17th European Congress of Herpetology Veszprém, Hungary PROGRAMME & ABSTRACTSPROGRAMME 17 th European Congress ofHerpetology Veszprém, Hungary Univ 22–27 August 2013 ersity ofP annonia 17th European Congress of Herpetology Veszprém, Hungary

PROGRAMME & ABSTRACTS

University of Pannonia 22–27 August 2013 Chief Patron: Welcome Tibor Navracsics, Deputy Prime Minister, Minister of Public Administration and Justice

Organising Institutions: Societas Europaea Herpetologica (www.seh-herpetology.org) Hungarian Ornithological and Nature Conservation Society, MME/Birdlife Hungary (www.mme.hu) University of Pannonia, Department of Limnology, Veszprém (ornithology.limnologia.hu) Hungarian Natural History Museum, Budapest (www.nhmus.hu/en) Dear Colleagues, Local Organising Committee: Judit Vörös (Hungary) Welcome to Veszprém, Hungary! Bálint Halpern (Hungary) Gábor Seress (Hungary) Júlia Tünde Gál (Hungary) The 17th SEH European Congress of Herpetology is hosted by the University of Pannonia, and co-organised by the Hungarian Ornithological and Nature Conservation Society (MME) Scientific Board Members: and the Hungarian Natural History Museum. Gergely Babocsay (Hungary) Participants from six continents (39 countries) registered for the event, giving the meeting José Brito (Portugal) Salvador Carranza (Spain) a global perspective on the science of herpetology. Miguel Angel Carretero (Portugal) The scientific programme includes four invited talks on topics of broad interest (evolution Dan Coga˘lniceanu (Romania) of reptile venom, invasive species, biogeography and systematics) and 113 talks in 11 different Claudia Corti (Italy) parallel sessions. On the symposia day, four symposia will be presented on relevant topics Mathieu Denoël (Belgium) such as invasive alien species, chytridiomycosis in Europe, monitoring of Natura 2000 species Trent Garner (United Kingdom) Bálint Halpern (Hungary) and herpetofauna and transport systems. There will also be 120 poster presentations divided István Kiss (Hungary) between two poster sessions. Luca Luiselli (Italy) Wine will be served during the poster sessions and a morning raffle will be held with Petros Lymberakis (Greece) valuable prizes to be won. There will be a contest for the best student talk and poster, and for Sebastian Steinfartz (Germany) Zoltán Tamás Nagy (Belgium) the first time in the history of SEH European Congress of Herpetology, a silent auction will Ljiljana Tomovic´ (Serbia) take place. Sylvain Ursenbacher (Switzerland) The welcome party, the wine tasting in Balatonfüred-Csopak Wine Region and the gala Miguel Vences (Germany) dinner all provide further opportunities to meet the other delegates. Judit Vörös (Hungary) Although August is not the best time for field herping, locations have been chosen for con- Ben Wielstra (The Netherlands) gress excursions where sightings of amphibians and reptiles are guaranteed. These locations Secretariat: are the Kis-Balaton and the Hungarian Meadow Viper Conservation Center in Kiskunság. Klára Biszkup–Nánási (Hungary) On behalf of the Local Organising Committee I wish you an exciting and interesting meeting.

17th European Congress of Herpetology, Veszprém, Hungary, 22–27 August 2013. Programme & Abstracts.

Congress logo: Balázs Farkas Herp photos: Bálint Halpern Kind regards, Preprint: Artibeus, Gyúró (Hungary) Judit Vörös Our Sponsors General Information

Venue University of Pannonia Egyetem út 10, 8200 Veszprém

The plenary sessions (conference room) will take place in the Aula (building B), this is the place where the posters will be on display. The sessions will take place in building A, in rooms 1 and 2.

Congress bureau before and after the congress Altagra Business Services and Travel Agency Ltd. Örösi Pál Zoltán sétány 0172/19. hrsz. 2100 Gödöllo˝ Phone: + 36 28 432985 Fax: + 36 28 419647 E-mail: [email protected] Homepage: www.altagra.hu

Congress bureau during the congress University of Pannonia Egyetem út 10, 8200 Veszprém Phone: + 36 30 3400013

Opening hours 22 August, Thursday 14:00–20:00 23 August, Friday 08:00–18:30 24 August, Saturday 08:00–17:00 25 August, Sunday 08:00–18:30 26 August, Monday 08:00–16:00

Internet access Free Wi-Fi is available in the University halls.

Message board You will find a message board in the Aula of the University. Coffee Congress photo Coffee, tea, soft drinks, mineral water and small cakes will be served during the coffee breaks A picture of participants will be taken and later uploaded to the congress website. in the dining room adjacent to the Aula. Place: stairs of the Aula building Please note that it is prohibited to consume meals or drinks in the Aula. Time: 24 August, 12:30

Lunches Gala dinner Please note that lunches are not included in the registration fee. However, we offer you lunches Date: 25 August (Sunday), between 19:00 and 23:00, in Betekints Étterem (Restaurant) between 23 and 26 August at the venue of the Congress (price per lunch 12 EUR). Limited (Veszprémvölgyi u. 4, 8200 Veszprém). Transfer will be provided. number of lunch tickets will be available to purchase on site. Lunch will be served at the Meeting point: Entrance of the Aula Menza of the University. The buffet lunch includes: warm dishes (soups and main courses), Departure time: 18:30 salads, desserts, mineral water, soft drinks and coffee. Transfer back: Between 21:30 and 23:00 continuously to all official congress hotels Admission: Only with ticket, tickets are available in limited number at the registration desk Price: 20 EUR Social Events

Opening Ceremony and Welcome Party Field Excursions Date: 22 August (Thursday) Time: 19:00 hrs 1. Visiting the Hungarian Meadow Viper (Vipera ursinii rakosiensis) Conservation Centre Place: in Kiskunság Opening Ceremony: Aula of the University of Pannonia Date: 27 August (Tuesday) Welcome Party: Dining room of the University of Pannonia, adjacent to the Aula Services provided: Transportation by bus, English speaking guide, lunch Please note, that it is prohibited to consume meals or drinks in the Aula. Meeting point: Aula of the University Admission is free for congress participants and for their registered accompanying persons. Departure time: 08:00 Transfer route after the tour: Wine tasting with dinner in Balatonfüred-Csopak Wine Region Budapest Airport, arrival not later than 18:00 (23 August [Friday] and 24 August [Saturday]) Budapest City Centre, Erzsébet tér, approx. at 18:30 Date: 23 August (Friday) and 24 August (Saturday) Admission: Ticket, price 25 EUR To be visited: Szent Donát Winery, Csopak Transfer: Will be provided 2. Kis-Balaton (Balaton Uplands National Park) Meeting point: Entrance of the Aula Date: 27 August (Tuesday) Departure time: 18:30 To be visited: Kányavár Island and other sections of the protected area and the Buffalo Reserve Transfer back: Approx. at 21:30 Services provided: Transportation by bus, English speaking guide, lunch Admission: Only with ticket, tickets are available in limited number at the registration desk Meeting point: Aula of the University Price: 30 EUR Departure time: 08:00 Transfer route after the tour: Morning raffle Veszprém University, approx. at 16:30 Every morning, five minutes before the plenary talk we will hold a morning raffle with Budapest Airport, approx. at 19:00 valuable prizes. Registration numbers on the congress badge will serve as raffle ticket. In order Keleti Railway Station, approx. at 19:30 to win, one has to be present. If owner of number is absent, another ticket will be selected. Budapest City Centre, Erzsébet tér, approx. at 19:45 Admission: Ticket, price 25 EUR Special Invitations Presentation Guidelines

Free entrance to the Budapest Zoo and Botanical Garden Poster display Budapest Zoo & Botanical Garden is the oldest zoo park in Hungary and one of the oldest Posters will be on display during the whole Congress. Authors are responsible for setting up in the world. It has 733 animal species and is located within Városliget Park. Thanks to the their poster at the beginning of the congress and removing it after the end of the congress. management of the Zoo, the participants of the congress can visit this interesting place for free between 21th and 30th August. Free check-in available between 9:00 and 18:30 hours at Venue: Aula of the University of Pannonia, ground floor and gallery the cashier where your name has already been registered. Address: Állatkerti körút 6–12, 1146 Budapest Poster boards: White wall elements in aluminium frame construction. Each poster will be The service is available for those participants and accompanying persons who have registered attached to the boards with Blu-Tack® which will be provided free of charge. Push pins are for the Congress not later than 15 August. not allowed.

Free entrance to the Tropicarium Poster size: The maximum size allowed for each poster is 80 cm wide and 120 cm high. Tropicarium Budapest is a particular indoor zoo, where you can find Central Europe’s biggest shark aquarium. Thanks to the management of Tropicarium, the participants of the con- Poster position: Vertical. Posters that do not adhere to the dimensional limits cannot be dis- ference can visit this interesting zoo for free between 21 and 28 August. Free check-in available played. between 10:00 and 19:00 hours at the cashier where your name has already been registered. Address: Nagytétényi út 37–43, 1222 Budapest Set-up: 22 August (Thursday) 14:00–18:30 The service is available for those participants and accompanying persons who have registered No poster should be set up during the Opening Ceremony! for the Congress not later than 15 August. Poster removal: 26 August (Monday) 14:00–16:00 Free entrance to the Hungarian Natural History Museum’s exhibitions No poster should be removed during the Closing Ceremony! Thanks to the management of the Museum, participants of the conference can visit the Museum for free between 20th August and 1st September. Free check-in available between Poster session 1. 23 August (Friday) 17:00–18:30 10:00 and 17:00 hours. Topics: Climate change, Ecology, Monitoring Please contact the information desk to pick up your visitor’s card. You only have to tell that Wine and snacks will be served in the dining room adjacent to the Aula. you are a SEH 2013 participant, the clerk at the desk will have the list of registered par- Please note that it is prohibited to consume meals or drinks in the Aula. ticipants. Address: Ludovika tér 2–6, 1083 Budapest Poster session 2. 24 August (Saturday) 17:00–18:30 The service is available for those participants and accompanying persons who have registered Topics: Behaviour, Morphology, anatomy and development, Distribution, Diseases and for the Congress not later than 15 August. parasites, Phylogeography, Phylogeny, Population genetics, Systematics, Physiology, Tropical herp biodiversity, ENPARTS Symposium Wine and snacks will be served in the dining room adjacent to the Aula. Programme for Accompanying Persons Please note that it is prohibited to consume meals or drinks in the Aula.

Accompanying persons are kindly invited to join the delegates at the Opening Ceremony and Oral presentations the Welcome Party. Accompanying persons are also welcome to the Budapest Zoo and Botanical The length of the oral presentations will be 12 minutes with 3 minutes discussion, 15 minutes Garden, the Tropicarium and the Hungarian Natural History Museum. altogether. Please contact the technical staff if you have additional requests. In Memoriam Olivér György Dely

The SEH 17th European Congress of Herpetology is dedicated to the memory of Hungarian herpetologist Olivér György Dely, curator of the Herpetology Collection of the Hungarian Natural History Museum for over 50 years. Olivér György Dely was born in Nagyszalonta (now Salonta, Romania) on April 27th, 1927. He studied in Nagyvárad, Nagyszalonta and Budapest. After graduating as a museologist from Péter Pázmány University in Budapest, he started to work in the Herpetology Collection of the National Natural History Museum in 1950. He published 83 scientific papers, mainly on the members of the Hungarian her- petofauna, discussing them from taxonomic and systematic points of view. During the 1956 political revolution the museum building was shelled, and nearly 40,000 herpetological specimens—including historic type material—burned to ash. In the next decades, through exchanges, donations, and collecting trips, O. Gy. Dely rebuilt the collection. Thanks to his devotion the Herpetology Collection now owns nearly 20,000 specimens. Besides replacing the collection with material representing the herpetofauna of the Carpathian Basin, he conducted several expeditions to Egypt, North Korea, and Algeria, where he gathered hundreds of specimens to enrich the Herpetology Collection of the Hungarian Natural History Museum. In 1981 he organized the First Herpetological Conference of the Socialist Countries, and in 1991, as honorary president, he hosted the Sixth Ordinary General Meeting of SEH in Budapest. Olivér György Dely passed away on November 19th, 2003. In 2013, on the 10th anniversary of his death, the 17th SEH European Congress of Herpetology is held in Veszprém in commemoration of his lifetime achievements in herpetology.

Judit Vörös One of the most exciting book projects of the last few years was the publication of the first Chimaira: a company portrait edition of Bourret’s “Les Lézards de l’Indochine” that came into being through a cooperative effort with the Natural History Museum of Paris. After the manuscript had been slumbering in an archive for some fifty years following the author’s death in 1957, an admirable initiative by Roger Bour eventually made it possible to publish this milestone treatise in 2009. The range of subject-specific literature was broadened over the past years by adding the fields “Invertebrates”, “Mammals”, “Botany” and “Ornithology”. As early as in the mid-1990s, Andreas S. Brahm realised that keeping close contact with The history of Chimaira Buchhandels GmbH began with the opening of an individualistic clients was a crucial factor for a specialised bookshop such as Chimaira. It is for this reason bookshop in downtown Frankfurt, located in close proximity to the Zoological Gardens. Right that large parts of our book range can be viewed at reptile fairs (such as the Terraristika in from the start, maintaining a combination of new publications and antiquarian stock was a Hamm), meetings of the DGHT, as well as scientific symposia of the SEH and other organi- main goal. sations every year. The countless discussions with bibliophilic terrarium keepers, herpetologists Next to entertainment books for both adults and children, Andreas S. Brahm had a par- and veterinary medicine practitioners held on these occasions have not rarely culminated in ticular interest in subject-specific literature of certain fields. As an experiment, he had a small new book projects. sales list of herpetological titles typed on a typewriter, copied and distributed (ca. 1992/1993). Major aspects, like species conservation or the systematic advancement of our knowledge This first catalogue of natural history titles was met with an unexpectedly strong response. of biological processes, are also part and parcel of a more recent project: the series entitled The following years then saw a systematic expansion of the range of both new and anti- A Life for Reptiles and Amphibians that was initiated by Fabrizio Li Vigni, the first volume quarian titles in the fields “Herpetology” and “Aquarium Keeping and Ichthyology”. Starting in of which is to appear in autumn of 2013. 1994, extensive catalogues were published regularly in August that are by now mailed to some Li Vigni’s collection of interviews portray a colourful and multifaceted mosaic of global 10,000 customers—this year (2013) thus sees the publication of the twentieth catalogue. herpetological work in that they provide intimate insights into the professional careers of New customers are most welcome to request a copy of the latest issue. several dozens of contemporary herpetologists of all ages and origins, with all five continents A first attempt at publishing was undertaken in 1995 with a treatise on African ethnoher- being represented. This idea cannot be over-appreciated, because it emphasises two points in petology (Segwagwanyana) by Ronald Auerbach. Originally chosen as a name for the pub- particular: lishing sector, “Chimaira” also became the logo of the bookshop in 1998. During the period (1) Every scientific discipline has to address its roots and history, if it is to justify its doings leading up to the turn of the millennium, further titles followed at a rate of one to two per and goals. This is achieved here for the period from about 1950 to the present day in a con- annum. With Dr. Gerold Schipper joining the team in 2000, the publishing sector was con- vincing manner in that it is an extraordinary compilation of interviews with prominent tinually expanded. The number of new publications or new editions has since increased herpetologists from the world over. The portrayal of the past history of herpetology we owe to between five and fifteen per year. to the three volumes of Contributions to Herpetology edited by Kraig Adler, while A Life for The range of publications extends from guides to terrarium keeping, via monographs Reptiles and Amphibians creates a link between Adler’s works and the present day. on reptiles and amphibians, to dissertations and field guides. The programme of Edition (2) Over and above this, the interviews are also meant as a stimulus for all those who are Chimaira here feels obliged to continuing the specifically German tradition that started in young, enthusiastic and interested in modern-day aspects of biology and who harbour the wish the early 20th century and has always been trying to create possibilities for a fruitful co- to enter a career in science or take up an occupation in nature conservation or an affiliated operation between amateurs and scientists as to broaden horizons to the benefit of both. professional field. Providing detailed guides to the captive care of reptiles and amphibians is what we regard as our contribution to a responsible interaction with animals in human care. That in doing so the protection of endangered species is to be respected and improved and the acquisition Visit our shop at www.chimaira.de of captive-bred animals is to be given preference, is a self-imposed directive we always strive to adhere to. Dr. Gerold Schipper A more recent branch of Chimaira’s publishing sector is dedicated to the field of veteri- (Publishing Manager) nary medicine. Here, the focus is once more on providing publications that can help optimise Phone: ++ 49 69 96 86 48 91 the husbandry conditions of reptiles and amphibians in captivity on the one, and convey the Fax: ++ 49 69 96 86 48 92 advances in a field that has seen rapid progress in the past decades on the other hand. www.chimaira.de Table of Contents

