c Syrna a GargalianoiKolpos Messini Anydro d o Chora Akra Maleas Anafi KykladesOfidoussa Thira a n Kattavia Petalidi Skala Molaoi (Rhodes) s Arna Sikinos Emporeio Chalki Rodos i Pylos Longa Papadianika Kimolos Ios n i MirtoöMonemvasia Pachia Makra ) Akra Tainaro Gytheio Polyaigos Zafora s Methoni KoroniKalamata Velopoula Antimilos e Strofades Areopoli Falkonera o Evrota Karavonisia s Akra KastellouDiafani s Akra KapelloNeapoli Folegandros s Pélagos Milos Anafi Astakidae Proti ) Ananes Thira Chamili Divounia Vathia Karavas Akra Agios Akra Sideros Messiniakos Potamos Karpathos Kolpos Akra Akritas Akra Spatha Ioannis Avlemonas NOTIO AIGAIO Sapientza (Cyclades) Lakonikos Kythira Schiza Elafonisos Kolpos Avgo Irakleio Karpathos SEA OF CRETE Dragonada Nea Elasa Siteia Palaikastro Alikarnassos Limin Chersonisou Kythira Malia Neapoli Zakros Rodopos Souda Akra Lithino Kastelli Agios Nikolaos Rethymno Perama Anogeia Kastelli Arkalochori Kalo Chorio National Capital (3,093,000 In 93) Krousonas Kournas Ano ViannosKasos Ierapetra over 350,000 Elos Koufonisi Chania Asimi Sfakia Tympaki Moires over 150,000 Palaiochora Chrysi over 100,000 Ormos MEDITERRANEAN SEA Kolpos Almyrou Chanion other main city Antikythira Dia other city

Capital of administrative region KRITI MEDITERRANEAN SEA 0km 28 56 84 km Kriti (Crete) Gavdos Photo 1: Physical map of Greece. X. XXX Photo 1: Political map of Greece. X. XXX 14 Introduction Introduction 15

Photo 1: Meadow in the mountains of . P. Pafilis Photo 3: Pindos, the “mountainous spine” of Greece. A. Trichas

Photo 2: Rhodopes Mountains along the northern border with Bulgaria. A. Trichas Photo 4: Vipera berus can be found in the mountains of northern Greece. J. Van der Voort 16 Introduction Introduction 17

Photo 6: Venetiko Islet (right), Schinousa Island, Agrilou and Ophidou Islets (left and center), and Keros Island in the back. ?. ???

Photo 5: The peak of Olympus, the highest mountain in Greece. A. Trichas Photo 7: The green Megali Diavati Islet in Skyros Archipelago, hosting an astonishing population of giant Podarcis gaigeae. ?. ??? 18 Introduction Introduction 19

Photo 8: Maquis is characterstic for the Taigetos Mountain area, near . Photo 10: Phrygana on Daskalio Islet, out of Keros Island. ??? P. Pafilis F. Wüthrich

Photo 9: Degrading maquis on Keros Island. P. Pafilis Photo 11: Mountain torrents like this one on the northern Peloponnese, dry out in summer. P. Pafilis 20 Introduction Introduction 21