Programme ...... 17 Plenary Lectures ...... 33 Symposium on Alien Invasive Species and their impact on Natura 2000 amphibian and reptile species in Europe, with special emphasis on the V4 countries ...... 39 Amphibian chytridiomycosis Symposium ...... 55 Herpetofauna and transport systems—towards a better understanding and impact reduction ...... 65 Monitoring of amphibians and reptiles in the frame of the Habitats Directive . . . . 73 Oral Presentations ...... 83 Poster Presentations ...... 199 List of Participants ...... 315 Author Index ...... 329 Programme PROGRAMME OVERVIEW ORAL PRESENTATIONS: 23 August (morning) t s e s o : t f s r t f s e n y o o u e l u h o o u l a t s i g c p n u a n d r n s r , n n s u n a g o r e a o v a o i e o f o i e v i u d n t u i A e i p r u t a g c t t p a l u e L A r i a x g T a u 7 o f d s d y l i l B . t e r A D e d 2 a o e r u d o r n A u n o n k v s p e n G p t r o , a E e c a i o . i a u t t m f d a p e s i p a o T i o m r n n h M d r n n s p o i n e a t a s a f n t o r o , l l n s o o l e r p i e i i n e i a e e e g a a a t h t e u x y t y t n d c h c b m t s n i a a s p e i m i g i a n a d c i o s s l r t e t s v v n h é z h s t e n s k s p S i a a c c r r o o i - s r k n n ö a p t e n e y a a m d w t r d a h i o s s e a e r r r a c o m e l n e B h s n r i o n a p e a p n n e i t w P o P o t : k k k e t d t o a t m y d t o o n f g a - e u l d e a s a a c a c n f l a f D r m y M e e g z g e m e e i o n i n u o P e u l r u a r r a i g R S a l n S d f r c g . p m i o b b b p d n g s e o . o i c n w s u f o o r n c a E n n f y e e l v L m i e e e h s r c i e n t p o , e e A M o g a t s d t c e g o o : n f f ) s a s u ) l n o e c r e , f h f n n s n o n a t , l , f w e e a a y 6 a o M i e i h a g o o y e s s i i t W o u r e u n c n i s m t t 2 p ) a i r c r c i b g i e m C o l : u s a o C C o ? i i a b L m r e t a a f l o l r m l t l b i i r r t t y p y t i r m l a a 2 l o n o a a e g a i o r s o a a p h h a t i o o C e H d r r l o s n p t a e h i m p i e p r c e e h e p C K l m r m a E t s g m , n n o v m n p h d a e e h . n a y . d P m e S e u w n c m o e t t r i e e t r l d n s a s s e T t V a r Á n e . o t d l o s m g s s y y t r w P u e a l : i , a a , a e s V o f N y i i S S o t a t R t y d e . M c a r n s g i r t m s : m g i d e s a b c n e a n s n o g r r b a n e e y t l e c n o l n i e M n t r a a c a a z l t l R r e a n a u v a a e t s e a a ( , i n g i m i R j s t i o e S H l p i F ( o : : f o m f n o b g l e a b : e e : U l t : i o s H i r s o o n B s v o t a m v m r e s i . i r t g r a i t r h . e o h . W e i s t J : u f u t n i a a d o p t c s i i p w c v d S . t A p c g , i r A a s s o s d e e — n c m d e d f l f a u n t n a l , u n o o o i , l T s u a n i t e a l m K n u i o a m i f M B d B n i E d r p t p h a i u á i n g R a P . a h s n e , s o g s o t c e e o O n : a s e i p o r t z t . t m C r m o T A p f u G i g a n f n o t . t o ( n a l l y a y p e n i n p , t p a e P B b r o i . m a o A t i e p s s i a b o S S e t h a e p D K i o , d t m i c r : Z t f m u N i r n u a h r y m E l R : : s y r , . e k h ¸ k k i s i l n y b g g t , n k f a s p o i e t S P g E c : c o i a i g e a a a o s ü t r S n i n n m m z W c o s a t p e y m e n , e e i g o y c a h i i o h a u F k e t r r R s l r u u r e n n t l e s m r r e e f a n e ) i i i i l l n l e c c g S f y b . z n i . b b e o o o s s s i o ü i f r t M d n f a e e t l n s á S F a t t . u t i c d o o j i e c t t t u o a e e i i v . r C s v n o . h O s . R G S i e r r p p p c c r e e A E c o E n n d l , u a p U s . i f a f D s s : . a a , n a G l U i f f M a a o o n F l : t n S H d o m m e y : . 5 K d u : l r o s : : a o o , t Ç , , a f J r y y i a u r , H H d a b l . h e r r a 2 o . S n t t g r M M e , n p a r S S t ë , i e e o C C G s L m m . 2 2 . i p a A T m a i a a e T n o r e m o ó n d d : : n u u a T T m , u , t r h w c d n e n l i i o p l k i R n n n i i i n n o f m l S e a s s ü i d : : o n a m m a n e g a a v o F i i o o z o o o s A o r r t b P l m m A v u a á i i t u u i i e e a W . a p p a a M I e P o i i D e s s m m l c i H t Ü L a a S S l t s s t R F e V . R S s s a a e e . c . r e . e . n . . m o m o l l l h h N . . s n . A A h e e . y . p y y a a e P h p p I I o E L N s ( s R m A s S S C G M S T P S C R G B Z H t S S r f m m y y e ) : r l s r S S r s s a i o o d e u t d f t l d r o n p n i a d i a a e g r e o e , d v r i n z u s ´ c p n n c i h e a e v a x f f i l i m e s n r g t o s l h t e t d e o o v i s s d n a a p e e n a s t e t e c t d i i o t e n i s s r i o g n y y j e s l e a t b l i a r f u n l e b r u f i l u a g g n e i u i n n o y o f b e t g i e s g i i m o o o t c M i a v a a h l a d V l p l i i ) z b d r r a l n p l t n r h s n e l i p o o v s i t . m t o e i o s a l c 2 n r L e s o c c o a a h t l r u C i o i ( d n A n k m c o k k s r c r m t a t h i r p E E g i i r o n r , a a e e r s a a e t y w D d t o n u n i t e ´ n e e e e c b l f m a r i n e b u i a g o A i r v v r r r r v t i a o a e g h n v d e f s a s b b b e z u v a e s H r e i c c o e a u o i t G r e d m l o e l e e t g . h u r i k n l s i n u o e e A e t e c h I e o i y e e t s f f o n e a P t y s y e t a c f f c r s y y n . h c d e r r m p 4 p n r h : h i t a e n o o S a a e g n n i u n g V e u a r y S 2 n v p p r i o c e e L n e r V n c n C C r , L i n s e d p o ( a a f g g i i ´ o t W i d c c G a r : r n o n o l i r a o o o U i t i d s n n g g l l d s a : n a o t o e P v s r a r r r i t e y y t o o t e o e d g l n p r e t h r i l f a i e a e e u j s h h e n l e t t n n O a u o e z i g g o v o z n P i e e o s t i p f p d e a a h p r i l o r o h f a c s r g o c g s d e l l d i o r a s v e h m g a d H r d o n s y y e o n i t e n d j a v f e i i P n M r n p r n n h t f h E e o c s i h t e i i e m c c P a a H a e D r s l S e l i P P t i a 1 u n r s p a g : t . a . a b h d i o o e g t x l a y o d i s t l S s p d u t G r t s u y m M E o u y i b r , n a r e c e m r e l , t i p i i r g P : : o i a t a e a v a l l a o k k z a e o g e g g p c , o t i c c s u r n s s s d ö e e a l o i h p p a d r I l v r f f f f q z z f r R a e n n n r — p o a d l s o g : o o o c c o ö o n e o l l s s a u r a o a a i i e o s r r l e G e i i e e z s C a n s c c r t r d u T t y y y y s w e e e l l i i i e g m : y b b t l r t . u i i H s c p l n g g g g . s t r l E s A h i i i t t e c n a J a H H s e l d a e r o o o o s a e g m A t i E u g . h h l e i l l l p p l l e l u h e e r s u , u . . . f k i n o s a n e e i S v p p t o o o o n r ’ e r f h n t i u d i a r r D t a c c 1 c c e T t e D á t i G G A a w C r M u E M k d s m m k k p , l p P T t c v : E E E E y n . e f a i , , : ´ a n i i : , l e a a c a n a n s s e o n . o d p o i i t p i k k e z n r R e e o n r o d s n c a t y u í b r a i v r r o u í ö ö R i o i e a e , r t á s n f t r t m a n d g r r f r b o t b b s i r c a , ñ M s h o r m l k e o o o u d n ö ö e k o u m a g a i n ñ u e i . e e s p l i T a c h a o b d e t E e r a e e T T r n e A c r g I n : v a i f f j T r M n r i y O Z g m M M 1 i u . . ´ e F f f t u c r n e u n a l r 3 t e - f f f f d i J J r . . i u k b i , s s t . : o o f s A u o u g a a J o o o o o s 2 v h J n h o , , y A e i í e h n n s e i r t n Z , o t i r r d C C o l t L n , r o t c , l I r r r r v s c . s a A a a v i d e á v n z c a n : r á f n P a a i i i e j e W t o a : u t u u u p n e a a . e E a l l m a n t a s o e v o a g c s b b n . t l i a i o o o o r n h r h r l l p , o h g i i a e l s e t i t i i i e m m s B G o e e b e t e M l t e o a g ó e o l a d d u t h h T v v v v p p b r M o n c v r r . . r v b s t a c , i P n e e ö ö e L p p . a B a a a i c o a B i M u o , e H i ´ e d r r c c l : E t n K t h k O t t . n h h h h k L i g : v o . s s h . : a e c s m m H H i g , e P e c a e e e y S , a r v . i e o , i n k n u a b r r a a a : u e l e i J L e i . . e t , s i k O G o . B B B B d r c f 1 1 r t á r o r W d d e z o c u f i , i l a l e b , , y u t l a n b y R e b a W W r r r z o a h n a a q t l o t u n n a J r r o C n e e i a e r ñ l : c r : : r p n i e n s o j j i o o e a c h n r v a p á r r r . p r o r e o e v a a i o i i i r B i i n c e t o f e t b n o ó B a o s s h e a M A m r a a s s y h f . B B t o B s y I G C e t n f t g e s s t e . . r h i L h . a f a a l o h h h c m u . . i . . . n . t u e e n i e l e f n h y . m a d i o g a P C t b o d a o c p m f p w X S C A A C M M C S C O K A L P K s o r r n u r c t m e l a s u e A e p i p r 0 0 h 5 5 0 g 5 0 0 e 0 0 3 1 2 5 0 0 0 W 4 O 1 e 3 c : : T 3 0 0 5 : : : 5 0 3 : 4 2 0 : : : : : 1 : 2 0 R : 2 1 4 4 0 1 : 1 1 0 1 : 2 1 2 1 1 1 1 1 0 1 1 1 1 1 9 9 – 1 – – – – – – 0 0 0 0 0 0 – – – 0 0 0 0 0 – – – – 0 – 5 0 0 0 0 0 5 5 0 0 0 0 0 0 3 3 3 3 3 5 0 0 5 0 0 : : : : : : : : : : : 0 1 3 4 3 3 0 1 0 3 4 : : : : : 3 : : : 1 1 3 3 4 5 : : : 8 2 4 7 9 9 6 0 : : : : 1 1 1 1 1 2 0 0 0 0 1 1 2 2 1 1 1 1 1 2 8 9 9 9 1 1 1 1 1 1 1 1 1 1 1

18 19 ORAL PRESENTATIONS: 23 August (afternoon) ORAL PRESENTATIONS: 24 August (morning) l r l , : c s : i s i a e a s s , i d t e s r t k m n e s s d ´ r e r t s c e a c s h a e w t e r i e n b a n i l c n n a s l s t g g t o w f v z l F o u o e o e a n i o a a n i u o o t o d l b r . n c e s t g p v , n C o r o s a i p f i k : i n l a e r s e r e i - e e r i C l e o e r e g l p e o d t e r n : e f e t d e t i h a r u s t l d r r , s f r r D i s o d e a i r l t t n l m e a f l a g e e c o h w u a r n k g l t r p o B c i i ? t o o n u i n T n d e i n v c a t g e o c z e Z c i . i a r c W t n n s i l r a n b c s f d c E i i n e l e i . a a o m e o V a a f c i s r u o ö T i B n . a b t i v g e t t c e r a p o , r h i r m i a i i o : d a m m R M ´ n h t g e n n c i i r t o m O f s e o m s i d d a ’ i o c n u i t t f e . r r , p b o . n r a l s i o e e h o s t i r a n s a i . k I c e f a J e r r A i u u c n t i ? n e m o y v D i n l n e L t h d a , f s f d d r p y r c u n c a t a a c f , c i z a h D H g e l , e n i m o e t i o m a n r d e t t i s ˘ d a i n o i m f u s g o o . n a s i r e l h s m d e u p r E n i r e g y e a n d g l c y r v t n t n y e s l t g d o t e i h a h t i s k r D u a a t e ? o d n n i n I n a s r t r o i t o c p l i i e c e t l s e l a a o T , a e e f p c o w N v s . n y c e c S r a e ) a n i i r n ´ i d s z g n o p i c t : k l i i n t B e t i r i r y f a i y h n e t a . r e a Ç a t n i n c I n M l y n o y i i c v s a r e c a v o g o V h p l n e i e r , e o h i n d u u r : o . a f r n a G b e w á t c i r l a s l g e r y m o c i C e j o d f i e n g n t i T P l F n r i i , o l s t b a r t f e n f p o h l . a r o t c t t M t a h t h y o e a n a s : i e m l o r e g e o n s s n i i e t a a t c e f p s f f c n f u t p , e n p y d s r A l h i d o r a h t r a f h i a i h i e u r r s i h h t e o o l f e . e i r n p k o l t t e ) r a e d . y ö t a l H m c c h p o o s t A w l a a a p b i ) e d u i a s t e b u n S h e s a i e d c u . D M B i s f c l t t n r s t M N n p f n s o n p n P i c i . i o m H y s o s A o i f n i r m f n e : . . m o d l A 2 s e p , 2 r ; ( d t l a n s l r y l p . d r o b A d : o c i o a d e o m m e l E G e k , e a s o o Z o r e p S n a o t x c e v c i W m e : n s u r n n t . i o c o n a e t a : e i s d b h o a e z , n y t s . M m s p s t m i s o i a m o , l l s t y a l l z e r r i c i d p h l r e g c e o f i a a s m , d u k t a a t o e d e o R t o M r e u d p l t h e o i n i m : m r l f e r m a n H n r a a r s H v T o s t t x i t r n e i s o l , o e , o n e a p t n r a V o a C s r o t e e l v i o w i . f e s y : d m n r a g ´ n s u r l e p a t o r c l y t p i o r ) a r g . r s t s p r R ö i i R f h o i m o o i i a o e e i e : n e e n R a o o e n p s i p H s g n f t n c u G u d a s c v r s c L A p H d C s f s g s n s a t a i c d i a v e a o r V P , a e w r c c r , i e u t n n o i . . s i e i e n . , a n c . . ( n m r d p n N e A a g v o c l i c s g h . a l s t r z o d R a b i m d a d k i m a S o o t o o r t a R r y e h l n B i r A e i a G o o o . r : L o u i r i u r l n a : A C t l c e r o c n . s d O , r e g a n . e y t m l e t a o o s b , r k S s p , e b a . . p s p f s s s b v t e i o s i a . f o u w i s e l t ´ : y i S p n e R A i c u h a i e i e e ó t r d f d , a o c e o l T i a i g d e h n n M A u l i i b y a e i A N s , l d h . , ) a y w h a i g y d r ˇ h l u i r l v i n P t o e K d , c e m c a i i D o n t n p . t e k . : a i l - B , l r e d r t o ´ a g e p i d i o c t A e h m p p : u r . n i h a l n e : d m g y o u l B a u G h a d i f ) i p l ö o t n c K i p t o p f e h m a p i , z t p p e a n p M s r v k B u s o l P a i a d h v o . P . e e o t r m r a s g s i a e m , e l n C l c R r t M c e a i ó e a f n e l i m , u - f i e , o . p h m o n r a . p g i s R y i l V T y A r a r i o p N s . s r l . . n f l t ´ n I u 1 W s T h y c a o o P a n w T o e k o t a e s i . i n . : e . e h i l g o f ó e f s ( i k l l . c r f f 7 s n b , i A a E , i s a c u n . M a J . r s T s c l D d a h d c l g s o h s e h P u o o e L o e p u . e n c s B C i é 6 s o o k o . l , a u o l f s t r g , l e s n e u n r c , s Z a A n c t o 1 e M c r e I r ´ : l P . o C l n e s n k c . , c i c t n r e a i e f g o i e e r s a i o - y y i i , P f r , e a M g – e y y s e M a V y u C , a r f f i a n , F v , e N e t n l m d e B r b u k i t i s a t s e i r f s i x y c g e g a o r 9 , e : o . l g g , c i g M a j o s c o r i e m o . i l y . t u ü r o f y . e a o s l c i B g e l d t r m d a a c 6 o h c o t t d r a n t a t a z r o , o c n e r u n B d F t j n k ( e o H L r r i V i t l i l k l e y s t e u i o a l l n n z t i c l 6 r f n e r s f . . t I n e a M a d u a c z l s M i i s B o a a s . k , 1 f i d e G t o o Ü a h E h y i e n n r c i a s r o o c n o d t b f ˘ ( s , a á S e A t a B e d t i . Z t t n s w c c a . n . e L p b n ö a c c a ó Z n S x C n p H r i M e N n r o s e c r g C l i e h a i , . u D P . y e . v n a a t e T o . k E d E G a , . a n . B D , E E e e i d B f v l P n o u o e N e m n t q . n l M ( . i . B a l z ´ o s v o a S 0 J t , c l ö . e n r a I M : : s . y V i m i o d e u . M t a , l : d : i t , a M e h G p S . . i t c S p . m C j s C t A . h 0 a K a o i a e f s z n v a : a . h h p y d v c d J , G s G n a - C h t i , : y i , t a , K i u n . , s s c L z T s r n J f . l . i p g l , s o s 2 2 i t i D o r o o e n p r m r i l P ´ , 2 2 r V i k r , c i b t - . o l M a n d l 0 , i u : , d a e n o r a , o u e g i c i l e n k a a y e d o n i i o , n r r a i n C J , d e z a a ´ i e s D M a i f r a c t o m a l ˇ , g s 2 s l r P a o c c f t n n c h p g n k a a n l r n n s a w f i o b b a n u i e t o l s r v t i l v i i r - : : d e r i t e m i l r t : : i r l l t c e t – t e s l G v a g a t n o o a l i e s e s e u a d u r r r o o m i m e i H o g e p e o v t r r k P o i l r a u o ó a i i i a a o s o h m i i a m y t v m i i o r . o i i a 0 r h o e d o e h F B o o i e . c r s s r d Z N i J d n V s s w a a d L S o p r l K a a Ü M 3 c f T R C g u J A c Z S W m y T t e W s s A t d b . . S s s o o m b : e e M A t . . c . n . O e e h h r o a . . . . z e . o c h h . . . . . g e e . . . a r . e e a r m n o . n i h 7 r o . . c a r n n n . e h n e M a i h M i i p E f a f c S C M Z A B B O S C E S M D t H e S F a F D m t A o i s p S C A J S C D S C C M D A 1 Z c h g n , , s , r a , i s u n e e o s e z s l : r e s a t t k t o c c r o v e h m e p d i a i t n a t e y i x - h g n t c i d t u r n c t n i t : a s v c i g d f , c l d c t e n ö m b s e i a i t i o a l r s i i w e x t e i l o n u s u t b g l o y a d i h t t s g R t s e o a m e t o b n u y c r a n f d e o c p l t n a y p o u r . s i e t l i — s m s o s i h o t e h y o a s r a r f s f h i a t W n p s l y i n t y f r o n b c h t r p e t p r O i i a , e o o M p g e n r . s p L f — o p g - a n o r x v l o d u o a n n a g i d . . - o l o e s l e i s o . e t e o A d h t r o r x p e e r n m n r a d g e P l v i p o g e e n c n a e f a i n g , e s e t u P y r M r u v s l M a t f n v , a g g n r t o o a h b t o u g s l i o u a r n i , e u n t , n h o o a o v r a a e r n O f o t j e s s o l l e l b o i a e p m k p e o m f s r a g e a g s u e a r o a . i h r n l y g r s g y i f N b r e t s o e o r t o ˇ i z o o n g r B o s n ( s o f o s h t l a u e a x h h ’ p o r N l v o e d l B f g o n e o h u u t y y f h n d s P o e i n B y a l i p h l e , p b n u m , c e n o i t e f t t o n h f n r : s d i i h I ? y a i C o . e h s a c i i n a a d c n e s t k a r s i i r o p J k F o n d i . p y p l n e d v o a e t e e a , g G c w m r o c u . t l g a . r s h r i g p d e L y i i o d d h n o l h n a . g t l o o ö r p b — e r o o t l a i m t c t a s d v I , h r t g D c d c n s e M a i s h n e y b A , e o T i u t t , e S a p a o n d s s , u e b - d N n t y , s n l i g n i i t i s h s r i ´ S i s c : a r i d . e e n c o z d i . a a n u c a e n r y r k u c o o a i k l e P : O l e l l z g e n l r u v i s s e n r e s l e e t g v P f d a n s l t e l y : r G a : M a s p u e l n o g g a l a b d n e r c e o p e s a n s l e g g d a , d d l , a s p i o n i o e g e o ö f o p l b g r s l n w n l v h e n n l e c a o o o K u g - o c l a v i o s e i o a p e o a e o n y v g s t s - v l b t t e e e H P i i o n i i n I a t c a c s n u e a h e r l a h R y t c v y i C - n i r t d g e . r m i f a h c r y J i a i u y — K a n p r f . a e h s a h T p r t o c s T n y o t V j g . d y g m s r s s i i r h o c . 1 a s d : a W o a n : P o e 1 p i : a e i l e r a n . g J n i n o n l t t o h n r n p M m y P f v a n p r B t d , D e i : r o r e p f h ? a i p s e o a e , r e m S o n i l a s e f e t , h . t n r v s t p u i . , r b a h r i m s c M l l i t e r g y i g m s g y g F f d . a e c t u r p d o e ) A e r k s n - o o L h r c y l l o s e i i , h d o o o í n d o o r e s a o o , r o c f a s e n b p . J e i u K e z s e n l i , B u o e t ö r o l i f : c d g a ) e n r i p o s t i m m J e a n o n o c h , o C , l y s g d c s ˇ z g V . s y g o r s e r a m s e x i N y e n t i s o R n D . e t t i g H a a e R f m o s . n o a o t u . i h l e o n r n h o n r n U s m r , l o a C c i l i u f V n v J : y G i e e r r n . p e v c l m l R r l a o r r s o t r p . a a e y i o t f r l o e i p , s a h v a o . n a i i , d k g , A a e i e e M i t l ü J o l s , e a y i i l a F a n y t G o h n s v v x t W t t n r . s t o l e u b w g b r : M b o e , t n m e c , d n e b b a i s e a l h d a n a e v a . o o r p : i l m M a o i r e , t t s s i y r e l l : t n : o A r d n y s a f i o v o t n d p c e b e f s n i a P l V ü ’ d S , r n h i h I d t u a m g B c u h z r o r N c e a ó r f h a h c a I z s a o e h c e e n , e f i c h l . s s o s ? o e B t c a i p r o i s l : n . l i l e i i u I i i p . i m s s o t t p i y l t z s i s l c l e e P r e u f t i a S r e i y P x . F n y r t e u n i r p r y t o a t t h i i A r o y t s e b r i e . i m s d p M W a N F A e R , e r B M m d a E e o . a a k l h s r c h w i e A c h p d h n t a c o U : n o ( a i o e l n , . C c F , . - B a c c . d n e d . V d n . b b a o r s e d o l g p r o s m n a , s p o a g e ) . l p r o C s a o S L , u i ü f p n t i o o n a n E d e f p a i o C s a e R . e l m d u n o a e A o b l l t i f g s l t h m x i h . a d c m s M a r S i r . i n , n r o l l o i . G , e e h t r y l o s o i e n , h h o e a l i f E t o e V o c a . e t t l c y e l , S i T s o n g l a n a C , p a n g e A V o n P e n o m f h P r m h f i h d o l s u o a l e l r T d e a l k r M n c u : v ( , t v d l p i l e o s e n t , o t u g p s a b o c . a n i l r v n ë , W P b s i u n r : e a i g l n e u e e d i p , e a i f g h e r o e a n u t o e e n j c l D . s a o d a o d d : a a t k o m t o n D o i l r c T n G d m o v o n h o m a u g n d k ) o e i J n e s s c g z i , a r n p r ö . l m l M n m S i e i l k o e n o e b n . : s t a c n e a ü r , n p f t b e k u s o a t u a l v a e E o o y : e e s ) c v t e h : e . a D e o P g n a e a r a R l i z t d e n e i m l t m y o l c H e r u g t a t í r n e p a d D e t z i m . h o K a S e r i e f ö i . a p . r . a n l y v l i s o e a ˇ D n r o t C y t g z s r 1 r y l h . . s t a S n f ö s m n f h v M i m r i . d . A P a n S h r a a h a c B l e d h B o i t y l h a ö . D S R o i e d h d f o r f i r u L e D . e o n o y l t , a e h s n n . : , V h h P ( . c p . r n n W n i C T r e H i t n , . L e i b o N o C o t F P p g r s i s , a i m , Z k P . J d o S i s t ö o e g B t p f . i d D e M c s : , s r F B s r . o s . a e e a h . M a , N k o , . k n o k s i . o : W e e t , e s - i a n n k d : h e e d n m s t t k e g i B : r m , g : M o y r u A G O a r i c d n S : o a t l , A i n , a c t z t t s p i e a d t b c , a t S G n s d a b a i g i H e l W r U o l s o e i k r n i r i . , r r v F z e L t a O c d o , , n s e k . a , s c r o r , e m t a o r p 3 3 o t a o r f k t r m A r r . z . r o h s a , r e l e t . , a u 1 1 r h p o d r h i f t : b e e e n , o f a a r n n u i l l z u e t x i n w a a p e y c o e l f ´ i s s o a b y r y y S G r W b í b ? k c s a l h e u l r e z e s b n s e k d b C A e e f i z o h - d r n n n m b t b y t e s e r l g a d l t k n o n n c f : a N r h t g : : a e o e e u f s n l e r u n u n a n m l r a S a s n o : : c o r n t n e a g l o o G o a h i i a e R r u ö r r o i e o i S n g g e o a o o i s a g i r ˇ r a e a n o o s i i e r r a s r i a i i i i e e . h . o c r i e u o v n o a o i i i . m f a r a i . i i n a a e o t u s s i t Z n a f P m t P D J r r p i a a M c d W f t a K x r s s H h M a t o m c w n T m a a H K H i f e i s s M a P R M p s a r o R S R n l A m t e B . s s . . h a e . M . . M w c l o h h l . . n a h . n i S . o . e o e . u e e o h h . . . . . y a t t l . . . o h . n o . . . . e e V p e e e e o e r . o . h h h P C n n g f ( S N c N c P s e r S r V N S S C U O F S C C S C G S J A S L B d i c n b a i M d a t s t S C W M L A E I C S C S R A C P 0 0 5 0 0 0 5 5 5 5 0 5 5 3 0 5 5 0 0 0 0 3 0 0 5 4 0 1 0 5 0 1 : : 1 0 : 4 1 0 0 4 4 0 3 0 : : 4 1 : 3 : 0 0 0 3 5 : : 5 3 : : : 0 5 : 3 : : 4 : : 0 7 : 8 3 : : : : : : : 1 : 2 0 6 : 4 2 1 1 : 4 0 5 4 6 6 4 1 1 1 2 7 5 : 1 1 0 1 6 1 : 4 2 1 1 2 1 1 1 5 1 1 1 1 1 1 1 1 0 1 – 1 1 1 1 1 1 – – 1 9 1 9 – – – 1 – – – – – – – – – – – – – – – – – 0 – – – 0 – 0 – – 0 0 – 0 5 5 5 0 5 0 0 0 0 0 0 5 0 5 0 5 5 5 0 5 0 5 0 0 0 0 0 0 1 3 4 0 1 3 4 4 0 1 3 4 3 0 1 3 3 0 0 1 : 3 4 : : : : : : : : : : : : : : : 3 : : : : : 3 3 4 : : : : : : : : : 4 1 6 7 5 1 2 4 4 4 4 5 5 6 6 6 6 0 0 0 0 1 1 2 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 8 9 9 9 1 1 1 1 1 1 1 1 1 1 1 1