rain in most of the country and snow in the highland regions, comes during the win- As a result, ambient conditions are of paramount importance for the field her- ter months. While all of Greece experiences a pronounced dry season during the sum- petologist. While both regional climate and local microclimatic conditions will shape mer months, this period varies substantially in length and intensity, and can range any- the composition of the herpetofauna in an area, prevailing weather will also determine where from 1 to 7 months of the year. whether one is likely to encounter a particular species. Different taxa can also differ A surprising amount of variation in climatic conditions exists between the differ- widely in their specific thermal preferences. While a cool and partially overcast spring ent parts of the country. As a general rule, higher elevations tend to receive more pre- day in the Cyclades can be warm enough to lure Aegean wall lizards (Podarcis erhardii) cipitation. As clouds are forced up the mountain moisture condenses and comes down out of their hiding places, it will likely fail to make harduns (Laudakia stellio) appear. in the form of rain or snow on the windward side of a mountain, in contrast, the lee- Alternatively, while on a hot summer afternoon in Ikaria, it is still possible to en- ward side remains comparatively dry. Although this can be observed on a small scale counter Ophisops elegans in the open, Lacerta oertzeni will have probably retreated in- on individual mountains, it is particularly true for the Pindos mountain chain, which to their refugia. forms an effective barrier against the prevailing northwest weather fronts by running Despite its small size, the varied topography of Greece offers a diversity of year in a roughly north-south direction. As a result, receives significantly round herpetological opportunities. During the winter months, when snow and cold more precipitation than the eastern region. This is also reflected in the fact that there conditions prevent herpetological investigations in northern Greece such as the moun- is substantially more surface water, in the form of lakes and rivers, in the western rather tains of Pindos and the Peloponnese, most species of reptiles and amphibians will be than the eastern slopes of the Pindos range. As one travels eastwards and southwards, active in the lowlands of Crete or the . On the other hand, when the low- precipitation progressively decreases and reaches a minimum of less than 400 mm in lands of southern Greece are baking in the summer heat and most reptile species be- the low, “desert” islands of the southeast Cyclades. A similar pattern but on a smaller come inactive, a trip into the highlands of the Rhodopes or the Prespa Lakes can be scale can be observed on Crete, where western regions around Chania receive more particularly rewarding. Thus, the best months for reptile and amphibian observations than twice the precipitation of the far southeast corner around Ierapetra. in the Aegean and as well as the lowlands of southern Greece fall in the Temperature regimes in Greece are influenced by three key factors: latitude, alti- months of March through June, and then from September through November. In tude and distance from the sea. For example, the average January temperature in contrast, the best period to visit the mountains and the highlands of northern Greece northern Greece is as much as 8 0C warmer than comparable temperatures in Crete starts in May and lasts through September. and the southern Dodekanese. This effect is further exacerbated by the higher average In practice, there are a number of ways to make the most of a herpetological visit elevations found in the north of the country. As a result, the winter landscape of to Greece. Driving up or down a mountain range can provide access to those eleva- Florina or the Prespes Lakes region is regularly blanketed with snow and the overall tions where conditions are most favorable for reptile or amphibian activity at any giv- conditions resemble central Europe. In regions above 1,000 m snow may remain on en time of the year. On a smaller scale, the aspect of an area can also influence the ac- the ground for four months of the year and above 2,000 m this period may extend to tivity of reptiles. In our experience, not only do north-facing slopes receive less solar more than half the year. radiation but are also covered by dense vegetation and have more surface water avail- Greek summers tend to be hot across the country with average July temperatures ability than south-facing slopes. As a result of these microclimatic conditions they are in the southern lowlands ranging between 24–27 0C. Even northern inland regions best visited during the summer months. In contrast, the snow melts first on the south- around Kastoria and Serres register average July temperatures above 22 0C demon- facing slopes and reptiles may emerge from hibernacula in early spring. Lastly, if con- strating the pervasiveness of the summer heat. The only areas that may experience rela- ditions are marginal, it is worth focusing search efforts to those particular hours of the tively cooler temperatures are the where the tempering effect of the sea day that are most appropriate for reptiles. For example, when summer temperatures in in combination with the prevailing meltémi winds acts to moderate the summer heat. southern Greece become prohibitively hot, searching during early morning hours or even at night along dry stone walls and ravines can be a good way of finding snakes and lizards. Climatic conditions and finding reptiles Because of their ectotherm physiology, reptiles and amphibians are more dependent on prevailing thermal conditions than are mammals or birds. 138 Species accounts Species accounts 139

Lyciasalamandra luschani (Steindachner, 1891) tuberance on the dorsal surface of the tail base and nuptial pads on the forelegs that are well-developed during the breeding season. The protuberance is thought to function in predisposing the female’s cloaca to accept the spermatophore. GR: Nyfitsa (Νυϕιτσα) • G: Lykischer Salamander • EN: Lycian salamander Notes on biology: Lyciasalamandra helverseni is active from October or November to • F: Salamandre de Lycie March, when mating occurs. This salamander is nocturnal and seems to be more active on the surface under cool, humid and windless conditions, as well as during and after Main synonyms: Molge luschani Steindachner, 1891; Salamandra luschani rainfall. The rest of the time it hides in underground burrows, cracks and crevices, Boulenger, 1892; Mertensiella luschani Wolterstorff, 1925 rocky walls, etc., usually in groups in order to avoid dehydration. This species repro- duces independently of water. The protuberance on the tail-base is rubbed against the Range: This species is found on the female’s cloaca during the ventral amplexus, but its precise function is unknown. The southern coast of Turkey and on the female usually gives birth to 2 fully developed young after a gestation period of 1 year. Greek island of (Megis- Sexual maturity is attained at 3 years. Longevity is estimated to be 14 years. Adults feed ti). mainly on crawling invertebrates. Habitat: The Lycian salamander oc- Conservation: This species is listed in Appendix II of the Bern Convention, and in curs in various habitat types such as Annexes II and IV of the EU Natural Habitats Directive (under the name Mertensiella Mediterranean maquis, pine forest luschani, see Remarks). It is protected by national legislation and is listed in the and scrubland. Shady slopes or val- National Red Data Book as “rare”. Small island populations of L. helverseni encounter leys are preferred, particularly those the same problems as all other rare species. The population on Kasos Island is the most with a loose substrate and limestone endangered. Main threats to this population are habitat destruction due to over-grazing crevices that provide efficient hiding and collecting for the pet trade. places. The species is abundant near human settlements, inhabiting loose Remarks: Before the recent taxonomic re-evaluation, the species was known as rock walls and ruins. Mertensiella luschani helverseni. Identification: The maximum length is 140 mm for males and 150 mm for females. The tail is usually slightly shorter than the SVL. The head is flat and somewhat longer than wide. 11–13 weakly defined costal grooves are present at either side of the long, cylindrical body. There are well-defined parotid glands at the back of the head. The colouration in this species is highly variable. The dorsal surface varies from pale brown to orange-pink, brighter on the parotid glands, with brown or black spots irregularly distributed across the back. The head, tail and sides of the limbs are bright orange-pink and sparsely covered with brown spots. Unlike females, the males possess a spike- shaped protuberance on the dorsal surface of the tail base and nuptial pads on the forelegs, best developed during the breeding season. The protuberance is thought to have a function in predisposing the females’ cloaca to receive the spermatophore.

Notes on biology: Lyciasalamandra luschani is a strictly terrestrial amphibian, mostly active during the cooler winter months when it can be quite abundant. Mating also takes place during this period. Mostly nocturnal, it seems to be more active on the sur- 140 Species accounts Species accounts 141

Photo x: Lyciasalamandra luschani male. K. Sotiropoulos Photo x: Salamandra salamandra. K. Eleftherakos

Photo x: Lyciasalamandra luschani male. A. Trichas Photo x: Salamandra salamandra female. Southern Peloponnese. D. Escoriza 142 Species accounts Species accounts 143

face during and after rainfall, and when atmospheric pressure drops. During the sum- Salamandra salamandra (Linnaeus, 1758) mer it aestivates in deep underground burrows, rocky cracks or crevices. Reproduction is independent of water. The protuberance on the tail-base is rubbed against the fe- GR: Salamandra (Σαλαµανδρα), Vrohalida (Βροχαλιδα), Voidaki tou Theou (Βοιδακι male’s cloaca during the ventral amplexus, but its precise function is unknown. The του Θεου) • G: Feuersalamander • EN: Fire salamander • F: Salamandre tachetée species is viviparous and usually gives birth to 2 fully developed young after 1 year of gestation. They can measure as long as 7 cm at birth, and weigh up to 2 grams. Sexual Main synonyms: Lacerta salamandra Linnaeus, 1768; Salamandra maculata maturity is reached in 3 years, while longevity is estimated to be over 10 years, excep- Schrank, 1786; Salamandra maculosa Doderlein, 1872 tionally reaching 22 years (!). Adults feed mainly on earthworms, slugs and various in- sects. Range: A species widely distributed across central and southern Europe, Conservation: This species is listed in Appendix II of the Bern Convention, and in northwestern Africa and the Middle Annexes II and IV of the EU Natural Habitats Directive (under the name Mertensiella East. In Greece it is found through- luschani, see Remarks). It is protected by national legislation and is listed in the out the mainland between 600 and National Red Data Book as “rare”. The Lycian salamander is very abundant on 1,800 m elevation. Kastellorizo. However, general habitat alteration and loss, habitat modification from deforestation or logging related activities, urbanization, loss of genetic diversity (a Habitat: Salamandra salamandra in- small population phenomenon), are the main threats for the viability of the species. habits mainly deciduous and mixed, sometimes conifer forests, where it Remarks: Populations on Kastellorizo are assigned to the subspecies L. luschani ba- hides under logs or stones, in cracks soglui (Author, ???). or crevices, leaf litter, and other places that provide shelter and mois- ture. Populations inhabiting anthro- pogenic landscapes and deforested habitats can be considered relicts of formerly forest dwellers.