20 21 ORAL PRESENTATIONS: 24 August (afternoon) ORAL PRESENTATIONS: 25 August (morning) - ) - t d a n : o c o ) - n i e r d e y n o s a c a s m h n r a , i j l i i l d i d r i t i e l t i d t t u u e o t r a e n i r d g d s e o c a n r i s , d t d l r a a o r p m l n n r i o P , j n e o t p w f y s o i n i e i p m a e m l i s a m e y p . r d n m , t l s t s s a w h i v e a e g a i l v e l i i x r a m n h o x c o r g o r R e i n m t o d l a d s s D b n t h c o n t e i , c s E a g 2 h t n i i a . m o c i t n . g t l o n o l a o o D t a i a e o i s y a r f : l n l P t n a C l n a E t e a n s v e . c c c e : d p h H h e s a e o e g i l s l c i , A o a o r . . b t t . N y y r c l r l o t n s e a r , p d h a e h p e t w : o e l i n a a : r a m a e r o b s C a . c d o r n O c M t s r r d e a l e o g l m m S r c i e u m n h n i u t n , t t . e e e r d n l i : t a y d e p i d t g s f . o s o c J a m u e a r . o E c p o a t i i , f F n i n i t x e l n n S t r o e d a e e k c , s a , i R n a B a e i e a d y r a o . e e u d d n s . e p v e m i e n e e h v c s , a i i j R d i a F e f r g t t r g , a m y i o x c f h z r r m d R l c D . V i e ? n n n s e n N a e D o h r o t a o t n l a t t j e t u h s a a e l a t a : n . f i c o e i l h b l E . m , , s u c a r i c G b s ) i y y a y t y u S t a p m n a 2 e o S e o u S s k o r e u : B r ( a g E a f n ; g n l h h h i . i M r l f u n S e z o l k í m g a d k , l e r g n . r o i o n : s t c c S ( l s c p r ë d , r i W o a l . i t a K e r g s i e n n C o m t s e l l c a n A o m i e d e o o r r i n y y e e l a s d . e n t h e a e n f a n n n D t e i d t o c e e k e e , o n h e n t t e h i d i i e a O g a a i a m c a a i a a t B l . c o m e d d o h d c n e v l l d s i i M m f n R h h , e : s - e n J e e t i g a u a c c a a r r i o n m o a n c c r z l o c t a , R . b b o p u D n e S , , e p i t n v e i a n p r u r s i i h e e r a s j m S n i b l V r - a g t j l S e . h A : i h m m s i : e l c e t r o i d . h h t a - u c . m v e t . a g m n e n a a e e n a l o , i d s r u - u a r l l u n o e a o i r h p p p n s r M H n a d u m l o K R o e a l H e a o t t n g d á n i z t c f i c m v S a o u . i s . , e v S a m t t , — a t r e i p . y R k n l n z m m i d i s h ´ s e E c s i o s t e n z i s a l d r u r e a i a w 2 B e w h T c D N n s e d t c p i n u M a e i : e P c r u A A u e , t n i e i d e ç i i V v e c b o l r a s , - i g S s r a ; l S s s g r ˇ i l a a . f t b n F n f i s s d t n e n o : : s p n i o , e i e n i r z z ˇ ˇ i n . i : J l i . v d m m n e w a f h y r o a m y 3 3 k . b t v S i r c o u d d h i a a a : e l e I S a e i i c , g n s o . i t s o s c l l g h A r e s h S r c i s s r l m o . e n i ( . . h s r I t a P h . i h n l : e G n e a a e s a e o a i n , C p a P u b , - o t l c h e d A l C H F v e A m m s s y n r , v d p n o t . e e r i t D a a a h o d - n B B . r , u i t F i s p a e f . R : e s , j t , e d o ? s t u u e o n n w u t s i s P x t a L L e i - , l a l s n f F t i . . i i m . n i f e b d n p i n e m o a s a n v h o o t o e , u e e , d m d n . a s s i ? a g a o n r i i B , r e k V V t t a u m o z b s c ˇ o u p z s e z a e y R s s s T b N r i n n e N o o a s r i M o i b s a t n n r f e : : á e r i e t o e s s o n m r r g t i S m h t g r a i i s . i k l i . a i f l i a e r i o r r . a s o , r p p f e v t o t s a n l d u u c e l o o ö c i i a r s t v e r , c i p o n i e i n B a l l c B a d n i t B h i c c m s s c t l e a i t e a a n a U b v M n c i h D M S M i B e r b p t n t l m m e s s l , i a i a a P . H c t n c n . a e i p y f i i . . n l l h h n : h . , t a . . i a d o o g . y y d F e e a d e m . s a . . C a a h i h s n n r s o x k h k P d d n n y c g i t p S C C M t W d A G D S C F A V D r i m d v s . S O o t t s . e c d p u u n e p a i D f e J a L E . a ó r a o r n f E d t o M , m i : g ? m , . . U b e y p a - K S A n o s T d s t l n n P i e , n b n x g v g n l a . : d A e . n i . e e s d . a i c i n n g O g l o e , n k o e a i o n P s o t o F P : e l o t o e n h h i . l r a i b t r n a p o t n n e i n m e t c i n . M c i t l , h f L J , r c r p i i n i i h h e p t s o o i a s t a a J e s a n e ? e t a i h t c t s n , a f s a o r s r d o d o . , m m s r g r : u h c e w s n s s i t , e v e g e t e i r m i v v t m h a y p m t a p l b u l h o t i c n n n s s e a r r i b i y e g r o n u s o G z F c n a a o k r s i p n w o a c r n i e a t t e C i m t a a i b D n n a s r m e i L h d o c r e p a u , n a : r B e c e a s i r f u r m b u g t c c e d a a a t . t e o r z ´ s a s h S e p . c n a r t s s i a d e r c e a e . b b n o s o e H e K i a s t t t i n l e m e s G r h t a i e A t n . c z , v e v r m r N h o n n e a a m L t e e . t a i . n i s d a e M s p n P i . v i b s . M e i m d d i a t o i S u c t m e n r i s i r a a i . a e a t s , i f S s P n T l : i Ü . w o c a e s , t a i i t l a s C g i v e . n l l e v e e c e t a f m E H D r e t D l b i , r o , m , c t a . a z k m L O a A D s e c o s b . b l e g t C e o p g s s a D . a h h a a e y o p , z t o c e s u i i J n r e s a e : t s f s a r t B M , r e t a f n : m o s p t r i r n i B n i e i . a : e r f L o r r A n i t t G c e r , h h i a f r A e s b v H a r , , J o a i r w e l s 5 7 c o d g i o e e d g l n n . r i i a a , . r y a m u o . y d i n l u p : p p e u e a , r e a a u s g t t t m i n e e b t d d r o n t e P S e r r a o t e H S d g i i n c i i i e U n s s E ç t n n e v n r e i e y a w H n e i c ) e n d d s p t i H i t M : : a m m u h l z z . e m o i n E m e i s e o o b b e s n E o o e g n l t s r o f i i g g t c r r . ö ö h e l r e r a a n . , r r a i i l i i o e T i : u n i i A a a v u e o o o l m n f o V n h s e S h s s m r t g o r k p l e S R R F l i h a a , . h i 1 d w f f n C n a a d s M M R t i F s s s l a a r I t e M . m h t l t G z i S o g e e o a . H H f . . C c l h h r o t i o o . d n t - . . . f m n e e e . A r e t a c o h . s i . . . . n h n o e h o o i w s ˇ J t n i m i e l m e e : r — t p E c J D b J L J R l S C J M F D S C N E E S w C s t i v a g g e r r s a t , a o e Z p g i w a : o u h h c o l t n e o . L r n n c d r t l d n i t t . o g t n l i o i i . o n e i l e i e a . n c G f a o n r r s e R t f f T y i n h d a e p : R o n m i o . s A i t ( u t n z o o M n s , o o r o c p p f h e i f t t a t E s , e o i t o s s o w a i i c f l n i o y t A f d i f k e m o n e e e , l t t f n l t R w S i o a b n a d o ) n n . o i n s i e n s o o a m i e e i a e s r e s c a a s P t . . r j j t a e p a t m m l r A s c o o r i v i t s e e r i n i i c s n . - l a h r r D e a a p a L r o a a a i i u R c i c c e b t s r b p e e f d a b l f n ˇ ˇ u - s s e i N s v e r r i A . C e s , i M M d g s a p v d i j j t t r v e y n e s o r f f a o p i l l a h e p p g a I , t . u m a e o B h n e a e D p : : M c n t s r p d t B m o o r p s d e n . y u i n a p m s n e e v , n s p n p f l s 2 2 . m i a o l i e s t r o g e a b b M d e a e n n L m i o L r l a t . i h h s n a s a e m , r z f i g d e A s . e t m a a d J g a o o C t t i , e o o n . e a c s c a B h u j o o s r c t a e n s o , n a n c m m P L n a h , : u o e a P P s i i e h c i a s t q t i n i o s d o e e h h o e n n t t , , g b u u l y s a ˇ n n n e n n n e D i t e o n . . s a T k a h a t t a i i s t o j i i e i h l n l e t l i t l a a a n t g a f o b e e o i l d o o s r e m : a m r m i l r s s i ˘ i r a g a A a i i r K K e r t b s i n s s i v a e n e o t l T d o n n r t u a : ó r s s A r i n s o e g i v V b o o a a g r f m c r m t g e e s : : v n e e ( s s a K g l i a p b e r N a ? r k a i r l W o l l l s e s o s e r r i ) n a p p t i t o e s u g e e e u u i i m e e r a s u h o i i e h r i p u l . p U w i 7 i h D e o m s a w v t t o t c c d C h V s r n i a a p d A P m p p n i h i t a B t o m G d e t 2 v m m s s e o t v i U r o n r t t t a . K e p p . . r u u i i r l h e W a t h h o e o . v , . o r . . 9 e y y n r k a f . f s t t m s o n o e e s u . g . h a o o s 1 L a b p t e s p S N E f f r r t R C a a S C B D K W p c D S C J J v f S D H o i l n k d b ) M n e o , w , i i f a u . l o d a 9 o t a n r l n r n i e k o : r t 8 n e o t n A K a e w n a w c i h s a A d i e i a e 7 a r g . n i e i g t o o a r e l i 1 d n p s l b m W n h h r t g n d o r e g A y d i d s i i x t t i , u n l a a s u c n r e o a l a s n n i , h e h n a a c e x n o n r e o i s r c f e a d e — h n i a c n o f a p n p g i r i s s s i l i h i c s v B o U u t d n t h t t C a e I e A n i l t i d t s n o e t - n ( m l r n a e . m s a t e u i a o e e n n e a : e e h a : i n T f i i a o l g t a r a b t r n c o e 1 r e r m v n l e A h l i n a l p h : i o t s t n t v n l o e e e e t t . u t r f a i a V e a u n e G u l e x p e G o o g a e o o n h p I S p o g i y a n n h v l e g i ( i e e . i m g n n o i f c p h P e l M c e r t m : , i i d s r t u n i h n l a o f T M C s o d r a o e e a V t , i o l e t e R u s t a ) i n p a n a n s e ( n , o b : : r n u p n o s e t t o i y s e c e h a s s s l R t c s o s n s , e d n a a n y n l i a e n t h d r t e r h u a e t e s l p R i u i a n . r P o a e e a o i c e c h n i i e e c r g c i e f a p e s e P e r v d n N c i e n a a : p r p e a c c i V a r v s g o m S o t m n b p r i z h o : M n s r r i i m u p r e e e G e h i r d I . t e a o e t z o v m : a a a o f s U a , r t d t e C p M l t g h . p p z n f a g n c x m A n a m e o a i e r . t n n l r a u t n I C h . e I s T n r o S S a a a l e d n w i o v s c e h , s e e n a a S t b i n t i t R : r g . r i i t , : t F s o i n c h l n L T r s h n o t l n n u D r e l a a e , l m i s ˇ z i t r a n F n t h l t a l A m . I e d e a a a e . . o e e d t v n p e , t a d o n c e l n e f e e a f t . a . f p a e i m u l l s r w u e p c r : t i i l o o u e f e P e j l l a e i U g n o G e n i a a e o o n v l W l l a j t l i q e a e r N e t n U n r i o t . a s . n , s n e M c . c n c s h h h e o o a b B e r t M D i r e i a l e m A v A n J s e , e n a e i s e r A c A , i A D c i e r c i p m s . a . t : a C i i m h n e l d , R G g n g T l h v h p a l . r l i , e , r e e g . e a n c r h h v i V t a e á p u a n A r s r e n . . r g f p l p a o g D o a e i e v v e k d i e A s M T r S a a l n e i c , l n a e i i s a t h C r l o c h E g s f . p n n i r l r n r n , i e , N n L d G i s s m m o D p . r b i i o S s A F d o n : e v g i o p u a i a n , n t a e o ( f o i n n a . n , i a a n x r l i t t t ? . o D : t s o , n A g r h o v e R t o e l a s n a e m e k o ) g n w e v v e i i a o r i r e S l d A m c C e a t H e e c e k k s a a n a s v s h e ? a á t i i l : t o v D o n n t s p d r a o o k l o n n ö i e . i g , m t d t i a ˇ n i l G n c u c r t s e u r r a r g t p t e u t V r I I a s , y s n i V n A í k a l i u v d a M a e Z e n r c e F r e e a e a n o h l n n H a a e t c H u . a o n i a k s . : : i H y . o a i n S t i n i n n p p s . s o l h s . p p u h s z a o l u h r . i s v D , 1 1 r p i s f R a N n m c B l l a K R t e o B k t s A r d e o e R d c a s M k R k o e l n a n . i e l a a r o D . : . n . p c e . s l n o i , d i i i a , a W h S o k . n n M P i , n k k a l , V n m a c h W t - n a y n g e k t , R t S c m m i g E t n e a a K t z n , a a g ) z H H T i o N h o r a A E : W : R : a s ( . e l i f n f r f i k i M a , e s f g a i L o W e e e n I u u l r s , l - s m , n e s J n ˇ l . e e o c , . . i d t , e , s r e 4 6 i t o o i i w r r o n r I a o í a z e a r t c y a . n l : o n s r s : . l c i o r o b , c p c w l h b s s i l e r l y b Z n m s B B i o e v b b t i r v ) P J a d e g s o o h s z n i a á t s n c c e e l n a s a s e n n u r r e D d w f o o m a a e m e o G t i t t : i g m i : : l d z O i e s b n : : c s e n s e f a e l A r v a k e d t i v t e e a k h r e r i i i o o n r a e r r a i r n i r r d p p d r r s d e . e a S s i o e e o l i y a i e r o u a s n i i t i i t L u o a i c i p s i i r e e e e f o e n l g f d N p i r o v R o V S a c . t e s s b e a h B a a f e X u T c o t f l a D e l a a A R S G f . H s n B m m e s p p R e l t f i n Y a s s b p t M v F . . . . e a o e h H n v n i . . k l o h h b s i s a R . r . i D . o h h . h . M a r u y y a a r f . E e e i i u . c f p e . . . . a p e o . n e P C K r p a r B o B i h s C P W S C G G R P D C S C C I J J L a o h d b g ( a z D o S C G S M C V C S C M J C S C M M 0 0 0 0 0 0 5 5 5 5 0 5 5 3 0 5 0 0 0 0 3 0 0 5 3 0 1 0 5 0 1 : : 1 0 : 4 1 0 0 4 4 0 3 0 : : 1 : 3 : 0 0 0 3 5 : : 5 3 : : : 0 5 : 3 : : 4 : 0 7 : 8 3 : : : : : : : 1 : 2 0 6 : 4 2 1 1 : 4 0 5 4 6 6 4 1 1 1 7 5 : 1 1 0 1 6 1 : 4 2 1 1 2 1 1 1 5 1 1 1 1 1 1 1 0 1 – 1 1 1 1 1 1 – – 1 9 1 9 – – – 1 – – – – – – – – – – – – – – – – 0 – – – 0 – 0 – – 0 0 – 0 5 5 5 0 5 0 0 0 0 0 0 5 5 0 5 5 5 0 5 0 5 0 0 0 0 0 0 1 3 4 0 1 3 4 4 0 1 3 4 3 0 1 3 0 0 1 : 3 4 : : : : : : : : : : : : : : : 3 : : : : 3 3 4 : : : : : : : : : 4 1 6 7 5 1 2 4 4 4 4 5 5 6 6 6 6 0 0 0 0 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 8 9 9 9 1 1 1 1 1 1 1 1 1 1 1

22 23 ORAL PRESENTATIONS: 25 August (afternoon) ORAL PRESENTATIONS: 26 August (morning) l , y a t k t d : i d d a l e d n n c i f n o z d o i o h s e a n t a o t c r h n y l a o e t t n o y m a a s s e y i o k c t e l , t e g g r t e s n e s P a i n i t o a a v l a r o s n l t t o m i e l e i i P e e i n a r n t a n n d r t M i s i h o b a . i p a r n r e u d h c n v i c r a n r n r e e a a s a s t e v i t o a e n g a f M m i T z c a l ö s e e W i d e o s c V : , g u a t l m u r t L w y ( s : s e a i e s u g n - i e o n r n t e o a i - r r i c t t u n n e t u v p o i s d i R e i h B a o a s l i r r q a h s l t e a t t n g t n e a . e e i r c u i b e n o u u t a e r i n s A n h a l b r o f i v a n G o P ) t e a m f r : b u e s r s L g d o , t , a a a m h a n p s s o z t u n n n 2 d a c n f i s n e e a . n i o m i a g n i W u s a a n S e b i k i Z s d n p i r n z s a o r i . a s a e f d u i i l . m n b r i a a n u U s h n n n A g o a i k t a L g n o g I p s i t a O o . , , n n a ) i m l r B s o s S n r p i i d r , S a g n r s r a e i o b . e s u s t L i e R i o e c n n , o n f ) s n s u h f i l i r c i s i r R n a a a s d h : p l g t H u v e p e g i s o u , a a o n s p n f n e n l a u g z r t i f t g e n e d n p h t o o r a h a n b d l n e U r M i s s m r o i o P n l i t n i a a t b l o r e c t i r e a a r l s : r , d e j o i i g t t a S f i a t c r k 2 a P z f t s l a t o f k n a r s i i n i ( m c . n c u a t a i e m a s i c a t c o i i i s a s a M l n N v d a z e p e i C R r n m a s n r D e i . s h t a h . m o n D . e l i R e : l T J t d r N o o u r : - i c S n o s a B T i e t m r a i h y o : . i c a , . o r . G s i a e n t , m , c s k b n z y u n f l r o i s y c S t a t e m : R o W h N o Z h i e e r o r r e o g u r p v c , n r c , w Y , r a a v a r h e t i u i e n n n i i é y s f n a n o T i . i e m g g g l s d h k p g r B G ( n o s P n r i o u o e n h e n a ) n i e i n i i p t n . o e . R s s i c a i n M s . t t i i a l e e i t l n p P m t l S y o h l r o t l b t c f o i , e a s T i d a f w , h f z a c , e i b S o o w n v h h R , h v G n t V r c t o i o r k n p e r u a . l ( r e a e Z a i e d P z . S e C e l n a : i e n S S p s e I i v r l z s N . r e n . n . o s f i l D a s ) e h a e . e f p , a e d d d v a l i R c o W t B : n r s k d a b n D I p s t t u o e i ) n s l j s h r i , N e . i f , h t o r o a e s n a , o : a p n t a W r r a r ö P . u i o v s l c i w c r c u a f a : u c L h a t p r s e o a m F c r e . u l g P , t t e d t l b r s i o d b N l c , s n r s l l . ö . g e a e i l n z t M t s t h e w n l g e s e : e c e t a . a a e s - a o i e c a u g t h o a i o s h o . V h i n e s s t e r N e D n d C s f r c e M h u t A p c i c s K n p c r s o k n s e d l S . t i g y l t a e A i p i , a l n u , m , e r e p r l J o m t e a , S s r o s n e a t S r u . s n Z e o e r T l b m e a a m s i u o n B e o r c c : t k a e , ( r i r c . í C a i h S r . s t t ó t d ó l n g e e ´ R a y r c t a v c A s g e D d a t a w Z r r s . g f a h L S i , i a e t o m r A o t r d n t n t g a H i r l a . t : i u d f t A e : r . g i i e i . o n r n h . . r t n a K i e n , J N e t s e r r c s i r n y n , r P i g o m i u P n i M P i a n n e a a J t A i p r t y t o T a R E . n u e i l , y k F . a z n u t i s b s : s a e r f n c s l o n g P t : o s . b e i , i g , - c y s i , t n , N i i u M l e t r i d n A a r u e l o o s A h y G c z n a b n v W – s p l e b a l t s o h h e d f n e a g t e E P e t e D S r e i a u i s . a , s o n e a n v , e C 9 9 . r s r d n u e r p p n h t y i . r p s t n , n r u e r a n i d u e k o l a k h t n A t o m u n m l b a l T a i d C i t í l m o f i t l l e e c l i n o - t u e p c r o n n i p e t a m h á o , s n l a u v s i m c e h o c g A r l n : : d M u t t h r t i i a r u e s g l a r : a r s o o m e d o t u b r r y n a s u H d o a e p a r u n u . a t v , G e a i n e e i i e n o e n i o i i t a r t G A l r s i e M n o g g r s a o g r e s s p n e a . G p n M a a n e a h r i K C w Z S a e U H c i t r : c H a Z M 1 i s s a M t n p t m d h n - t n t e N o s g c e s d . T e . h n h h v . i i e . i . . . . . t u m e e s s e i e a . . f a s i r t p o . J . V t h i z a n p n u a r e ? e i o c A D N b w o S C G B d J R K S C B S S A ( t v D A E a d F . r l m s t t a m m a u t n g e r m g i e a i u - r o M m s n u i d r d d b n M p d n r m e H l e i e s o . e a a o e a o e x n ) o a n , e r e p f l , t r i c f k i o , z e h a o e r B d i 6 v r r i R a c u a u s u o d g s o p d t y 1 e l l s i d a , e n s o R P e a t i g C q n e r s i u r 2 b t p 9 r u e r c n R r s u a e r s , m o a y r : : 1 1 . n i w u y e c n o e f a h e r i e a t t y t o e p r n d n r r 0 n n c t e , t s u t R s n v s : a o n l . a r e c e e t v a a y i i 2 o p h p i n s t a m t H u o , 2 o r e r t v s g e v v o d e h o s e – b o é s e i d c s . n a f k e a e r l i t s . o o n I r p b h 9 i a , v t r r J m t e n i e e c c n H r r l o n o n t , i G o r i i e . c d t . s 0 t t o e e o u c r h v o t a , a z u s e : s s . A l o r a l g s e e a r t c , e a 0 b o t s p S N v p K f r e m y n i e D a a a l s 4 h s t o t a P P s 2 u n t t l n n a h . D t n n s e c t o o i e s t e i , d u i m o U i . . e a N i a n m a e a n s g h r , n e r n o e i h R n h t ( r w i p S g s i a S S m r p m e v i c m h m a a e t t o r a R e o p b h s T , i e e o e r i . z c e a i d n n e , , m . n i u a r h l i x f i p n t n : n t A : v r s s i r n n n n J n u t g g S l o o e y h g w i h a o s o i e a c a i i o s a y ? o i i N h o o : t a a t o o s t i D i , h i . ö y A s s p n d v i t g i i t t h v l u s r z k k n p o e t t m : e c l i i s u s s d t t c d n i t a a e e o a w u e u A c B p t a l d r u n c a u e r i a b v u p i a a l r e n j m r u u r l n r o i i t e h a t t g a o i o , m . r r o r t i P u o a a a u c c i t b a i a t l h H r u r a M M r c o t n r e H v t n s f t r e l S M g g t w s s t m c t a c w . e i t d i p i i o i r i i i f i c h . . n o W g o . . r u n n d o . . n e y r O n u m i n r a i e d t i n e o e p i o n g f , e f t t s S a s f m o m r f A m n i S C I T M C D C A A D L — m T . s e h i a d n e i c a e g . n t y r A e t n u l e i ö A s t n n T n S o a o V e , g i o c : i h n f i d f , l a l d l i r R i r t . m t e a , a f e r i n o t A r e s d y n o e s e r i . t x I n a y S p t t r a b e i r t o s n s s e n z h h a s t a f s i N ) , r e a u n n u d c x a e t c u D a n f t - s i o l o a r r o f p e f i r e e i e h o e e r o : f , D a o t e M l e t d , d c b o o f b c s s s e m C b v s s t o a v i e M l P m p d p N m n r o u a u v i a u e e n : o n e . n n a d n k n t . a l e r o p . r y n - e 1 t b a r m d m r a l o e k i o R s m r r c n n e n o l c o e i c r i t a e N r e r d a e W i i n i i l C e e l O o a t u . t r t o t Z a t o a i ( e a r s l t m e h v , u l i s e p c c s t t f b c , n i r m , i c t d l t s . y t h b o s n s n v a d a a a e y a a l a r A s M u s e o t m i T u o a s o x i l n t a e y R s e i n g u r . p l t n o e T e h : u e a . a n M B r Y e i e P o i p a d a a e a s t t e n . r n o h F d . m o t t t d T : h x . D s i g s n s N i e W b l t e r y P — J R r h I s t m o a e , s ( s e e t s i y e e o a o a , t i i g h s . ) r h e A r T l , r n , e a l s i i l d e c v u v t n c e t s i r d e . e e o e r t r : t n t h t i o o n g e B c t L m G e l v n m o a n o r n t f t e a u r a a d s a s g h i o c o o e w a d o , a i o e o i w o . i t N a n l o t a e s a o y m u t t h r a n r c C t t A s c c n a u p p e o a h a s P , i f a v I d s u o r o l g x p s M o S e s F N l r n . a g n . - s i u g e w v i S e p c e f e w n e w e i n f i n C e a a K a . p a e . b r ; p r f R l i n r : i e t B A n í k l c s o h i D V d o o e c d i u s S . e o l l p . n o p E n c o s a A , D n e r m s c R l k . x e m s n g u i r n t á c . i e e e e h e c i i n n h a p c U e a o i e i c s t . o s d r r P a a i m m I N M M n s s t h r t e t a c z D h a s n f l y e , e i T y e i P e i t b o s n , a i E r t . c r a t h m : , a o p x f G , c s v i D l v : c a l h h s e F v s A . n u , n s d e T y ö m s m s g e W o t x e i y n . e n u d r r i K l o F l b i n k P , . r e r - i c n r e a d : e C c a n r a r b e h o e f e a B d e e l w J i n c p . l h e , y i s C i y a a g i : p & h t h t f n n e e , C z t c v e a o S r o r t o r w v ) e t , ) p l p s e . n y t s c t d , l a i a i i G r n s s m s e a l r n t s : a n l a m m o x p o a t n a u g s e t i c h k F n s e n r e e w r A n t v d H s e , r o n u m o j o e e r n e o o i e i W i r s a p o e s d c c e n c t a f j o o e m c . d j u Y e e : e d d m o z i s P e l i r T i i c n l e m v o o a i e d o p : N G d a r z t S A e C f i b c t . u a e o a y s A u t h s o n t e r b g v t n n n t D M n v e e g G o n t a M d . n s . d v . e a K t r g n E s D . l e R x r e a e s r i e o e i i n w m r I p ) o s . o . n 2 . ö n l , m a g n n e a a r A . A a r n e i , r r c s . d c . r i p e J y L y i i t O l i A P u y u W c X t t c i T i : a A i c o . h f a R e t : r i á V v p d t c l M u ) T . , . S o A n , , r , A 1 N v x l n S l B e , i j t l s e . t m s s T s e o e J p l e f c . , a w e ( 9 . , t s s e S a n C s ó a e k P , n n o . a n : a A . k : d s h : a v , u v u i e e 3 - b l p , a a e i d k e t e n s a x r a a u i t d i . a : a l r n , v r D C r s a b c z E H T n 8 m G a h s t s z o g t a t M e o n e e L H z t n M e n W e o n u a e g r 1 e f s k o e r t u U . u w , s , a 8 8 a s : r r e a - - u . G e r e h f u n c . n , n t t i r o t d t e o s . i n r a p n o r , B e n e n n i a : d c i d i i s y d J m v a n b t n a a m e S Z f , c o p a b i L a g o s i í r a - d i e r l i g s r s e t l r m t r d n n b t l i u B r i l s t c n t a a o c u h o a : t f - h : : c e i e j s a P n r m s j a i i e c a e w i e g r u T T o e A a a i i o o q m o r a h a s l : a t r i d m M i r r ö s e e e l o a s d a e s e ( ( i j i u i o ( i v i i a t o o h . F i p b t i i t m c r i . W p h e i l f r n u e f n e e . u o c s s e a B i e n w e t u n ö a a t t c s f . i m f u t R M r A W N n i . H e M b F c p s s e n D d A d a m M p f M n . h e a e F u s n a P n h l W C o h h m s D p B . . . . w w . e u . n h e e . m S o u . y . e e r i v e . n o n . e o d h c . . . . p i h l o o . y e e e . . P C l . a p a t b c g S g p I S C J B E A A T C S C P F J L L W a s A a e h r n P N n t f n i t G A a S S C G J W E S D C P C t 0 0 5 0 0 0 5 5 0 5 5 3 0 5 0 5 0 0 4 0 0 : 5 3 : 0 1 0 5 0 5 1 1 0 0 4 : 1 0 0 4 0 0 : : 1 : 3 : 0 0 0 1 3 5 : : : 5 3 : : 0 7 3 8 : 4 : 0 3 : 4 3 : : : : : : : : 1 : 2 0 6 : 4 : 2 1 : 4 1 4 0 5 6 6 1 1 1 7 5 : 1 1 0 1 6 1 : 4 2 1 1 2 1 1 1 1 5 1 1 1 1 1 1 0 1 – 1 – 1 1 – 1 1 1 1 9 1 1 9 – – – 1 – – – – – – – – – – – – – – – 0 0 – – 0 – – – – – 0 0 – 0 5 5 0 5 0 0 0 0 0 5 0 5 0 5 3 5 5 5 0 5 0 5 0 0 0 : 0 0 1 3 4 0 1 3 4 3 0 1 4 3 0 1 3 1 0 1 : 3 4 : 0 : : : : : : : : : : : : 3 : : : : 3 3 4 : : : 4 : : : : : : : 4 7 1 5 1 2 4 4 1 4 5 5 6 6 6 6 0 0 0 0 1 1 2 2 1 1 1 1 1 1 1 1 1 1 1 1 1 1 8 9 9 9 1 1 1 1 1 1 1 1 1 1 1 1

24 25 ORAL PRESENTATIONS: 26 August (afternoon) POSTER PRESENTATIONS: 23 August : , , , e e , s s a s t n r . u e k e ? i n a n s c a l n r s n a L o e e c i a e c d l r n , o l i s h n i r f e n i h m , e t o u f t n k g r s o h a u t g t z u f a e a s i r s a a r o g t o i i n i n r l o n n n a l u e h e l b a a r i M i a p a b e n e - t t i o a c i a o P b i n e R i t h p e d o d s t o e t h P c f s a u b a r l m h z . i e r v p i n e , i c o d a s c d t p y o , a a d e e e p f d o i p r r a o A a o p r t h l o r l e n r l s n a w r z d r e a u a o a m r l n , h f p t b b h S r m t o a e a e o j r l n a p e a i l n B o i s a a t n p l A a l a A h h a Q i d a ( s t t m c s h u n o r s e ( ˇ l H a t y o a v c c d s e i t a w j a l f a o r d h c r i e o l w t m u i d , n s o r i t u g K t f k f m o e r a m e d a g o y N a L o u á a b r v o f l e N q n o . l n f r ( i o t s o i l s c n n b i n o S W i C a o e g f n r y a i e o o t s a h p b A d s d i o s f n i n a k . . l e g v k h r s t a i s C n N s n p s w , t c n t P f f l h s á n s e i n r t c a B i r s n i e a s i t n o u w - t c o o . a o e M i e z a . b o e s e l d m s f i e e t r c S o J t k o h n l h l t h , o e a i e o m K t i l u W t n o m c t t n a t d l o m o m t , : c t s y c o w b f a f , i t r : o s r c a A s t a a e e o S f u y i t e ˇ a i c n o , p a n o y a n o r m o r - e I t p i . p n a i o o m i s s g t r h h t e o g e i l G c o t v m s r t i e f s o y A a o n H s s G n a p s a e c k t g m . r P m e o l o r i u i r r o r a t e i e l y , i - e l a l r r f a t i Z e n m p l a o V g g e s c r r t o o a a i i s b v e a k d o n e a e c b s r . h s a , n e a t i n r n h t r f c o r n v p c s a i c i i t e i i e l t r h i t g t p r R e e P R o S p l e u o A t s v t h t t i a a s s n t s a M t m l f e n n c . . s . m o : n h i n , y r u p i n u e d r i i e a n i n I a p n . g o a ´ c l e s c s l p a g L o t r n M e h o G e n P i r S a a o o i s , o a ) n e r i m . c e d c d d n n o . ˇ i : a c i ? c y e c p t r i , u . t F e 2 o n k c a d h c t a S s m a a t t s r n z a n m r c s c á t v c a e o i A t p f d t l . u a i . e a a m o k n c a a a y s c e r D e e a a l R i s t n r r r d b n l a , a o e : a l e e f n l g r n s i i m o V C f l s j u a , b r o a D a g e n n e f o o i i a p h l o b d i a s D n . t t é M h n a o c o w o o s a n t E o h r i o , o a n t p a - S a G s i t l u e w h r s a l i c i r a y o p m . n A e c o M a r n n o o e t l i g l t m c s . i S A e e e p l n p n n e P c a o l O r R b l i , ã a B a g a a a n p o b e a f n i u R H m : h s : d t l o A i I s t t e o v t x i e i e f . r : t t n a t t , p n i a r s a r d T n n s o a : a d i e o n u e z , c e f a t s a a ´ h a r c r a e S E E s s a d á r u r i . i p o o s d a t a i r d n d p m u o e n a h n d C i i p t t i l s s s : r : o , v o o e e n d a r D a e t l t u B l e e D o l I y o n l f y a . b l r t c g t o i k r A a n p C e v i a a c g r l a n e , m n n a , s i i p i n d : . i a r i t c n a u r L c e a a P g : y e t t s C e a e o e s n d d e e z i r e n n s a t n A m a f e A i . r l i S o t u c C a c d c g u n e . e r l ˇ a ˇ n . z o p u ( c h i b t o c z J M a e a i g o d o i t o : t a e i i i i a e t d t r a . a e a n n a d i i f g v r r h A f r c , , f L g e t h n n f i a v x t a s N h e a t i L n e a r o l E d n L ´ r s i P a n c e M c o C , r n . d e n c s t o m o e d m i i n e i c , p ˘ á a o r a K b . L l h s p i : u . h b l s h a t P . e , o i i n e a p n v e e o o r r e t e p g r i C i n p . . e e c a t d R i k r a A m s B a e A v m a y e o m t u r e c J C d z M o b r r r y i , c o l g p P i u a n . m : a e . a n m e d g , m e i k t s m q S c a a r g S , , d u i i o i n g a ó g j t s b d . i ˇ C o y e R e c r F o p R n e V o R á i s i a s , a p g s n g , n a n F o - r i l o e r r í M n h d n d . H r i o c r n S . v l , y d h . b A r e S S : k u f a r a á i n i r a t i o . f Z , i m n o l o p a n r r - r i s i p . e c o c : l k t t s h s r P r f R S h i b r D u a s e a c i a a . z J a i e e a s e k e m e p a p k S a a M e k . e r . u a u r e r G s r o . r e l M . d t , t h h s T T K ö s P J s a i á J t u g e f i d e g d . a , ’ g P r n e D i B F d r . n n . ˇ a r n i f n k c C : , e i . l e P , ´ C v , v n c r T h í - o n a D h o e l F . h : . a ö r i a o i i e i e r o o ó d u t M . a a . k r . e z h n t t l : . t a M e o J t D , z l s r g r e v a N e l v t p d p e p r i o e o s a - : n p V L o i a G i M y a k o z o i s i g f s o a C s M l t l , l e u o o K l , c d t M - n o i ˘ a e h n s t o e a r a g . i , h e l ´ . r p n a D . i c i x c l u n f o í , g a , c i t t r k w , B A i l i J s l i r , D e e i i o c d n t i a s t a n n s t a n m M P a e v . o a a t A o ) s n e i s a n r l s v r ˇ . s U . u a e a , i a d a o g a e n e ) i i i : n C e V . c R e : l l e v a a t r a y n , y i e F d s o r u r v a g a , p 1 s m i N r s t L d i l a i n z r . . x r n r M a u a o a J N . d . t r m N m t e h n s r q C i u n o 1 i a i r o r h z - S , s e t i a . i e e l a u z t M h y t r . o a o e , n a u a A i l s p a K s K b , : y ´ p i A z S y ) v g s G t r p . r r M c s s l M c S a ’ s . a r n i y n e K T n n , o P s o . R H e . : r I d C a k S a m a n t C a . n t l : n , F i e r a e f , n . e a d - , a u h t i n a . e e o D y . y . c . l r r o ´ l e c o . d i B , c m n s A e e e g g , o I h i a t o z C a l k i s r o m i r r n p g r n d r s a , a C j a a M a t P i t E i A - , i s l i F c , i c u A H o n c a , a n e á v e H w e c o i m W o t s e s l h y n S C r , s n e j e , l s i z s , k . n , t S l f t k . i s ’ f . u s . o t D g l l b h f r p a s l h i r B a t n k E s r e c i e d e . . i a o a n y i i i o m n a . t i . - e h i e c a n k o l o i k D i a b n o á s i o i r c a a e S e o M s i S C G D P I S S d v b W H P t i n b i l l t r c á l m c n i h t l ˇ z n e C r n v l o s t e p f r a i e S a i e s g b l e z w v n e r i a e p e n u r o o t . z t o n p i e o a . t g r o n o o b e ) u a f r c m p n i c i I L S B L M S o e m b i K S s s v 8 e e M a e r R l K T t H a M u a D O o h e ...... e l C f G o e 5 . . t o l s a . r h . . . t l f u . . r . i a l m . . 7 i i R o e t n i t E A E p t t ( E M S I i s 1 a w k M D D M M M T P E J M B M M L S S t f p d : o e g s s r r n ? s d e n d c u f a a i s d e i r l l n n t n v s o t r o a t a a s a i i n , a i l t r i h c a l f z - p a s s d o t a o i ) t i r g t t s i m y n l o l i n f e i c n p 2 g n r c e a r u o d o r o e 1 b o i a s a h e e s e i c n t g n h n a r t n a t n y c n 8 t r p e n e a c h o e n x t i s 1 a m e n a e a e i r u o : h i r t u o i a t t o i e t e n e r l s t d e , r i p i h l i e u p n r r r u e f o m b o a e t a s t t o d n e a f u i g r p n r u s n o v a o i v e k c d m v w c n l o e a h g e p a E g r r o s c t a v e d u ’ a s n i a m l p a c g i u e e e p o p r n t r e n f c a t u t s o d L o i r n y o o n t a e p e s c b c e t y i f a t i o l e c r r c p s m n i e g r o s b h t n l p d a e c e a b f a i n e n o a e i e c - s D t x n n p a p f p i n v e n n d t t f c r g ( t e o e e a a h a m r n t o a E i e e w l a r o r t n d n A k i f r i i S s h t d e r o e h e : o m t o d a s a a a l o o n : a d e : e u l s c i r s m g z r l a r n l B f p - l i h a m l c v e i s a o r S s f e b d e l n u d i n c m r u i é h o o a s u n t d a h o m m d y r r n i R e o e m n g o i m p p r h a e c i p o l t a n r é e a ö r r : i a r t t M n a n v a e p o a s e l n e H g e o H ´ a c o c r i r m i R h c t o o o m t f t i P h t i h i y a . n r i e a : r m o b p . t s v t i o h f t n m m r : a p o g f m a p i n T f t i r i r f e u r m ’ e i u i l M u m a o o l s O o e h ˇ e a o : p e ó n d o r s - s t u c d t 1 s - l b r a c n s . o t a i s a k . t o i c t e l e e s l s T n e d J a a i i o i r o i r e r s e i l n v h l n a ) v g i i r i u y e . t e d ö s g , c i p L a a g a o M e m t t s i u s c m d k c m t i i e o i ‘ n r d n d B n M i i a e n e i i R f s x t t , s r t o y c l l m l c f E e o o n n r i g e , . a n d e p o a . u h n e r o u a i h l c i e s m e : t i p u o c e t y n p c n s h e r i ˘ B a d i t s t y v a D t o i h p s D t f R f r R W l e l h e c u s s t l t r e g a e f e o o e t h , a i o s v , . , i a t e s r p a n s ’ p r u p e i o g ´ n i c c A t t v r l i l h n y f s o r a n t d i n i u i i p a t t n a i e H W e m o : e t l n t C o a o B v g x e r V r l V n i i t l o . a a r a m D l h c R r : t D n h e . i o . o a r í o n u : e t . y 3 a a l s t h h i o , t P r o g u n d . e C J a t s n r n n i ? d a s t a o i R D i l m e t e b l e : ) M n n C i n y , , n B e e i e C r l l i a n n a l e i h d e W : r d , a a i y r p n , : i o n i n e e . e d a c M e o , a o a i i n i e i p c : h n n d m i F a n J a r j i s f a l m h t n t o í h r t i a a c p . e n - s o i a t V a u e r H t l y l t t d s i e o r , e o i i s p s o u z l a S m d t p t t r t a e a : t g t z m l e i , n u t e n P e A l y i f s i r . e o a a m g r z a l l d S e a e l r y y a n l o y p O t t a r e M e l . o z a n . r t h o ˘ r a s a i ( B r u i a n S . i p S í x s g n s n e P n s r M a r I t : c M e g G t a z a e n n , d f a a n . D N : e o r V x o , y s r . i f r a n i D a m r R o . G a . p a i l n r d e f , . e F n r e a r e . e J I n t o n o m d l , r u r . r , s t C J h M t o t e e C n o e n a a a ´ s n c , c B g e , B e e a u p a . z n l g e u , m i E s o , n i a i p M o h i t . t s h z t i r l t o : y l P i , s a i ˘ a r z n i a r s u v E á p c , e r t l ) n . l u t m p e e o t h h t a n m i i . h e a o r n i e r o D r o p n a a V l , F r o a c r p r o a t o i S r l p i F p c v t a u e n d a i a o i , o e t u w e n , b i H c l ó N e h a o . t p . a e m A u s t w n e n s o t a s r a n B s p m h i b n b i l t i : n P S L S h s e e s ˘ a h n r h . a h e i m B E i t o a V u s s a a I H e t v t t e o M ( a r . . , . f t E p u d g e e ö e B i - a n u . C k t , z P c m n r K a d W h r d e o T o n . V : a i v H m g o h r . g i t g o s y a d B w e j . i r e c , l i e h t a t P r . . i i l r , r d a T o 0 n : n M f S r J c y n s a r R c a c . n e c i i e a S s t M u s . T h e t 1 o d e . e D s , r a t o , D n i s p E l , c : n a d . M . c b o . n l e n s h g i w e ´ a a c d s . b c g W a o e i J . s K c o r o a a u z d a n i M , c r i E o n r i g n ) h i X n h c t i i o s d m e c K : p t e , n y R b , e i t v f i , r a n s a , r r o t i l o a a s r i a r , t d U l n . a a o m r T . g r i o s t K e a o i h c i l e a e a s M c s a d u r i a t a o i l p I L t a m . e s f l : a s g e c i ö B a n i A . e t t c c K t e n f l t . n o m i s n a s h , ó z d i . h e a r C a t p s E v F t b l o . L h o P , h t a a i n r e . t a y r c o e P a i i e v n r o g S a a . l i L . , e e T N n . . a h c c o r I f s . E ( v i S C J M G R C C a e t m T á d i o p h . n M h , S V h t o o r d . n a m s h a . H S G y M p R a f l f a y a r . r a , ö . t Z e e t S d M . S e y A s a , . r . f a l 0 , e n i c h 0 ( n E e c t , L L s f u ´ P . e c 5 5 z 0 5 e 5 , d a i 3 , o 0 i t 0 3 u h K c n r y i 5 x i t 0 i c 0 o 1 : , C 1 o H . t s , : ˘ a e a , 4 t l r 4 l 0 0 o 1 0 : a e : r C 0 n 3 a s e l l 3 , o : i 5 : : a r : l : n g n i n S 8 3 : l : , n i : e p o u g y r a y e c 6 z 4 s a 1 : n , n 5 6 d 4 a i 1 e 7 5 s o t h 6 o a i i c y t o s i n 6 1 , e i 4 1 t f B m k w 5 1 1 g 1 s l 1 1 i c t 1 t o – l i c 1 1 – h c a f O z n m 1 e h n a t n k i – – i n o r . s – l – g – o e e n – o r – m d m – r o l ˘ – a t 0 – c o p c g i a . 0 e – p l i 0 0 5 5 a A o a a 0 0 0 n 5 5 c m 0 n m R a 5 h i O F 0 B I i H R L o E H 0 S m M o 0 1 3 4 e a B 0 1 3 4 i D d r e 3 0 1 n d I e r : l O : r e : : : : u l : : : : . l l : : : . . f . . . c . . . . o . . y o e . . y n h . 4 7 5 4 4 4 4 5 5 6 6 6 6 o o p e h r n n c A C r r P a i s P t f E d i 1 A S s R 1 1 1 1 1 1 1 1 1 1 1 1 C K R S E A A B C D S F a