Identification: This species reaches 250 mm in body size. Females are generally larger than males and possess relatively shorter limbs and tail. The cloaca of the male is much more swollen than that of the female. The tail is cylindrical and shorter than the SVL. Parotid glands behind the eyes are prominent and always pigmented. Two rows of poi- son glands run down the back and the tail, while two more run along each flank. Dorsal and lateral surfaces are black with large yellow to orange spots and/or bands. The ventral surface is black or brownish. This colouration and pattern serve two pur- poses: cryptic, where the spots on a black background allow the animal to hide on a dappled forest floor, or aposematic, where the bright spots indicate poisonous skin se- cretions.

Notes on biology: A mostly nocturnal species, it spends the day hidden in favourable places. Females are active through the day in the breeding season. During rainy weather 144 Species accounts Species accounts 145

Photo x: Salamandra salamandra. A. Trichas Photo x: Salamandra salamandra. Taigetos, Peloponnese. B. Trapp

Photo x: Salamandra salamandra foraging for earthworms in leaf litter. A. Trichas Photo x: Salamandra salamandra. Taigetos, Peloponnese. B. Trapp Species accounts 197

AGAMIDAE

Laudakia stellio (Linnaeus, 1758)

GR: Krokodilaki (Κροκοδειλακι) • G: Hardun • EN: Rough-tailed agama • F: Stellio commun

Main synonyms: Lacerta stellio Linnaeus, 1758; Stellio vulgaris Sonnini & Latrellie, 1802; Agama stellio Boulenger, 1885; Stellio stellio Moody, 1980

Range: Greece is the western boun- dary for this species, which is distri- buted through Turkey, Syria, Leba- non, Egypt, Iraq, Saudi Arabia, Cyprus, Jordan and Israel. Within Greece it is present on the islands of the east Aegean (on the majority of islands from Lesbos to Kastellorizo), the Cyclades (Mykonos, Rhineia, , Paros, Naxos, Despotico, Antiparos), Corfu (introduced in 1915), , and some areas around Thessaloniki.

Habitat: In the eastern Aegean is- lands the species appears in a variety of habitats including Mediterranean maquis, phrygana, oak forests, groves and other cultivated areas. It prefers rocky places or stonewalls. In other areas of Greece it frequents olive groves, dry places with phryga- na, and stone walls or archaeological ruins. 198 Species accounts Species accounts 199

Photo x: Laudakia stellio daani male. Lesvos. A. Trichas Photo x: Laudakia stellio daani. Ticks are common parasites of rough-tailed agamas, as the one in the neck of this specimen from Paros. ?. ???

Photo x: Laudakia stellio daani. Rhodes. A. Trichas Photo x: Laudakia stellio daani. Samos. E. Razzetti 200 Species accounts Species accounts 201