26 27 POSTER PRESENTATIONS: 23 August POSTER PRESENTATIONS: 25 August i s , l , s s l i n i r n d n d a n x n s n a n e e a a o e n d a t i u o o i i i u o , t s : e g n i d s i e t r s t s p d a e a b f t o t a t n n a r s i a n i e t p z a e i l , : o S e a a u a e n r i v o a l n u r s a i b i z h i o : i r r s i a c p l r b c t o c r r b s e o s u r n e p i s i a a d o t g a e l m r o a s f u r e u a e p i t k h r a s n t h F u r t y v l a a F n t n t v n r x r r d t s t m o a t i e ( a o i i r E i c s r t d o u e a i u c u a r l u i t n v h a d p l . e e t s t c d e d x m o t e n x s s e r a i j b n e q E d ˙ i z r h i e j s e i i t e p i e i i h i o r e o D t r u r i r o e e h e u S d s t n e f m a e o V s r f t a t i n n h s t r s ( c s s : o r r a f r l m B o o M e a a M y a c f x e e p a p o o e w m r i d b p d u B g r x ) m t a . n a p n e o . d p t e a i i l o h n c m h c d J e n r n n f s u s m o n e l r a S s c m h i c y e s c i i R y a m a x o c a t , f a a l e o a o n o i h r z c r n c e s e i h n u i a i l d , r i n n t t o s r r O i o e h e e a i t H t e i y a t e g d a t p a t h i N y t a B r z o i l d e p i , a a s N g n t S i a s b v n o e c s e t e p m t i t c e h i y c f i c r l E o s l s l i e s f y l s s i a a . s i o c e D y t t s l o d e e o N u e w n e r r y a E h y w s n b r S c u y t 2 o V a a l n r e r n l t P r e u a a r d n s o i r a i n r e u R a s t . d n i a , h y i a s p e t e r o p n h s o s c t o a n n s o t G e s k F e c n i t a o o i t n b i n c l l n o h g t a . e G i c o l p d a t i a r r . i g r M t a f i a d n b i a a a o e r o d w a T a a i L o e a r t n h g t l e d t c y t e n A r r t e d t e z p v c s R a m . i s i a n a : v p t b f t e e g n a p r a d i i l o a r , J i á k o s r i a s r n v d g l p l a b a b e r s B l r s o r e n e B a u e t r o , a d t e s L a S p o w m y u e a u s s m e l e s a Y u t p l e n a r d O : e a n p n a t d s s u o t e . t h e l l h b i n A r s b r r S b B o i p o p n f o c r a o n : s n n o m k a s i m u t i m i u s a d o R m g l i s o : o c a : e r u d i o a t p u e s r e o e h w h s c n d r e d s s o u a , t ) b g n a w t u s c c n i t e s f l n r n h n s t s p s e i m k i f e a e i d o e l n r i G i h e i t i a a f v s e r g e T d r h e z o l l a z g e l a o f z ’ a o a i w P n u i ) e . p g t e r n o s a w f w a o h s s r c n a a f e f s d s c u r o n a l f t o a e t s r , e r n T c n i i r g B a i r V m t S u n g y e e f o i r a H n o F ( t o g c n e P u d o o s R r g i d a h , c B o n : H c . p n a r t B a l i o a p : i . t n n e i e e 1 t i i i c r n n a s n t t m r e a i r m s ? n S H u : C n t y o . f t n e c c Y s o z g e a i t 0 n a a a s n r n e m t e l e r e e i r F a b l , . n k e n o . s P c r s u t v v n e ) t o i u a r s a t R L i a z 0 a e s o d n n f e o i a l n a l a n c . e i á a r e t t u e i f e r I f f . t p 6 o r o 2 i e u a n G e . C p l s l t h s t e a e c s o c f e o e s t i e e m C P e t s l n c 4 t i S o H i w b , h o g c n , e p , W o t g o l m E A m e b y l D b v h n i y i r r i o a n r c t y m 8 . r : o l m e r f . e a : h o n a I . t t r i , N e i n u : e e b 1 n s á Z D , p n h i u o i n d a i a e t c p t o e R s s o o t m k h a i L l a l o e r c t n O L k e i e h u - u , n í N . l : v e p M o d r e n E o n P . ˘ a e , o l l s u Z l a a t i e s o b n e d , T : m e p r r P o p á d u C h b , F t M d n e a a g O o l e : v s o h t b l e u u m f L h o m r e t : i e i o h M n h o v a i c p e r c a s . a , m a m c t o ( l n : C k f t t s t o P n m o o r y i a Y d r o r p G i g R n , f z e r A e i s a o R a a a s M A s k f t g : e r a . s e R s h s r é f f u a a r . C t n r y e : o o . n n o s r : n ) e p P e t n i e g n g n . L y : n w n i a z t i l f i r s a , d v a o o n a h h l . t n e ˇ a r r i N c t r . V e o s s l d e w z r o o n s e i a e e i n c l c t t d y M R g s a o r s e e g M s i s e i k i c e s l e n y e a h u . r a . n i h A M n I e e e e v a u D . a t e i n i e s i c s i t s ) r a s , L s F o i r c c z , t p l B u i w S b i u b z r e c c e , i . . e r , P w h u u P ( u K e a d a ) n g h . C K z . n r l e s n w a o u v k a H s O e Z , M f a n y t . e t i e l - i i d C o p . y a a m i A i t o R . e l h s o s i t t o . F o h r u p V p o c A . t , t , i a o s . r d r C r t L l I i s t n e l f S t , s e l o a s , , i . A . N u c o a s . d s l k u r l S m D e ) o l m d M r T A e : T p s s a U e , z l k e i a t s n m . M c p r a s i a e s i e : e s l d t ö e r R i e d p y e i . r , ( f e . é p c t , P x G i p C x t n i , R o a e a o c O r l c r r v e r m c r t n t l d s h P r - i h n z a p e . , e t h S e a o c W p ó n i a , á d r . . á a e ) a e t a ö l g y n . y m e a a h a n a F a a ( j C , S l N n o a : o f t , t B r k e o l l m a ó r r v C S s o a ) a r t , e y l o p e y f i A A a s n A T e , c t b e f : o . u k . g s t s o x n s . 3 g u o i y u L n h g c : g . M l e n p k m e r n , , a . c y E M i o o . t n a a s s c e o s m m P 9 u e e p l e i A z e i l . p f J t M , Z a m M i t á D K l o a n a l e o e t a o : o . n M t i M e r w 8 e m P o e - n s l , a C u r M a h : b n ) o i B i v i e ) , a C . P t s b , o , . i e l a r 1 b a v H p n f s d , r r i V - d r t o e A P , o s l r r i s s y f k , . c ˘ i r a o h i e d h l C a o . l n c i T C o , o e n o m d . t r l i m u a r s . a a n o e b . ) a a a r t a n o e t c ö n c j e t l k i d i n a a t r Z y á s o t B i j : n t a , o y í , a r d a o h ü r S o a c l a n n v j r a c d a i s u c i s a t o m t s R t b v u u e a M a u n u , e d i s i t e , h s z e H M e Z r e ö r i a a a p e n i , n t w r u y l i g A d o . s á . ö y n B v o c c v G s t o q n o n p h E t s t n r e . h . i p , a i i d V g á ˇ a o n N i r ˘ a d p d T s P S i . o Ü r V R e t o . Z L m t i a r B e . t a h r s t r é K s m e a o D e t m a . l h n u e . , A n A u , t . B . p ó t m i k d . v s V n . c c o r o . a . J z u R c l K , T n z K M . n a n i i i e d , t E a i n p J n y E a t i e r a , t n s e - i e , . E ( u u . i . a l M a A E a G V S s s M i . ( l T m j f r ( a a f r , o . N a T o o z h N i a c n , g e e n ( k a y o M f e , L i t k s , , , , J , i s . n B N R c o L n R n d n e s c i , , s e m i g o a a m c e g h r e d U e a n o m d i a C o , ( n , ( n s m s l k y l m G l s i n k i e o t ¸ d , t z , : i s y e , , n e l n n h l r i e a i m l a r l t i n t : e i . a o g d m i s r s y r c a e g z t o l M : y s i a u t c ó ü z . r s p n e n a e t i e u r i i m r g á n l J o a l n a t s r y r y c m g z A á e r i a e á á l i a h I o o c m e e s c a r , s n v o n z ’ r m s i n o e i e r t o u i o u t o b g c t y s c g , i ˇ v s r w u c h o n e r g s m s f t b s r m ñ p d , d H S e e i t t l e r t i i o y a y r j i r z e n d p O a d k a r n a h n l e t a a i a e o r y a s i é t l f á a a t ü l u o t t o p n á r m e m y y r r h é . . g r a é o r a e s a r p i i n u t l i c ) p m o t t e a o o a e o t d r r d a L a b e c r D l z h e e N S S S G H W n r r m m 3 a l a o e p e G b x K M G K I s s K B V o M Y l G o S p t P S a a F n h n l l o . u d w w . . . t S m K r K i i i p . . o o . o . a . u m 4 o k c . n . s . . c . . . . r . . . a . . e e i e o o i u o . f p . f e n n o g x m . . . a 8 e M n d P p c a N m a n o a p d H Z Ç E q r o V a n e S M i ( o P N P P 1 A E M S Z Z H M H B E S M D T D A B D R A m r P H e b H A s ) ) r e d s a n e n a o r h n s n i o l s t n d e n n o i r x e r d t a o w a r f t t i e n i o n o o e e f n t r o i n p n a t i i c e o i t u s a f t l o o t t a r a t ® n s g a S e e n l n r s a a u c y a i n s o h o n m w l p e h l l f i a e m e r i u o s t r t p s p a o a s c c m t a r u u e a f o l y e i o a i m s r p r r o l a t C t t n t a g t t m p t p p h n a n ( e i n a o e x s d p s o r a i a n t l i n r l n m s o o i c n w t v o r e r i e p i n n i d o i e k d c t e a e o o u t p p h C n i i p a n t n g a t i u a r r r o c e t e i i r o n e n r L c i o a a o p a a C p d i f e m c t e d d ( t n i d a o z ( m v d i f a r P v h a c o t i n a a m i n n n c n a a m i s l t l o n n c r l n i a l a i l l o g k a a r d p a a l a o e a d d n a i n c o h o a r t a h g e l o r r s i s r o t t a t d d u b c y o a a e r f l S i n n s n e e n a t o c r i m h s n o n n ( s n t e i e i P b p t p t o o z c o s a n n u u e p s a a u i i i a v o i v l a g a r l d l l s m t t i o n a o t y t e a m n e p i d g i r o a r l s i d y m m n e a o m u u l f t y S i d e e e o x e e n d t c r n a n i b m n c : s ( r t r o b b o r G e d a c o a N s a e m i i n n o o d o w e y c e g n t c t n s f e i l r r e r e o a n i i i e y l h o i e e v e o n r r t p d t t e i t n t e l e t c s p o b s s d e n a o i g x s f w o n i t m u d m o i i i n i r h o i s i a t y c o e t i c d ) e o e c g o a s c s c r s o k t d d N s t ’ p p i s l r h i b m i o s f i i e n s h e i r n t t n n i t c t y o n t l l a l t t d e l i a o s d l t c s y t e v s i a s x t o n a a e l u a m a t i c a i i s e r ) m A b s e i i i i u e b n c t c e a r e r s a r m d : t t u a n n e i d r D : a a p i s o i D l r a a u g a a e e m e i a o d m b r n E l a a n l i m c l n i d u r a t r i e r i r t : i e - v i f n p p s l b a i a a r m n f s e a a m i s i i a A x s h p r d t o s c S S h h f r s a t i e o e e g u z m f u e r e c d t t e f a h : s ? i i i z s : : o a D s m i p s v v e t i r e o e o l h l n s h i S i c p t i y f u i z z a s r i R e t i a o t c p l e e e o l t i e e c e a e e L t r a D f o e r v i b r n g r k f c : n n S d l l f t t m a v b p r t d p a a a a . o a : N s t b r d p é é t f i ) o o r n r f a : n o s a a e i d a d h h e e s c e u n a d a n a l v n u o y t e e d r c V V T s s i e c e a M a m o n y s t A h d i a n o o t c ) l e l u g f h t d h C . . a s e - r a o t a m A P d , a s l B f c i : a n t e h n t j i n n t b n i r m e c a ( l n Y Y i l . e a e a L t t n e : e i c r m m : i e a a g e í z l t t u a ˘ ( o a f p c c n e e a a , , e a m a n i ) r g A f i m i i f b d e d e a a n r T i g l a B e r B i z l c m s g o o : a e . i r c C a t g z a r o n k l l l m m b e h e e o i r d s : i z a u s . o d r s l a t d g e n o : r o i r i i o r u o n u u I h n a ˘ r c l v c i e f z u u a n T r g r M n z s l r C i e t i e e F m r t . w t g g m a V e g o o s B l a a n o e a : ) - o B i s , n a e i r d e z r d p n i L f G w c l n i o n n . r t g g n z i a i y i a P e D t A a a e b s n l p n s l e n o n y r l . t n a e o E v r e n t l i v i n H A A c A s i C c l i t , s i : a P u i e P l t c e u y i t p : n s u a b g d n a d . u a . . , n c e . s . u D t í v u a a i r l a o y c : v i e n T e I I R e v t t n n t e E a n o s d i l t e t q a w , e m n u a m h i i p A G h n i r . i e e r D t , , . a o s n o e f A d h a s : o g r t l e v z e e a c r a i p b e i f o , i c , l a a i e e r s T , t c k á e a o e s E : i t m g n t c R g m v y a r r H t e a v t y t v J e t r n m i a í p a t h , ( i i e a e u l n i m , e e e e a r t M : t . l n f r a l u M a p . t a y o o l r T i a n x r e a s a H i r s q v v y l l d t s t y d m n r u a r . t o u i i a k o g r : . n t d r A o E d R k i e a y a c a s l : e e u e a V n a F n m r r u t i i u i r o ( e s á p h é o n ö h s i , , l a b B R R R c y p a ó t e e , s a e r p c . h t o x s , t z p e R n o i t a C f c M t y s n h ó i a e u y e , , e k c i z e e a o i t t e e i ó v z H S M B y n e f p m z d t g . H s . z o k e F o g s a s o b d g r e p h n d e f i P r i p n i d d i H L . i . J s . y o , r c t i T o i s e e n s e ) r . e s o : t e A ( i p l H s o P s l g o s : t u a a l s o k a e r , w d . r e a H z z t c ( . R t r a s C l D e d e o A R W f a M o ë , D d d n c r o a e g e e n a l t a s c p - p o . l s R K c i , C Z l P p n B i a a , b e , : n l o . p e : d M e n ) a t n o a u u o d ë y M r d t . . a r s a y . m , i i A i i e c z f w x l . . o o k M o n o d n e o n g g n n . s i d K s r o e i l p t t J l R t i i , o t t c o c e í a í í i l r u b d l , e S a t s ö t l Z C a e r e B t D d l a r s r i n o t . n m d y r t r r , o t : t a l r f b k . t t l g a o p N a . a u n r r I r , , a e y n s u e a g r A a , o n r á a h o h d d D i f é o l ö o . m o o ) u c e g n u t a S z w e s c t t t e e o , a r h E v G k c z u n n i i m á n o o a o t i s . a t p G c a i o o g H D r á z p i y E s q o , i r T i l a i c n n p o g a . t S f f e , l l a s n s a b t K d v i n R a . M v o e R R . . a o e v e , G Z . ñ ˇ B t l c v a e e V l á a i o o i n á n a . d A l a r M K a P ö s i d i R o J . n l l a . s . i g n t . . P i i s r y f u . . o . t P r d d S n i i n G o P h : v t d J . , é e , r b a s v o a n , M , a : K b A d a a k ) r l ö i c a g I L , a y . ) A a o Z e n a c , o O O d a . k o H P n r , g r o r M i ´ i l , M n c a a o . , o P x p p n r ( m e , u a t o m , d z c ) i s , a b , a e h y - V o r r , , o d t á h n , s s c o f f r u l f i a i K s i m a s l i m y d c T , g o o z i c v i a n t n n z l . l h t o r d a d r n n k n z o o a i s e h a u c í y t s g o u S n , e g n , s k c c o m t J i e t í í e o a s l l a a p n i l e n e a i a e m i c i i c d n e s k r t t r o a b v e o r z b g T á t i l o f e c d i , b y y p p e p u n o r d i ö b z g r v r r a r ) é i s o g e n i i a p p m t e c n s o o r i t t o r h . i r r a o u n a o s m a i o t v U o i t z e á a a i i á c s r a o a t n j l o o n g a o a e o t e i p a r H L E G s s u a a f t m i n w i a m B P S s t t . F D D r C M B l P D K i o h t G V o n W S a t h e n p t T a a o a J S e . . . . n n u h R U M M i g r b i K a . . . . n n e . . . l h r s s . . e . . c a . c . . . . r e e s e f f p a h . g . . . . y a y c . e R S u B e w n n a i o u K ( s s T v S l i G o s s d s d c N r ( N R b e t L D E G G I J J K M N N I I B G B M T E F M M R C t C (

28 29 POSTER PRESENTATIONS: 25 August POSTER PRESENTATIONS: 25 August s t s s n n g s e o , o o m - i r s r o t e m f i : p n c t a g n f s e i a l n s i a i s l n o n i e f y u i k y ‘ s t o s n t : a y p r u g i m e t e o e k s C o g q i i o a - r v h ? a t r e v p n : m l s t l f e Z a i a i d s r a t ( t t s h u v a f v s o i a a c a i r ö o u ) B s o l e e g k d h e r e e e P B e y e a s m s c h n a n l u i l k . t c n p . e h r t n r a n d a a i c i a e e f i o i a p c t o i r o N n r t t t i V t W v o a a c s s c r b T e , r p p . , i l e G e s a e f g m n c d o g a n c h i o D n r w e e o o L n a n a t n n i t p t . k i i g e f a e e s g f b L o a n i p g E l o y a S m n e o e s ; a o h t o a 3 , m e S l i . h a A a r c s c y o t i N 4 o V i u : r i e i : n R G r z l T 8 t h c s a f c . i s B d 1 u e u , : : s f e p a o o ö r d o r r E a a e s , a n a d r e f e L p r e e n s e r , S e a l l P A ö e r o d ( s a L s g g n g g o i t m . g f c d a t V o s o u o e P y k n n n o l i i n o c ö e i i . i n a a i g p s c t , y ) d M i J g r s r r z s e a R o R R a n e h s i f t . e l e b i o , i z g n l i e . i e l . . l i r P S b o t A h e l i p l o t F y e h . i E E c : e e c a s k D u t t e r h z m h s e p e c , , b a s u , d a t c D e r r e y h - P , l i a p o t t i e e m A u w f , y m L s v : S l l u w c E r t i a o r N o q o r r i . g g i e n e : p o l c r a e s l l n . P r e n r a n n n o n o l á y i h r s P i e i i d i r a h , I e s l w e o p h l t s s a s . i s d ) y a R R s v M . o g a e n a p e o l i T v n A l t o d F n . . . a L e n n e i d , L e i o i r P m ( G u , o , g a h t R i i s o k i . g t d t o M M l . m f h i i a r l s N , L o l S P e o W g i n , , u o k u y b p t : n l l i e o o a i c s , , . u e a u i r h : i s s l r k y g d d r h u i n h t e b o c í a B a r k e e c i T M o s ö ö - i s e t o n f l d l c , o a v b y a . d o d e M ˇ g n z t a y r m o d t u H H n d i s S g t e e g o i n o A l h i m t n c . . n B a c g a . n n W b n v a r a , y p h a d a . b e y o o o h . i u L W W l A i G H m a p i s o W b y d S m t y D o C , , . s a C c a r l M . . o i n y a r n n , s l P e i c h , i o a d n , s i t . r e e V a C l r n l l a o i l s n h , i P o n M e i i a e d r , h g t s o d i o e s s t l s n l i t : n o t t M , y t e G a o s g r n i a a p a r s r t u m a k a n r n f i , t a b . l e a r l v n e e i c H a S c l w x o e B h m u v a o i L E A ( a h s l t k G s t m t s k i s k r u t . s n d t . c r r c a m I a s c o e i , i o r a a n P i d n l M s i a a a ö i l v e u : d A d g e h o e n o h t t , s u r o l i . i u e P t i r e L b l l e i c , n h n a e H i n t o G n h h t A z s l f u y . T u a K e b a c o . t f P n l o ö c . a l o , D o . k f S p h i i i . M h r a c l P e r o n T . e t t n e t a g r , o , p l h s r . P r e K u , g e a , t y o a a r a h J n g f l R N s d i . z d a o , n A a n C c r c o a ) e l t n t o , u g i , i T n l n e b p s e d c n . r y a , ? a t w s H h l u R k i p o a p P u ) y o a o r e o b u í n e l - t a r a n G , o w s i o l a N e s g o k f , f k a b n a r n d o n e l t n i g i í r r : k i m u e m a i p r a r g ˇ z e o ’ l l e t t i c r a t e ) g o u o n i c t t ß n c m p o y l s n a r l o r g d - s a s a N i g a h e e a o s o o u o a a a g n y n v e k a t o t a i l u r a n l u r . i p p a t l s a L i l d u g e y S r K m T M V G p e G H y a H l T W o a r t . a L n G R e l e e n . . . . . h . . r o . v a h . e e . f a . . . m A h n o I A E l P P h o H i ( P J K M R P A K K P T S T V Z t D T ( c ) n : s ) s i , s e o t r i u a i a l n n a r a s , t i i f l t i d o f : r o c n i u i r i o u l t i a l z e t ) t t p d b s s b c t a a r s v P a i i i a d o u y , t n n c i e , t e r a a A l l r r t i . b r m e a n t r c i a b e t s p e r P o r . H F e s w , s a r l l r a s m c o e i J t v o y y o m , i r e : . i e e a g t a , L S t e d , z s e r g : h c s n f a S g i w n M d a ˘ a , e U a v a k e l ( n y s p a t o p d i d a i r a g . c E P n e , s a p r o f l i e n J N i e w d r e n r u r m s t l I r . a . a n l n l e i c v a e y o , . g z n b E c a , u . r l P o t o e F a i e A i i ( X k O i t i o s T s l a u y A t t . p t . t , e Ç . e s . a m o C ( S s s f e m e P a J h s a J ˇ e c s u l ( g , f u s E i Z u o a t p c n , , p a e u a i , l b t d o i o h t r r l s l s e s a c i i , n i i n s e i c u e i i a n r n r l u u g e m e l e e k M b s r H g k t t a k v e i í g t f n r n t g d g r d i l s a , i t i i . n c a d l i g o E d F a l t a d r i d e c r e ö l A h e a i ˇ d z e r d o c : i s M u s t s t i D a n h w p r h g H a o r i s t e e c . n f u N o s . o e e c M B l v F o f p . m e u á P i o w n i G h p h n S f J s e n t : a D . e e t p M l t , G i u b . . s s m e . o g k p e x ´ l n h c h o n , r r A . r : s o p i a t e e T t r e n a a N e n i e D i H d a a h i c r v i V f c e e t , r o i f d t . e r h , , t e l A : n f S p , p n h l s s o j w z m n S s d h l f M a ö o y l i c a y g o o a . ˘ t a a o n o t t i i n , o n n . r v r y k h l A L c i d n r S r g d s a e a f o i t l s a a o o i a n l f u i l h r e h i . a i y l i c t , e s ˇ r a o n g I l r g t M I e h a i o r s t i p h v s E t S l a e r h h i a h h r o a z . c e n a . , A r a c n b t l o l a t f a i e e , b p n g r e s e w r F o l h u e u r o c S e i c f u k o Ç v a z á e t f e l A u h g t r j s s e g , i D i H i r c a e m n . o t , i a o c d o n n s H l s i o m g r t n l L v g l y f . . e e u a t g a s e r A x I k u u m a e f J f n y . o o s H o l L y e t h a d i t c g r u o n u , t . f V t i o m e t e M y r i a : . d t e e n a o K t n o E s e l x . r g B m e s M h i P d e t , y e v a o o o a D e g , r e l n p a i f r v t l F a o a o r r y . ö b . , c l k b ´ i t l r l i c e , D n a n r r a a o o s e n m . i t i r t t i i f s p c e c u B i . y E i v e l f n t d p t a e i u M r e M v t n u k h m e P . h n : i a o l r E e a i M a e n n M i m n d d , m J e i , a d l n c o . p g h i o r a , r d e e . b s s o p r o , i d l b r . m W c s m e s c o e s n e e l t i e c á c d n A e n u r l e L t i n ö s e r f a , n s e M A a l a t r t k e t h e f D r r d o o i i , a n d n s p a a i s o i n d R M i f , N c ü : e i p e r l n p f c i z , : i a e h a a s b r l e i o d t g . e h o t e r e d d n i l v a u k o e c l e p D t g d i m i M r e r u r n m : e á p n G o d — n s l l f e a p i D ˇ r z l f t Á a e l s i l a e . l m b i m : A ö P l l o . r G o o a i S f h n e , o y P t d e c ) . h S o i e u o l a t l : i v . u a t h d t E . S R . e o r n r h s n s l s r y t s I É , a h e c u P f u r l a n f t a , . a . s d c p d H : K l , , n n e ñ a e o H J a d c a y d e ) l h S a e i i l t t A s n n . n n a i M , k ó r l e D , t n r e i . e s . i h e C t e , a f r i o u g a e i a T m C a a : s a i g h r o r s í , c f n r t s L r B : r i a i h d r D s d , a s c n e . r o g t M e d i S o L a e y t i a B i , . c i e a g e e r k A c n e s k h , e f o c k B u : : t k A e a p e a d . g h T o a f ´ e c e n s c . g m e , h s a R ´ l i c a C p k t r i t m s i P , s n a c f e l k e h t s e e i o í a d L s i o v i R e f , l E t L l t . o o h k e a c n o r , g t d i v n a g s u r t m s d l e B , e l o l i c t a s o r c F , a a e s r z v u a A o a n A ˇ t z e o z M g a n e v r o . u S a s i r o ˇ ˇ e S o z a a a i n . . k i c s a t s h o P t I K P B l o t f l z : n M b a i r r v t o i g f i . i d i x i B l o v a a s e c l C e , t . a g ó l D l m t . P n r m y e e M M o C l l n N n . o e e i z n c k J o d r C l . o e b B h . p R E i G . á s h a h , , a i t y g e a . y t n e . e y a E e i l n R ) t n l i 5 e C . B t A , s . f r A P e S o h h e p K s z G r a L e . G h N L h h e 9 . : d y R . I A e e e . n R n h . g V t , h o . a R a a t S . , . ( , . p , 8 r I d s t a S F i e e n r : : d d J : a W , e n P J , 1 d F K a . a Z i y g m ) M z h c r r i M n r N : i a a ( : v a k o , t g a r i , p , l h s i i o l , , , n , i c e e r : d t o s e z z l e g , r e t u K e , r , o t r r i i t t g t s g s o a ´ t r d c u e a a i s a s e s n n n n d d a e o e á l t i k , a i n M a l e r o t e u s e ) s g n h m m m i l l d m r a a g g n s o u g t - y g o s k o t l r n c s e r n o a C n g t l n i d a r o p h l a a l s h ü u i s r a d o i o ö z u e a p a a e t n m a g s i l r e a l j i e e t m o m l z l t b r o o ö r o ö ö ö u u p a t e i r z e e e t a l n o l s e c m e i a l W W o u W h u h r ö l c l h g u t i c C B h B e Ü l B D R J D R d T M y y r t u P G V a K u L M e y . . t o . n d m s s x e n A A V . . . o o e e ...... c i . f a h h . . o . . . h y h h y y . h n f . . v f c . m A N r n m o P N ( F m e I B W u m a s i W e B P ( D D D P B W P A D E J F S C S D N P P P N V T N

30 31 Plenary Lectures From historical biogeography and shifting range borders to Functional and phylogenetic insights into squamate genital structures, phylogeography and dispersal with a focus on chameleons and monitor lizards

Jan W. Arntzen Wolfgang Böhme

Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands; [email protected] Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; [email protected] Parallel species distributions often reflect similar ecological requirements and are frequently interpreted as evidence for shared evolutionary histories. However, over what time period The final conquest of land by amniote tetrapods was not only successful due to their kera- may this hold true and how deep in time can we look back, really? Early on in my career tinized integument and the calcified egg with an additional embryonic envelope, but also to I adopted the comparative approach to deal with this type of questions and I worked with the possession of an intromittent organ allowing sperm transfer outside of water. This organ pairs of amphibian species, such as in the genera Bombina and Bufo, Lissotriton and Triturus. called penis is plesiomorphically unpaired and is retained by all chelonians, crocodilians and A keystone for fixing the history of species in time and space has been ‘enclaves’—pockets by primitive birds, and—of course—by mammals. It has got lost in modern birds and in of a species range geographically isolated from the stock by the other species in the pair. lepidosaurs where it is still missing in sphenodontids while squamates developed a new, In my talk I hope to show how my work, from historical biogeography and shifting range paired structure which is called hemipenis. Hemipenes show a great variety in shape and borders to phylogeography and dispersal, has been inspired by enclaves and other remarkable structure of both macroscopic and microscopic details. Some of these structural peculiarities biogeographical patterns. have phylogenetic signals and are thus of taxonomic value while others are closely connected with function, including behaviour. Lizard families with a complex social system and evi- dent epigamic attributes tend to have uniformly structured hemipenes across species and even genera while others which lack a marked sexual dimorphism have much more diverse genital structures makimng them diagnosable on species level. Iguanian families are exam- ples for the first group with the chameleons as a more complicated case. The second group is best represented by monitor lizards which can often be diagnosed only by means of their genital structure. It was discovered that, as in the penis-bearing groups, also females have a homologous, paired and miniaturized structure which is called hemiclitoris. The talk gives an overviev of squamate genital organs and discusses the problem of sexual selection and structural stability, also in view of respective organs in other animal groups such as molluscs and arthropods. Moreover, it correlates genital-based phylogenetic hypo- theses with those derived from gene sequencing as compared with traditional ones based on external morphology.

34 35 Invasive cane toads in Australia: the impacts of a giant frog in The deadliest lifesavers: reptile venoms at the forefront of medicine a strange land Zoltan Takacs Rick Shine World Toxin Bank Initiative, ToxinTech, Inc., P.O.Box 6266, New York, NY 10022, USA; [email protected] School of Biological Sciences, The University of Sydney, Sydney, NSW 2006, Australia; [email protected] Reptile and other animal venom toxins have evolved over millions of years to harm, im- Almost 80 years ago, agricultural scientists brought American cane toads (Rhinella marina) mobilize, or kill prey and predator in seconds to minutes. Accordingly, toxins aim at vital to the sugar cane fields of tropical Queensland, in a futile attempt to control insect pests. The physiological functions such as nerve-to-muscle signaling, blood coagulation, and blood cir- descendants of those toads became pests themselves, spreading across the Australian tropics, culation. At the molecular level, the toxin-target interaction is characterized by high affinity, and even setting up satellite populations in cities as far south as Sydney. The toad’s invasion potency, and target specificity. Consequently and from the medical point of view, the animal has had devastating impacts on native wildlife, but also has enabled scientists to explore the kingdom’s toxin arsenal causes up to an estimated 100,000 human fatalities a year, yet also evolutionary and ecological consequences of toad invasion, to learn how a tropical ecosystem posses an enormous potential for clinical therapy. Today, about 15 pharmaceuticals are has dealt with the novel challenges posed by this toxic frog, and to develop new ways to com- derived from toxins, including first-in-class, top-selling, and major life-saving agents. They bat the invader. are used to treat a diverse range of diseases such as heart attack, heart failure, hypertension, diabetes, and chronic pain, and are taken by tens of millions of patients globally. Most of these pharmaceuticals were derived from snakes, in part due to the large venom sample requirement for existing analytical methods. Recent technological advances however, in many although not in all respects, enable the utilization of tissues instead of crude venom, have reduced the quantity of venom/tissue sample requirement, and allows for high-through- put processing. Along this strategy, we have developed Designer Toxins technology that per- mits the creation and screening of target-specific combinatorial animal toxin libraries based on either, the toxins’ amino acid, mRNA or genomic DNA sequences. As a result, we have identified a novel toxin selective for a subtype of human voltage-gated K+ channel implicated in autoimmune disorders. It is estimated that over 20 million toxins remain unexplored in more than 100,000 venomous animal species found in nature. This highlights the molecular biodiversity of venom toxins as an unparalleled and a valuable source for the development of novel therapeutics and presents yet another critical dimension for nature conservation.

36 37 Symposium on Alien Invasive Species and their impact on Natura 2000 amphibian and reptile species in Europe, with special emphasis on the V4 countries IAS and its impacts on the Iberian herpetofauna: Civil efforts to conserve the Caspian whip snake (Dolichophis caspius) a review of conservation projects in the shadow of Budapest

César Ayres Gergely Babocsay 1,2, Balázs Vági 1,3

Asociación Herpetológica Española (AHE), Apartado de correos 191, 28911 Leganés, Madrid, Spain; 1 Amphibian and Reptile Conservation Group, MME BirdLife, Hungary; Költo˝ utca 21, 1121 Budapest, Hungary [email protected] 2 Mátra Museum of the Hungarian Natural History Museum, Kossuth Lajos utca 40, 3200 Gyöngyös, Hungary; [email protected] The Iberian Peninsula has a huge record of human-mediated introduction of non-indigenous 3 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, species (NIS). Since the arrival of the Phoenicians or the Romans, and later the occupation Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary by the Muslims, several species were introduced. Some of them make part now of the fauna The Caspian whip snake (Dolichophis caspius) is close to extinction in Hungary. Very few of the Iberian Peninsula, even as protected species, like the spur-thighed tortoise (Testudo isolated populations have survived, and only sporadic observations of individuals have been graeca). recorded in the last two decades. The populations are under heavy pressure from invasive But in the second half of the 20th century the number of NIS that became invasive alien plants and human activities. In 2011, the Amphibian and Reptile Conservation Group species (IAS) has increased exponentially. Some of the cases are known globally, such as the (ARCG) of MME BirdLife, Hungary launched a conservation programme to assess the status zebra mussel (Dreissena polymorpha), American mink (Neovison vison), North-American sliders of the species and its habitats in the vicinity of Budapest, and to build up a habitat manage- (Trachemys scripta), or the red swamp crayfish (Procambarus clarkii). A huge numbers of fishes ment plan. During 2011 and 2012 with volunteers, we made a total of 32 visits to seven habitats, have been introduced by anglers, like the Danube catfish (Silurus glanis) or the black bass of which one at Pesthidegkút was discovered only in 2012, and observed D. caspius in two (Micropterus salmoides). Recently, the Iberian herpetofauna faces a new threat due to the arrival (including the new one; on the urban Sas-hegy we made no surveys, but D. caspius is regu- of new IAS, like the raccon (Procyon lotor). larly seen there). Dolichophis caspius was observed 11 times at Farkas-hegy and 22 times at An increasing number of conservation projects have aimed to control or eradicate invasive Pesthidegkút, but only 11 (five on Farkas-hegy and six at Pesthidegkút) could be positively species in the Iberian Peninsula. This presentation is a review of the different projects, from large + identified as different. Two different hatchlings were observed on Farkas-hegy. Other historic scale (LIFE Trachemys, LIFE Lampropeltis) to smaller more specific projects (Eradication of habitats have yielded no snakes during our surveys. All suitable habitats are highly isolated, the northern banded newt [Ommatotriton ophryticus]). small in their extent (< 5 ha), and burdened with an increasing number of invasive plants and recreational activities. Most profoundly Ailanthus altissima, Pinus nigra and Populus tremula alter the habitats. We partially removed the black pines from two fragments (altogether ca. 0.5 ha) on Farkas-hegy where snakes were most often observed. Further tree fellings are scheduled to the end of 2013. The effects of the removal of invasive plants on the benefitted population cannot be evaluated at this point, but only a large scale habitat restoration and connection of suitable habitat patches can prevent the short term extinction of D. caspius from the northernmost parts of its European range. Our Dolicophis programme together with other programmes of ARCG triggered an increased citizen involvement.

40 41 Unwanted spring breakers at the Riviera Maya: the red-eared slider Monitoring invasive alien species in the Netherlands, Trachemys scripta elegans (Wied-Neuwied, 1839) invades the cenotes a multi-taxa approach of the Yucatan Peninsula, Mexico Jeroen J. C. W. van Delft 1, Ronald Zollinger 1, J. Wiebe Lammers 2 Stephan Böhm 1 Reptile, Amphibian and Fish Conservation Netherlands (RAVON), P.O. Box 1413, 6501BK Nijmegen, The Netherlands; [email protected] Johannagasse 18/16, 1050 Wien, Austria; [email protected] 2 Food and Consumer Product Safety Authority, Invasive Alien Species Team, P.O. Box 43.006, 3540AA Utrecht, The Netherlands Known to be the turtle species with the most extensive geographic distribution, the red-eared slider (Trachemys scripta elegans) has been recorded from dozens of countries all over the world. Collecting of distribution and population data by volunteers, has a long and successful Because of its longevity, fecundity and opportunistic omnivory it may locally threaten auth- history in the Netherlands. This also concerns (invasive) alien species. Next to data of her- ochthonous turtles and other aquatic wildlife and plants, and is listed among the top 100 petofauna, Reptile, Amphibian and Fish Conservation Netherlands (RAVON) stimulates worst invasive species by the IUCN Invasive Species Specialist Group (ISSG). and facilitates volunteers to collect and pass on data on other taxonomic groups, since the During a field trip to the Yucatan Peninsula from February 13–20, 2012, five cenotes (sink- introduction of online data entry portals. RAVON collaborates with eleven other non- holes filled with freshwater) in the area around Tulum and the Laguna Bacalar (a lake at the governmental organisations and the governmental Invasive Alien Species Team. They set up a border to Belize) were surveyed for turtles. Temperature, pH, carbonic hardness (KH), total multi-taxa (invasive) alien species project, to stimulate the collection of data. The importance hardness (GH) and conductivity were recorded. Turtles were spotted by snorkelling in the of collecting data on (invasive) alien species of all taxonomic groups is stressed by various waterbodies and photo-documented. activities (e. g. the development of an App, identification keys, a popular newsletter, maga- At all surveyed localities, Meso-American sliders (Trachemys venusta) were found, but at zine, websites etc.). In this way 25,000 volunteers are reached. This resulted in an increase two sites red-eared sliders were also present. An adult melanistic male was found in Cenote of records of many taxa, as well as a broader range of taxa being reported by individual Cristal (20°11'24"N, 87°29'56.4"W) and an adult female in Cenote Carwash (20°16'26.4"N, volunteers. 87°29'9.6"W). According to the Global Invasive Species Database, these records are the first An important herpetological result of this project, which can be considered an Early ones for the Yucatan Peninsula. Warning System (EWS), was the detection of the first reproducing population of American Besides these findings, research in hobbyist web forums also showed that Trachemys species bullfrog (Lithobates catesbeianus) in the Netherlands in 20 years. This population was sub- readily hybridize in captivity. Would this also occur in nature, the currently unstable situ- sequently successfully eradicated. Other relevant (invasive) alien herpetofauna species to this ation of Meso-American sliders could be even more complicated than thought before. programme are Triturus carnifex, Rana dalmatina, Elaphe schrenkii and several aquatic turtle species. Another striking result is a strong increase of RAVON volunteers entering data on other exotic taxa such as crayfish. This national project might serve as an example for the development of a European EWS for invasive alien herpetofauna species. A European EWS for invasive alien herpetofauna species, could also pay attention to invasive alien species from other taxonomic groups, affecting herps and their habitats. Therefore, we propose to scale-up this project to a European level.