Identification: Laudakia stellio is an unmistakable animal resembling to some extent a ANGUIDAE small crocodile. Adult total length is up to 30 cm. The tail is long, almost of the same length as the body. The body is flat and the head triangular. It is covered with small, asymmetrically distributed scales and plates. There are many spiny scales on the neck and the sides of the head. The tail is covered with spiny scales and it does not break easi- Anguis cephallonicus (Werner, 1894) ly. The upper part of the body is dark brown or gray with some yellowish spots on the back. The belly is whitish, and males have light blue spots on the head. The legs are long and the ear orifices visible. GR: Konaki tis Peloponnisou (Κονακι της Πελοποννησου) • G: Peloponnes-Schleiche • EN: Peloponnese slow worm • F: Orvet du Peloponnese Notes on biology: This diurnal lizard is active from early spring until late autumn. It climbs well and hides in crevices in tree trunks as well in rocks and between the stones Main synonyms: Anguis fragilis peloponnesiacus Stepanek, 1937 of walls. Males defend their territory by bobbing their heads up and down rhythmically. Females lay 6–14 eggs at the end of spring. The first offspring appear in mid-July. Range: Anguis cephallonicus is en- Laudakia stellio feeds mainly on arthropods and other invertebrates, but also on lizards demic to Greece and distributed and plant materials, such as fruit. throughout the Peloponnese, Ce- phalonia, Zakynthos, and Conservation: The species is listed in Annex II of the Bern Convention, and in Annex Levkada. IV of the EU Habitats Directive. It is also protected by Greek Law (Presidential Decree 67/1981). Habitat: Anguis cephallonicus has been recorded from sea level up to Remarks: Other Greek names are Kasidis, Korkofila (), Krokudialos, Krokotavlos an altitude of 1,200 m (Mount Tay- (Samos), Kroka (), Kurkudialos (Kalymnos), and Krokodilos (Mykonos). getos). It frequents humid areas in meadows, scrubland, open forest (deciduous and coniferous), hedge- rows and wooded stream banks. It also occurs in rural gardens and tra- ditionally farmed agricultural areas.

Identification: Anguis cephallonicus is a legless lizard with a body length of up to 50 cm. Morphologically it is similar to Anguis fragilis, but adults are more slender and have narrower heads. The number of scales at mid-body is greater in the former than in the latter species (34–36 vs 24–26). The back is creamy brown, the flanks and the belly are dark brown. The borders between these areas of the body are well defined, from the tip of the nose to the tip of the tail. There is usually a dark stripe, 4–6 cm in length, which begins from the middle of the neck.

Notes on biology: The available information on the biology of this species is limited. Generally speaking, some of its habits are similar to those of Anguis fragilis. 394 Species accounts Species accounts 395

Photo x: Zamenis situla. Samos. E. Razzetti Photo x: Zamenis situla. Milos. G. Kreiner

Photo x: Zamenis situla. Western Peloponnese. K. Sotiropoulos Photo x: Zamenis situla. Rhodos. B. Trapp 396 Species accounts Species accounts 397

VIPERIDAE Overall, the colour is very variable with a darker zigzag pattern on the dorsal surface of the body.

Notes on biology: is active mainly during the day and is usual- Macrovipera schweizeri (Werner, 1935) ly seen out during summer, especially on road surfaces. It is a good climber and dur- ing spring it often creeps up trees to prey on nesting and perching birds. In addition it seems to be a skilful swimmer, crossing small streams and lakes. It is mainly a diurnal GR: Levantoxendra (Λεβαντοχεντρα), Ohia tis Milou (Οχια της Μηλου), Therio snake that is also active at dusk or early in the night during the summer. It preys upon (Θεριο) • G: Milosotter • EN: Cyclades blunt-nosed viper • F: Vipère des Cyclades mammals (mostly rodents), birds, lizards (especially on Podarcis milensis) and insects (usually Coleoptera). Mating occurs in May. The female lays 7–11 eggs, the incubation Main synonyms: Vipera lebetina schweizeri Werner, 1935; Vipera lebetina siphnensis period lasts 30–45 days and the young hatch out in August. Hatchlings are 18–25 cm Wettstein, 1952; Daboia lebetina schweizeri Obst, 1983; Vipera schweizeri Nilson & long. It is the only egg-laying viper in Europe. It is not an aggressive snake and hisses Andrén, 1988 before attacking.

Range: Found only in the Milos Ar- Conservation: Endemic to the Milos archipelago. Vulnerable. Until the 1970s the chipelago (Milos, Kimolos, Polyaigos Forest Service classified Macrovipera schweizeri as vermin. Main threats are illegal cap- and Sifnos). ture by reptile collectors, habitat destruction (especially on Milos, which hosts the largest population), and accidental death on the roads. Milos has a particularly inter- Habitat: The island ecosystems esting and valuable geological history and its exceptionally rich subsoil has been ex- where this species is distributed are ploited since ancient times. Mining and the new road network are responsible for the more or less similar, phrygana and dramatic decrease in the viper population in Milos. This species is listed in Annex II of maquis being the dominant type. It the Bern Convention, and also in Annexes II and IV of the EU Habitats Directive as prefers humid areas close to water a priority species, while it is protected by Greek Law as well (Presidential Decree such as streams, small rivers and lakes, 67/1981). Listed in the IUCN Red Data book as “endangered”, whereas in the Greek and is usually found in gardens and Red Data Book of threatened Vertebrates it is referred to as a “vulnerable” taxon. cultivated land.

Identification: A large, robust viper Remarks: Highly venomous! As mentioned above, colour varies widely. The most im- (total length 1.5 m). The head is pressive specimens display a red background with a blackish rhombus shape on the large, triangular, and clearly distinct dorsal surface. from the neck. The eyes are large with a vertical pupil like in all vi- pers. The tail is relatively short. The head is covered with small, mostly keeled, scales; apicals: 2–3, nasal: 1, partially fused with the rostral, ocu- lars: 11–18, upper labials: 9–12,lower labials: 4–5, ventrals: 126–181 in 23–27 rows, anal undivided: 29–58. 398 Species accounts Species accounts 399

Photo x: Macrovipera schweizeri is endemic to the Milos Archipelago. J. Speybroeck Photo x: Macrovipera schweizeri climbs trees, particularly during spring, to feed on birds. M. Dimaki

Photo x: Macrovipera schweizeri. Milos. B. Trapp Photo x: Macrovipera schweizeri. Milos. B. Trapp 400 Species accounts Species accounts 401

Vipera ammodytes (Linnaeus, 1758) April to May. Females lay 12–20 eggs. Vipera ammodytes feeds on small mammals, birds and lizards. When disturbed it first tries to escape rapidly, hissing loudly. If its escape route is cut off, it coils up and strikes vigorously. GR: Ohia (Οχια) • G: Sandotter • EN: Sand viper • F: Vipère ammodyte Conservation: This viper is listed in Annex II of the Bern Convention, and also in Main synonyms: Coluber ammodytes Linnaeus, 1758 Annex II the EU Habitats Directive. This species is present in many protected areas.

Range: North Italy, south Austria, Remarks: Highly venomous! Vipera ammodytes is the most widespread viper in the and southwest Asia. In country and its venom can prove fatal. For this reason its name has become a synonym Greece it is present on the mainland for treason and death in country lore. and many islands of the Aegean and Ionian Seas.

Habitat: Found in a vast variety of biotopes, from dune ecosystems at sea level to mountains of 2,500 m height. It is found on rocky slopes with a sparse vegetation, phryganic biotopes and light Mediterranean forests while also inhabiting man- made habitats such as gardens, dry stonewalls, cultivated fields and even areas close to human settlement.

Identification: Vipera ammodytes is a small-sized, stout-bodied viper of a total length up to 90 cm, though usually under 65 cm. The head is large and triangular, covered in small scales. Over each eye is a single, flat scale. The eyes have vertical pupils and are separated from the upper labials by 2 rows of small scales. At the end of the head is a distinct nose-horn. The tail is shortened and ends in a spine. Head scalation: 10–13 scales around the eye, 9–11 upper labials, 12–14 lower labials. Body scalation: 21–23 rows of dorsal scales at mid-body, 160–165 ventrals, with the anal shield undivided. Colouration shows an astonishing variety. A characteristic vertebral band, consisting of rhomboid blotches fused into a zigzag, runs from the neck to the tail. The band is usually darker than the background, which varies from light gray or yellowish brown to dark red, and even melanistic specimens have been reported in rare cases.

Notes on biology: Activity pattern alters according to the season. Though usually ac- tive by day, may be seen after dusk and even at night during warmer periods. It climbs bushes and trees with ease, especially during spring. The breeding season lasts from 402 Species accounts Species accounts 403

Photo x: Vipera ammodytes meridionalis ready to attack. Western Peloponnese. Photo x: Vipera ammodytes meridionalis. Mani Peninsula, Peloponnese. D. Escoriza K. Eleftherakos

Photo x: Vipera ammodytes meridionalis male. Florina. G. Kreiner Photo x: Vipera ammodytes meridionalis pair. G. Kreiner 424 Bibliography Bibliography 425

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