42 43 Environmental DNA for the detection of alien (and native) species in Challenges in responding to invasive herpetofauna threats in freshwaters the United Kingdom

Gentile Francesco Ficetola 1,2, Tony Dejean 3,4, François Pompanon 1, Pierre Taberlet 1, Jim Foster 1, John Wilkinson 2, Andy Arnell 2, Tony Gent 2 Claude Miaud 4,5 1 Amphibian and Reptile Conservation, The Witley Centre, Witley, Godalming, Surrey GU85QA, United Kingdom; 1 Laboratoire d’Ecologie Alpine, UMR CNRS 5553, Université Grenoble I, 38041 Grenoble Cedex 9, France; [email protected] [email protected] 2 Amphibian and Reptile Conservation, 655A Christchurch Road, Boscombe, Bournemouth, Dorset BH1 4AP, 2 Dipartimento di Scienze dell’Ambiente e del Territorio e di Scienze della Terra, Università degli Studi di Milano- United Kingdom Bicocca, Piazza della Scienza 1, 20126 Milano Italy 3 Laboratoire d’Ecologie Alpine, UMR CNRS 5553, Université de Savoie, 73376 Le Bourget-du-Lac, France At least 10 IAS amphibians and 3 reptiles are thought to be present in breeding populations, 4 SPYGEN, 17 rue du Lac Saint-André, 73375 Le Bourget-du-Lac, France ranging from a single site to areas extending across tens of km2. Reports of further intro- 5 Centre d’Ecologie Fonctionnelle et Evolutive, UMR 5175, Ecole Pratique des Hautes Etudes, Campus CNRS, 1919 route de Mende, 34293 Montpellier Cedex 5, France ductions are common though most do not result in population establishment. There has been excellent progress in some of the main IAS policy areas. For example, there are now standard- Early detection of alien invasive species is extremely important for management, as eradi- ised risk assessments for many IAS. Specific control operations have been very successful, for cation efforts are most likely to be successful if performed when alien species have limited example on North American bullfrog Lithobates catesbeianus. Broad IAS policy has been sub- abundance and range. However, early detection may be challenging, due to the limited den- stantially strengthened in recent years, and legal mechanisms are being improved. There have sity of individuals. Aquatic and semi-aquatic species release DNA into the environment been valuable research projects on, for example, the link between introduced alpine newts through mucus, faeces, urine and remains. It is thus possible using the DNA present in the Ichthyosaura alpestris and the amphibian chytrid fungus. Despite these successes, overall im- environment (eDNA) to detect the presence of alien (and native) species in freshwater. plementation of IAS policy is patchy and poorly co-ordinated for herpetofauna. For instance, Analyses performed on invasive populations of American bullfrogs (Lithobates catesbeianus) despite the acknowledgement of the primacy of prevention, there is minimal action to address showed that primers amplifying short mitochondrial DNA sequences allow to successfully the likely key introduction pathways. IAS detection and surveillance systems are passive, track the presence of frogs in natural wetlands. Using eDNA allows a more accurate detec- largely relying on chance to detect invasions. Research into prevention, impacts, control tion of bullfrogs at very low densities and at any life stage compared to traditional visual and and mitigation is negligible. Links between government and specialists need further develop- audio-surveys. eDNA therefore allows early detection of invasive populations, or to confirm ment, with a more engaged and transparent process for acting on risk assessments. There is the success of eradication projects. Furthermore, multiple studies showed that eDNA allows concern about whether the current allocation of resources is targeted toward the highest pri- a reliable detection of a growing number of species, including several species of both invasive ority actions. Uncertainty about risks too often results in no action, even when rapid response and native amphibians and fish. Furthermore, recent advances of DNA meta-barcoding prom- would involve minimal cost. Underlying these issues, there is a worrying complacency among ise the simultaneous detection of multiple species. Nevertheless, major technical challenges many herpetologists about the risks of IAS. This has contributed to hesitation over action remain, such as contamination and identification of universal markers with high resolution. and has even prompted deliberate introductions. We believe that substantial action to address all these issues is needed soon, especially in light of proposed European Commission IAS legislation.

44 45 Genetic contamination of the indigenous northern crested newt Native or alien frog species in Bratislava (Slovakia)? (Triturus cristatus) by the invasive Italian crested newt (Triturus carnifex) on the Veluwe (The Netherlands) Peter Mikulícˇek

Department of Zoology, Comenius University, Mlynska dolina B-1, 84215 Bratislava, Slovakia; 1,2 2 3 2 Willem R. M. Meilink , Jan W. Arntzen , Jeroen van Delft , Ben Wielstra [email protected]

1 Vrije Universiteit Brussel, Pleinlaan 2, 1050 Brussel, Belgium; [email protected] Non-native species of the western Palearctic water frogs (Pelophylax esculentus complex) were 2 Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands established in many western European countries. They have a potential to readily invade new 3 Stichting RAVON, Postbus 1413, 6501BK Nijmegen, The Netherlands sites, where they compete with indigenous species and predate on the congeneric specimens. Two crested newt species, the Italian crested newt (Triturus carnifex) and the northern crested Hybridization between alien and indigenous individuals leads to introgression and genetic newt (Triturus cristatus), have a parapatric distribution. Hybridization is known to occur pollution of native water frog populations. In this study we investigate genetic structure of between both species in the laboratory, at localities where one of the species has been intro- water frogs in Bratislava (Slovakia) to find out the origin of local populations and dispersal duced in the range of the other, and at the natural contact zone. The Italian crested newt has abilities of water frogs for colonization of new sites. We apply two types of molecular markers been introduced to the Veluwe, the Netherlands, where T. cristatus is a native species. At other with a different mode of inheritance and evolutionary history: sequences of mitochondrial localities where T. carnifex has been introduced, populations appear to be stable and long- ND2 and ND3 genes and nuclear microsatellites. All examined frogs possess either “lessonae” lived. Furthermore, displacement of T. cristatus by introduced T. carnifex has been observed or “ridibundus”-specific mtDNA haplotypes known from central Europe, corroborating their in Switzerland. A study on the distribution of T. carnifex in the Netherlands revealed a recent native origin. While natural ponds in a studied area are inhabited predominately by Pelophylax range expansion. Furthermore, individuals with intermediate phenotypes were found which esculentus and Pelophylax lessonae, man-made gravel pits, established in the course of the last suggests that hybridization between the two species has occurred. Hence, not only direct century, are occupied almost exclusively by Pelophylax ridibundus. This finding highlights a competition, but also genetic pollution by the introduced T. carnifex could pose a threat to colonization potential of P. ridibundus and its ability to occupy new established artificial the native T. cristatus. A detailed knowledge of the distribution of T. carnifex and genetically habitats. Our results also suggest that rivers, river branches and artificial canals facilitate dis- admixed individuals is indispensable for management considerations. However, morphological persal of P. ridibundus to new environment. In contrast, a highly fragmented urban landscape identification in itself is unreliable. Therefore, a detailed map needs to be created on the basis lacking water corridors limits dispersal and gene flow between the ponds. of genetic data. Widespread sampling was performed on the Veluwe and individuals were provisionally classified based on morphological characters. An equal number of individuals were chosen per population for DNA extraction. One mitochondrial DNA marker was used for Sanger sequencing and showed that the distribution of T. carnifex haplotypes is more widespread than expected based on morphological data. An additional 52 markers were amplified by multiplex PCR and sequenced by the Ion PGM Next Generation Sequencing platform. The nuclear DNA data were analysed using New Hybrids to determine the extent of introgres- sion of T. carnifex alleles into T. cristatus populations on the Veluwe.

46 47 Plastic invaders: reactions of tadpoles of the invasive anuran Impact of invasive species on the herpetofauna of the Czech Republic Discoglossus pictus in front of native and introduced predators Roman Rozínek 1, David Fischer 2, Vojteˇch Balázˇ 3 Eudald Pujol-Buxó 1, Olatz San Sebastián 1,2, Núria Garriga 1, 1 Caroline Pereira Almeida 3, Gustavo A. Llorente 1 NaturaServis Ltd., Ricˇarˇova 66, 50301 Hradec Králové, Czech Republic; [email protected] 2 Mining Museum Prˇíbram, Hynka Klicˇky 293, 26101 Prˇíbram 6, Czech Republic 3 Department of Biology and Diseases of Wildlife, Faculty of Veterinary Hygiene and Ecology, University of Veterinary 1 Departamento de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal 643, and Pharmaceutical Sciences Brno, Palackého 1/3, 61242 Brno, Czech Republic 08028 Barcelona, Spain; [email protected] 2 Departamento de Herpetología, Aranzadi Zientzia Elkartea-Sociedad de Ciencias Aranzadi, Zorroagagaina 11, 20014 Donostia-San Sebastián, Spain Every herpetologist is aware of the danger that alien species pose to amphibian and reptilian 3 Universidade Estadual de Santa Cruz, Departamento de Ciências Biológicas, Rodovia Ilhéus–Itabuna Km 16, populations. However most of the support for deleterious effects of non-native and invasive 45662-000 Salobrinho, Ilheus, BA, Brazil organisms is based on anecdotal reports. Scientifically processed and analysed data about the non-native species impact on reptilian and amphibian fauna is completely absent from the A growing number of studies consider phenotypic plasticity one of the major forces driving area of Czech Republic. Only individual observation data are available at the moment e. g. a survival and rapid adaptation of species in front of new threats like invasive predators, thus dissection of invasive fish, brown bullhead (Ameiurus nebulosus), that showed the digestive being biological invasions an optimal system to study the evolution of anti-predator pheno- tract to be completly filled by ingested smooth newts (Lissotriton vulgaris). Most of the “evi- typic plasticity. In order to deeper understand the role of inducible responses in the popula- dence” is in form of mere observations about some surveyed species going lost from locality tion dynamics related with current biological invasions in our area, we first studied the plastic after alien species was detected there. An example of such case is the absence of dice snake responses of both native (Pelodytes punctatus) and invasive (Discoglossus pictus) anuran tad- (Natrix tessellata) in the watershed of the Berounka River after invasion of mink (Mustela poles, all facing native (dragonfly Anax sp.) and two invasive (fish Gambusia holbrooki and vison). The most important introduced species with direct effect on amphibians and reptiles crayfish Procambarus clarkii) predators. Native P. punctatus displayed the predictable set of (and other fauna as well) include the fish species Ameiurus melas (black bullhead), A. nebu- reactions against native dragonflies, but milder (morphological or behavioral) reactions to in- losus (brown bullhead), Carassius gibelio (Prussian carp), Pseudorasbora parva (topmouth gud- vasive predators. Invasive D. pictus reacted behaviorally to all predators, unexpectedly reacting geon) and of mammals Mustela vison (mink), Procyon lotor (raccoon) and Nyctereutes pro- also morphologically to native Anax sp. Our results support prey-predator specificity in these cyonoides (raccoon dog). The area of Czech Republic harbours at least two species of alien reactions and dissociation between behavioral and morphological plasticity in tadpoles, al- (American) crayfish species that are known to propagate the crayfish plague, which extirpates though some reactions in D. pictus could suggest unspecific plastic reactions against predation native crayfish species. Additionally these invasive crayfish are potential vectors of the am- in this species. To have a deeper understanding of the anti-predator plastic reactions of this phibian pathogen causing chytridiomycosis. The habitat changes triggered by expanding invasive species, in a second experience we confronted this invasive anuran with the most non-native plant species cause serious problems to both amphibians and reptiles. Examples usual predators in our study area: common backswimmers (Notonecta sp.) and common of such plants include Canadian waterweed Elodea canadensis with ability to overgrow the darters (Sympetrum striolatum). Results of this second experience further support the ability of ponds, thus making them unsuitable for survival of amphibians and their larvae. Terrestric D. pictus to react specifically to native predators and reinforce conclusions of the first work, plants especially Helianthus tuberosus and Impatiens glandulifera cause intense shading of river pointing to an important role of phenotypic plasticity in this system. On the other side, banks, thus limiting the options for basking for dice snake (Natrix tessellata) and lowering the each species suffered from a particular set of trade-offs between inducible responses and their temperature of egg-laying sites. costs, and interestingly we usually detected milder and unexpected patterns in combinations We have to admit that our knowledge of the invasive species impact on our amphibians and using introduced species. reptiles is still limited, although first studies are on the way we will have to wait until the results are available.

48 49 Tracking the expansion of an invasive species Xenopus laevis using Invasion patterns of ancient and recent alien herpetofauna in environmental DNA techniques the Balearic Islands

Jean Secondi 1, Claude Miaud 2, Guillaume Koch 3, Benjamin Audebaud 3, Iolanda Raquel Silva-Rocha, Daniele Salvi, Miguel A. Carretero Nicolas Cotrel 4, Sylvie Desgranges 5, Tony Dejean 6 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto Campus Agrário de 1 LUNAM, Université d’Angers, GECCO, Angers, France; [email protected] Vairão, 4485-661 Vairão, Portugal; [email protected] 2 UMR CNRS CEFE, Montpellier, France 3 Communauté de Communes de l’Argentonnais, Argenton les Vallées, France Species introductions are becoming a major threat to biodiversity and economy but its effects 4 Deux-Sèvres Nature Environnement, Niort, France are unevenly distributed in time and space. The Mediterranean Basin is the biodiversity 5 LPO Anjou, Saumur, France hotspot where introductions shape the vertebrate communities in greater extent, being the 6 SPYGEN, Le Bourget-du-Lac, France insular species the most affected. Due to their life history, amphibians and reptiles are among The African clawed frog Xenopus laevis has been introduced in several parts of the world the groups most widely introduced but also most threatened by species introduction. where it is considered as an invasive taxon now threatening local herpetofaunas. It has been Mediterranean islands harbour a combination of herpetofaunal elements: endemics long time released about 30 years ago in western France where populations survived in the wild and rap- evolving under insular conditions; natural colonisers from the adjacent continents in recent idly expanded. The introduction point lies on the Loire River basin which covers 20% of the geological times; and ancient or recent invaders transported, voluntarily or not, by humans. French territory and includes two wetlands of international value (Brenne, Sologne). Our Distinguishing between them, as determining the sources and invasion pathways, is not only ability to track and predict the movement of individuals is a key issue for the control of inva- of biogeographical interest but also crucial for conservation management. Here we analyse sive species. To this regard, tracking the colonization front is crucial to better apprehend the the paradigmatic case of the Balearic herpetofauna by using a combination of literature and dispersal behaviour of X. laevis in the local context. However, the task becomes more and original molecular data. The Balearic Islands currently harbour much more alien than native more challenging as the size of the colonized area increases. Classical trapping methods may amphibians (4/1) and reptiles (21/2). Although archaeology dates some introductions back to fail to ascertain species presence on the outer margins because of unbearable logistic con- the Neolithic, the Balearic Archipelago has increasingly importing alien species until present. straints. For these reasons, alternative methods are required to improve detectability and cov- Source regions and invasion routes have shifted throughout time, from the Eastern Medi- erage of expanding invasive species. Environmental DNA (eDNA) techniques may provide terranean and North Africa to the Iberian Peninsula and extra-European areas and from pas- suitable solutions. We provide an update of the colonized area by X. laevis in western France sive cargo transport or food/religion related to pet shops, nursery trade and tourism. Overall, and outline the conservation issues related to that invasion. We then present results from one native amphibian (Alytes muletensis) and two native reptiles (Podarcis lilfordi and P. detection tests using several eDNA methods in sites of low and high density, and compare pityusensis) are restricted to Ibiza and the less disturbed Formentera and small islets. Even the pros and cons of classical capture and eDNA methods for X. laevis. there, menaces have increased during the last decade, particularly due to predatory snakes and competitive lacertids. While the Balearic herpetofauna forecasts what could be an omi- nous fate, it also provides an opportunity to extract lessons for developing effective conser- vation strategies.

50 51 Alien herpetofauna in Poland—problems and solutions Populations and habitats of the Hungarian meadow viper (Vipera ursinii rakosiensis) in the Hanság area Wojciech Solarz, Katarzyna Kurek Gábor Takács 1, Róbert Dankovics 2, Attila Pellinger 1 Institute of Nature Conservation, Polish Academy of Sciences, Al. Mickiewicza 33, 31-120 Kraków, Poland; [email protected] 1 Ferto˝-Hanság National Park Directorate, Rév-Kócsagvár, 9435 Sarród, Hungary; [email protected] 2 Savaria Museum, Kisfaludy S. u. 9, 9700 Szombathely, Hungary Although there are no established alien amphibians or reptiles in Poland to date, the prob- lem of alien herpetofauna should not be ignored. Availability of exotic pets and number of The Hungarian meadow viper is one of the most endangered snakes within the European amateur keepers is increasing. However, the level of awareness about causes and consequences fauna. The West-Hungarian population of the subspecies survived in the Hanság area on two of biological invasions remains low. As a result, the number of releases of unwanted pets, pitches with dry grasslands (loess grasslands, secondary grasslands) void of floods and inland including herptiles, is alarming. In addition, there were cases of illegal introductions of rep- inundations that make up less than 25 ha altogether and are surrounded by lower lying marsh tiles for “fauna improvement” by professional herpetologists and dedicated nature lovers. At meadows, willow marshes and marsh forests. least 10 reptile species have been recorded in the wild. There are only anecdotal records for Within the frame of the “Conservation of Hungarian meadow viper (Vipera ursinii most of them but the number of confirmed records for red-eared slider (Trachemys scripta ele- rakosiensis) in the Carpathian-basin” LIFE+ project intervention is done on 392 ha within the gans) and Cumberland slider (T. s. troostii) reached a few hundreds. Single cases of egg-laying 1609 ha “Viper Reserve”. The aim of project is the suppression of invasive species, conserva- of red-eared sliders were detected but hatching was not confirmed so far. Even though the tion of and help to spread grassland communities favourable for the Hungarian meadow species may not breed successfully in Poland, its life span combined with the number of viper. In the place of higher situated hybrid poplar plantations and ploughlands 32 ha of new released individuals constitute serious threat for native biodiversity, particularly for European dry grassland are created that doubles the extension of wintering grounds. The surrounding pond turtle (Emys orbicularis). former marsh meadows and swamp meadows are being restored on 360 ha primarily due to One of the major attempts to mitigate the invasive alien species problem in Poland is a new the removal of Salix cinerea and Solidago gigantea. Further conservation management of the decree of the Minister of Environment that came into force in 2012. It imposes restrictions habitats is done by mowing and grazing of sheep, and it is protected by an electric fence to on 52 invasive alien species of plants and animals. Import, keeping, breeding and selling of eliminate damages by wild boar. these species requires a permission from the General Director for Environmental Protection. Within the frame of botanical monitoring the spread and general state of habitats was Breaking the law is subject to a fine or jail. Negligence leading to escape of animals listed in examined with vegetation mapping. We try to show the changes in the state of the single the decree may result in similar consequences. Restricted species include American bullfrog habitat types due to management with coenological snapshots taken on constant sampling (Lithobathes catesbeianus): its import and breeding is forbidden altogether, including by spots. zoos. However, to reduce the risk of release of unwanted individuals, it is possible to obtain The monitoring survey of the Hungarian meadow viper and its potential food animals— a permit for keeping and selling bullfrogs that are already possessed, on the condition that Orthoptera, Amphibia, Reptilia—is done since 2001. Changes in the number of species and they are microchipped the risk of escape is kept to a minimum. Restricted reptile species are individuals of the assigned taxa as well as the relative frequency of the viper sightings in a unit snapping turtle (Chelydra serpentina), painted turtle (Chrysemys picta), false map turtle (Grap- time were tracked on the two habitats with nearly the same times of sampling. The presence temys pseudogeographica) and all subspecies of Trachemys scripta. Import and breeding of these of predator species and the effect of habitat management were also subject of the survey. species will only be allowed for zoos, while individual owners and pet shops can only obtain permits to keep and sell turtles that they currently possess, provided that they are micro- chipped and the risk of escape is minimized. The major challenge to make the new law effective is awareness raising among pet keepers. A few campaigns were organised and new are planned in close future.

52 53 Amphibian chytridiomycosis Symposium Summary of Batrachochytrium dendrobatidis research in the Czech Environmental predation of Batrachochytrium dendrobatidis Republic in wild and captive amphibians during the last five years determines infection dynamics

1 2 2 3 4 Vojteˇch Balázˇ , Petr Civisˇ , Jirˇí Vojar , Martin Sandera , Jan Mateˇju˚ , M. Blooi 1, D. S. Schmeller 2, A. Martel 1, T.W.J. Garner 3, M. C. Fisher 4, Barbora Havlíková 2, Roman Rozínek 5, Trenton Garner 6 F. Clare 3,4, A. Loyau 2, F. Pasmans 1

1 Department of Biology and Diseases of Wildlife, Faculty of Veterinary Hygiene and Ecology, University of Veterinary 1 Department of Pathology, Bacteriology, and Avian Diseases, Faculty of Veterinary Medicine, Ghent University, and Pharmaceutical Sciences Brno, Palackého trˇ. 1/3, 61242 Brno, Czech Republic; [email protected] Salisburylaan 133, 9820 Merelbeke, Belgium; [email protected] 2 Department of Ecology, Faculty of Environmental Sciences, Czech University of Life Sciences Prague, Kamy´cká 129, 2 UFZ—Helmholtz-Centre for Environmental Research, Department of Conservation Biology, Permoserstraße 15, 16521 Prague 6, Czech Republic 04318 Leipzig, Germany 3 Museum of Nature Bohemian Paradise, Prachov 37, 50601 Jicˇín, Czech Republic 3 Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, United Kingdom 4 Karlovy Vary Museum, Nová louka 23, 36001 Karlovy Vary, Czech Republic 4 Department of Infectious Disease Epidemiology, Imperial College London, London W2 1PG, United Kingdom 5 NaturaServis Ltd., Rícarova 66, 50301 Hradec Králové, Czech Republic 6 Institute of Zoology, Zoological Society of London, Regent’s Park, London, NW1 4RY, United Kingdom The occurrence pattern of Batrachochytrium dendrobatidis (Bd) is not homogeneous at local and regional scales. Sites which seem appropriate and in vicinity to long-term infected sites, The Czech Republic is inhabited by 13 anuran and 8 caudate species, thus having the richest are not always colonized by Bd. We hypothesized that the presence of resident micro- batrachofauna in Central and Eastern Europe. All native species are protected by law and organisms that prey on Bd contributes to explain the observed pattern of Bd colonization. many receive conservation attention. The exotic amphibians are common in both public and The aim of this study was to better understand the interactions of Bd with resident micro- zoo collections. This context gives many opportunities, yet challenging environment for the biotic communities, with a focus on the impact of microorganisms on Bd viability, infection Batrachochytrium dendrobatidis research. probability and infection intensity in amphibian hosts. The first effort to study B. dendrobatidis in the area was triggered by cooperation with Sites in the Pyrenean mountain range with known history of absence or presence of Bd the RACE (Risk Assessment of Chytridiomycosis to European Amphibian Biodiversity) in were sampled. The decrease in Bd zoospore viability after exposure to the water samples was 2008. At the moment two Czech universities work in close collaboration on the topic and assessed using a method combining ethidium monoazide with real-time PCR3. This method several nature conservation and research bodies are involved in field sampling. was also used to assess the effect of cultures of several freshwater microorganisms on Bd Altogether 1562 samples from wild amphibians were collected between 2008 and 2012, viability. Based on the results of these first experiments several microorganisms were selected allowing analyses of host specificity and bioclimatic modelling. The distribution of the for Bd zoospore ingestion experiments and for a Bd infection experiment with Discoglossus pathogen in the Czech Republic appears to be almost general, but lowlands and warmer areas scovazzi tadpoles in the absence and presence of microorganisms. seem to be more suitable. The most commonly infected amphibians are from the genera We were able to show a significant correlation between the abundance of microorganisms Pelophylax and Bombina with subadult post-metamorphic individuals being in the greatest and Bd zoospore persistence in the Pyrenean water samples. In pure culture, some but not all risk of acquiring the fungus. Individual cases of mortalities linked with high infection burdens microorganisms were highly efficient in killing Bd. These highly effective species ingested Bd detected by qPCR were observed in Bombina variegata and Bufo viridis, but several species zoospores and prevented Bd colonization of tadpoles. In conclusion, environmental micro- showed the ability to lose the infection (Bufo calamita, B. viridis, Bombina variegata). organisms dictate chytrid infection dynamics. The preliminary results from exotic amphibians showed that over 100 species are found in Czech collections. Dataset of aproximately 500 samples proved that cases of infection by B. dendrobatidis and fatal chytridiomycosis occur in small private collections as well as in zoos. Obtaining data from “pet trade” proved to be a challenging task, bringing very little data. The Czech Republic is a great area for chytridiomycosis research and it allows unique com- parisons between the present taxa, analysis of bio-geographic, bioclimatic and social context.

56 57 Dramatic decline in the fire salamander (Salamandra salamandra) in A novel pathogen from fire salamanders (Salamandra salamandra) the Netherlands—a species on the edge of extinction An Martel 1, Annemarieke Spitzen-van der Sluijs 2, Mark Blooi 1, Wim Bert 3, Richard Ducatelle 1, Matthew C. Fisher 4, Antonius Woeltjes 2, Wilbert Bosman 2, Wilbert Bosman 1, Annemarieke Spitzen-van der Sluijs 1, Frank Pasmans 2, An Martel 2 Koen Chiers 1, Franky Bossuyt 5, Frank Pasmans 1 1 Reptile, Amphibian & Fish Conservation Netherlands (RAVON), P.O. Box 1413, 6501BK Nijmegen, The Netherlands; [email protected] 1 Department of Pathology, Bacteriology and Avian Diseases, Faculty of Veterinary Medicine, Ghent University, 2 Department of Pathology, Bacteriology and Avian Diseases, Faculty of Veterinary Medicine, Ghent University, Salisburylaan 133, 9820 Merelbeke, Belgium; [email protected] Salisburylaan 133, 9820 Merelbeke, Belgium 2 Reptile, Amphibian and Fish Conservation the Netherlands, P.O. Box 1413, 6501BK Nijmegen, The Netherlands 3 Department of Biology, Netamology Unit, Faculty of Science, Ghent University, Ledeganckstraat 35, 9000 Ghent, In the Netherlands, the fire salamander (Salamandra salamandra) occurs at the edge of its dis- Belgium 4 MRC Centre for Outbreak Analysis and Modelling, Department of Infectious Disease Epidemiology, tribution range and is restricted to three small populations in the extreme south of the country. Imperial College London. St Mary’s Campus, Norfolk Place, London W2 1PG, United Kingdom The largest of these populations, the Bunderbos population, has been very well monitored 5 Amphibian Evolution Lab, Biology Department, Vrije Universiteit Brussel, Pleinlaan 2, 1050 Brussel, Belgium since the seventies. For many years, the population was stable until the population completely collapsed in 2010. In 2011 four specimens were seen, in 2012 only two. A trend analysis In the Netherlands, the fire salamander (Salamandra salamandra) is restricted to three small based on the populations form 1997–2011 showed a highly significant drop of 96%. We aimed populations in the extreme south of the country. Since 2010 the species has shown a rapid at determining the reason for the decline, by examining the population’s genetic diversity, and significant decrease of 96% of the total population. Here we isolated and characterized presence of known amphibian pathogens, and environmental concentrations of pesticides the novel agent that drove this Salamandra population to extirpation. Molecular and pheno- and nitrate concentrations. We failed to attribute the observed extinction event to any known typic characterization, including infection experiments, show this agent to be highly patho- cause of amphibian decline, rendering the collapse of the fire salamander populations in the genic to fire salamanders. The emergence of a novel lethal pathogen in European amphibian Netherlands enigmatic. communities is worrying. We developed a diagnostic PCR that allows inclusion of the detec- tion of this novel agent in any amphibian health assessment protocol.

58 59 Chytrid fungus in Western Europe European plethodontid salamanders (genus Speleomantes): resistant to chytridiomycosis? Claude Miaud Frank Pasmans 1, Pascale van Rooij 1, Mark Blooi 1, Giulia Tessa 2,3, Sergé Bogaerts 4, Biogéographie et Ecologie des Vertébrés (E. P.H. E.), Centre d’Ecologie Fonctionnelle & Evolutive, UMR 5175, Giuseppe Sotgiu 3, Trenton W.J. Garner 5, Matthew C. Fisher 6, Tonnie Woeltjes 7, Campus CNRS, Montpellier, France; [email protected] Wouter Beukema 8, Stefano Bovero 3, Connie Adriaensen 1, Fabrizio Oneto 9, 9 1 9 It has been over three decades since scientists first studied the massive declines of amphibian Dario Ottonello , An Martel , Sebastiano Salvidio populations observed across the globe. Among several causes, Batrachochytrium dendrobatidis 1 (Bd) was identified as the causative agent in many cases, and the World Organization for Department of Pathology, Bacteriology and Avian Diseases, Faculty of Veterinary Medicine, Ghent University, Salisburylaan 133, 9820 Merelbeke, Belgium; [email protected] Animal Health (OIE) now includes Bd as a notifiable disease. The aim of this presentation is 2 Dipartimento di Scienze della Vita e Biologia dei Sistemi, Università degli Studi di Torino, to focus on the ongoing studies mostly in Western Europe e. g. the observed Bd occurrence Via Accademia Albertina 13, 10123 Torino, Italy and prevalence in amphibian populations and species, the identification of lineages and fac- 3 Zirichiltaggi—Sardinia Wildlife Conservation, s.v. Filigheddu 62/C, 07100 Sassari, Italy tors contributing to the spread of virulent lineage, their impact on native amphibians, and 4 Lupinelaan 25, 5582CG Waalre, The Netherlands 5 Institute of Zoology, Zoological Society of London, Regent’s Park, London NW1 4RY, United Kingdom effort to mitigate the threat of chytridyomicosis. Despite many gaps in the understanding of 6 Department of Infectious Disease Epidemiology, St Mary’s Hospital, Imperial College, London W2 1PG, the disease, we are beginning to unravel important elements of Bd biology and make progress United Kingdom toward amphibian conservation. 7 Stichting RAVON, Natuurplaza, Gebouw Mercator 3, Toernooiveld 1, 6525ED Nijmegen, The Netherlands 8 Cátedra Rui Nabeiro, Universidade de Évora, Casa Cordovil, Rua Dr. Joaquim Henrique da Fonseca, 7000-890 Évora, Portugal 9 DISTAV—Università di Genova, Corso Europa 26, 16132 Genova, Italia

Plethodontid salamanders are represented in Europe by 8 species of the genus Speleomantes, which are confined to small geographic ranges. Despite the established presence of Batracho- chytrium dendrobatidis (Bd) in their geographic range, mortality attributed to chytridio- mycosis has not been reported for this genus. The aim of this study was to determine the prevalence of Bd in Speleomantes populations across their distribution range, and to determine the susceptibility of Speleomantes salamanders to chytridiomycosis. We show that Bd was un- detectable in 921 skin swabs of all eight species of European Speleomantes. After inoculation of Speleomantes strinatii with a highly virulent isolate of Bd (BdGPL), none of five exposed S. strinatii were colonized by Bd beyond 2 weeks post inoculation. For comparison, experi- mentally inoculated Alytes muletensis developed increasing Bd-loads over a 4-week period. A partial explanation for the observed resistance to chytridiomycosis was provided by the antifungal activity of the skin secretions of seven Speleomantes species, showing pronounced killing activity against Bd over 24 hours. In conclusion, the absence of Bd in Speleomantes combined with resistance to experimental chytridiomycosis and highly efficient skin defenses indicate that the genus Speleomantes is a taxon unlikely to decline due to Bd.

60 61 Exposure vs. infection: can Batrachochytrium dendrobatidis Monitoring ranavirus-associated mortality in a Dutch heathland in pathogenicity be an issue to salamanders? the aftermath of an outbreak

Gonçalo M. Rosa 1,2, Richard Griffiths 1, Rui Rebelo 2, Ana Duarte 3, Luis Tavares 3, A. M. Spitzen-Van der Sluijs 1,3, M. Kik 2, A. Grone 2, J. van den Broek 2, F. Pasmans 3, Jaime Bosch 4 A. Martel 3, J. M. Rijks 2

1 Durrell Institute of Conservation and Ecology, School of Anthropology and Conservation, University of Kent, 1 Reptile, Amphibian & Fish Conservation Netherlands (RAVON), P.O. Box 1413, 6501BK Nijmegen, Canterbury, Kent, CT2 7NR, United Kingdom The Netherlands; [email protected] 2 Centro de Biologia Ambiental, Departamento de Biologia Animal, Faculdade de Ciências da Universidade de 2 Dutch Wildlife Health Centre, Yalelaan 1, 3584CL Utrecht, The Netherlands Lisboa, Bloco C2, Campo Grande, 1749-016 Lisboa, Portugal; [email protected] 3 Department of Pathology, Bacteriology and Avian Diseases, Faculty of Veterinary Medicine, Ghent University, 3 Centro de Investigação Interdisciplinar em Sanidade Animal, Faculdade de Medicina Veterinária, Salisburylaan 133, 9820 Merelbeke, Belgium Universidade Técnica de Lisboa, Lisboa, Portugal 4 Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain When ranavirus epizootics lead to mass mortalities of adult life stages, this is relatively easy to spot in the wild. However, in the aftermath of such an outbreak, mortality associated with In recent years, pathogenic fungus Batrachochytrium dendrobatidis (Bd) has becoming widely ranavirus infection may be more difficult to detect. We provide an overview on the spatial recognized as posing major threats to amphibians. However, while some species seem to have distribution and temporal dynamics of ranavirus infections in a Dutch National Park in the disappeared completely following Bd outbreak, other species (and even populations) persist year following a mass mortality event caused by ranavirus infection in 2010. without being (apparently) affected. Although correlation of Bd with loss of anuran popu- We monitored five ponds intensively between May–October 2011. The visits to the site lations has been well documented, the pathogenicity of the fungus to urodeles is still poorly were biweekly in May, June and October, and weekly in the other months. We noted the understood particularly in Europe. number of encountered amphibians and their condition, and we additionally monitored the Recently has been documented a massive decline in common midwife toads (Alytes obstet- chemical water quality. Ranavirus was detected in 4/5 ponds, causing mortality in water frogs ricans) in Serra da Estrela, Portugal, where the species have disappeared from several ponds. (Pelophylax spp.) and great crested newts (Triturus cristatus). Of all sighted amphibians Both A. obstetricans and fire salamander (Salamandra salamandra) populations have been 2% showed ranavirus-related disease symptoms, and additionally a ten-spined stickleback monitored since 2010 for presence, counts and assessment of Bd prevalence. Our study aimed (Pungitius pungitius) tested positive for CMTV, but it is unclear if this animal died due to to understand the susceptibility of S. salamandra to amphibian chytrid. Experimentally we this infection. tested the affects of Bd on growth and developmental rates in larvae, exposing individuals Amphibian mass mortalities were not encountered, but throughout the season dead indi- to different doses of Bd zoospores (low, medium, high and two controls: sham exposure and viduals were observed. These continuing mortalities are unrelated to water quality and water extremely high dose). temperature. Our field surveys showed no significant fluctuation in the numbers of larvae found over the four years, contrasting with the abundances of Alytes tadpoles that dropped dramatically. Both species share the same chytrid contaminated water habitats, however, no mortality or Bd positive salamanders have been recorded in any of the monitored sites. Additionally, lab experiments provided robust data to support that S. salamandra larvae are very resistant to the disease chytridiomycosis. Exposing larvae to a highly virulent Bd strain and in a manner that resulted in potentially lethal infection resulted in infection of only a reduced number of salamanders in the positive control treatment. After four weeks of exposures, no infection was detected in any of the other treatments. Sublethal costs exist in exposed individuals, indicating that successful immune responses to Bd exposures come at a considerable cost. Costs are accrued in a dose dependent manner during the larval stage and can be also ex- pressed accordingly.

62 63 Herpetofauna and transport systems —towards a better understanding and impact reduction Gully pots: death-traps for amphibians Low usage rates of an under-road tunnel by three Australian anurans in an experimental setting Annemarie van Diepenbeek, Raymond Creemers Andrew J. Hamer 1, Rodney van der Ree 1, Michael J. Mahony 2, Tom Langton 3 RAVON (Reptile, Amphibian & Fish Conservation Netherlands), P.O. Box 1413, 6501BK Nijmegen, The Netherlands; [email protected] 1 Australian Research Centre for Urban Ecology, Royal Botanic Gardens Melbourne c/o School of Botany, University of Melbourne, Parkville, Victoria 3010, Australia; [email protected] Gully pots are essential for ridding the roads of rainwater but can be a death-trap for amphib- 2 School of Environmental and Life Sciences, The University of Newcastle, Callaghan, New South Wales 2308, ians. Each year, large numbers of amphibians, both adults and juveniles, fall into gully pots Australia and die through starvation or by being washed away into the sewer system by surges of rain- 3 Herpetofauna Consultants International, Triton House, Bramfield, Halesworth, Suffolk IP19 9AE, United Kingdom water. This happens year around, but especially during spring migration. The vastness of the problem is illustrated by the numbers of amphibians retrieved from Many amphibians in Europe and North America regularly use under-road tunnels during gully pots in the Netherlands in 2012. In 36 locations, in three random counts we collected dispersal. However, these structures have not been shown to mitigate the impacts of road a total of 782 vertebrates from 526 gully pots of which 683 were amphibians. A rough calcu- mortality on frog populations in Australia. We tested the behavioural response of three lation of the numbers that fall victim to the drainage system came up to an estimate of Australian frog species to a 12-metre amphibian under-road tunnel (ACO Polycrete Pty. between several hundred thousand to more than half a million adult amphibians, and many Ltd.). We used infra-red camera units at night to view frog behaviour upon release from an times this number of immature ones, each year alone in the Netherlands (the Netherlands acclimation chamber near the tunnel entrance. In April and October 2012 we tested 34 striped has 7 million gully pots). marsh frogs (Limnodynastes peronii), 54 green and golden bell frogs (Litoria aurea) and 15 A test with live amphibians showed that amphibians will find and use climb-out construc- broad-palmed rocket frogs (Litoria latopalmata). During an experimental run, one individual tions. Our innovative research suggests to implement these constructions at European level was placed under the acclimation chamber for 5 minutes, then released and its behaviour was to prevent this additional amphibian mortality. monitored. A “balk” was recorded and the run was terminated after 25 minutes if (1) an indi- vidual had not entered the tunnel, (2) an individual entered the tunnel but remained inside, or (3) an individual entered the tunnel but turned back. The overall proportion of tunnel usage (i. e. frogs entering the tunnel) among the three species was 0.13 out of 103 frogs tested. The proportion of usage for Lim. peronii, Lit. aurea and Lit. latopalmata was 0.21, 0.07 and 0.13, respectively. The mean time taken for an individual to enter the tunnel was 14 minutes 24 seconds. The results of this study showed that there was low usage of the ACO under-road tunnel compared to the relatively high usage rates of the tunnels by amphibians in the Northern Hemisphere. However, Australian frogs exhibit no seasonal movements in migra- tory behaviour. Therefore, the lack of use of the tunnels probably reflects the behaviour of Australian frogs and our experimental setting, rather than the apparent unsuitability of the ACO tunnels. The next step in this study is to test frog usage of ACO tunnels installed under several roads near wetlands so that the behaviour of wild individuals can be assessed.

66 67 Prioritizing amphibian road mitigation measures at regional scale: Road mitigation using tunnel and fence systems for a protected predicting Bufo bufo mortality hotspots using habitat-resistence models species, the great crested newt in the UK. Is it working?

Cátia Matos 1, Silviu Petrovan 2,3 Cátia Matos 1, Silviu Petrovan 2,3

1 Centro de Investigação em Ciências Geo-Espaciais (CICGE), Universidade do Porto, Faculdade de Ciências, 1 Centro de Investigação em Ciências Geo-Espaciais (CICGE), Universidade do Porto, Faculdade de Ciências, Observatório Astronómico Prof. Manuel de Barros Alameda do Monte da Virgem, 4430-146 Porto, Portugal; Observatório Astronómico Prof. Manuel de Barros Alameda do Monte da Virgem, 4430-146 Porto, Portugal; [email protected] [email protected] 2 Froglife, 2A Flag Business Exchange, Vicarage Farm Road, Fengate, Peterborough PE1 5TX, United Kingdom 2 Froglife, 2A Flag Business Exchange, Vicarage Farm Road, Fengate, Peterborough PE1 5TX, United Kingdom 3 CEMS, University of Hull, Scarborough Campus, Filey Road, Scarborough, North Yorkshire YO11 3AZ, 3 CEMS, University of Hull, Scarborough Campus, Filey Road, Scarborough, North Yorkshire YO11 3AZ, United Kingdom United Kingdom

Among all effects produced by roads, mortality by collision with vehicles can be the most Froglife manages Hampton Nature Reserve, probably the largest population of great crested important. Amphibians are the most affected group and road fatalities have a significant newt (Triturus cristatus) in Europe with an estimated 35,000 adult individuals in around 400 impact on population dynamics and viability, especially in protected areas. Considered the ponds. Since 2007 we have conducted intensive monitoring of a tunnel and fence system most threatened by roads in Europe because of their movement patterns and need for a com- connecting two parts of the reserve which have been separated by a new road. Both areas con- plex matrix of habitats, Bufo bufo is a target species for mitigation measures. However, due tained terrestrial and aquatic habitats which makes this system different than most other to the extensive road network, precise locations (hotspots) identification is needed to apply amphibian road mitigation sites. The system comprises two large ARCO tunnels (2.5 m high these measures successfully. The aim of the study was to develop and test habitat-resistance x 5.5–6 m wide) as well as a 50 x 50 cm ACO tunnel. Here we present 5 year annual moni- models using GIS for identifying possible B. bufo mortality hotspots on roads located at 12 toring results demonstrating that while the system successfully connects the newt metapop- different Nature Improvement Areas (NIA) in UK. Also, it aims to inform the different ulations separated by the road there is very important variability in annual tunnel crossings, stakeholders (local planning authorities, regional nature partnerships, NIA staff, conservation probably influenced by weather conditions. Between 10 and 113 newts were annually trapped bodies and toad patrollers) about the potential critical road sites for this priority species in in the tunnels in different years with newts successfully crossing from both sides of the road different regions. Twelve habitat suitability modes and least-cost models were built using but with substantial sex bias. We suggest that such monitoring studies need to be carried out expert-literature knowledge on B. bufo road mortality. Models were validated with B. bufo over a long period of time (5–10 years) and should include data from before the tunnel sys- presence data collected from different sources. Model’s fit was good, predicting 80% of tem has been emplaced in order to accurately verify their success. species occurrence, despite information on ponds location being not used uniformly across the NIA network. Models predicted high-quality habitat and possible corridors for B. bufo presence in all NIA’s. From the network considered for analysis, 3% of 500 m cells showed 90% of possibility for being B. bufo road-kill hotspots. All NIA cells presented sections with 45% probability for being a hotspot, representing nearly 20% of the road network. TOR sites buffers match 30% of the possible hotspots cells. These models were designed to provide regional-scale data information on potential road mortality hotspots to different management parts for future study and implementation of possible mitigation measures.

68 69 Amphibian migration characteristics and tunnel use during spring Temporary and permanent mitigation measures on Hungarian roads: migration in 2009–2012 along the Hont–Parassapuszta section of the failures and successes No. 2. road, Hungary Miklós Puky Tímea Mechura 1,2, Dorottya Gémes 1,2, Béla Mester 2,3, Miklós Puky 2,4 MTA Centre For Ecological Research, Danube Research Institute, Jávorka S. utca 14, 2131 Göd, Hungary; [email protected] 1 Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 2 Varangy Akciócsoport Egyesület, IX. utca 40, 1172 Budapest, Hungary Fragmentation is a main conservation problem, especially in Europe, where providing access 3 University of Debrecen, Faculty of Science and Technology, Department of Ecology, Conservation Ecology Research Group, Egyetem tér 1, 4032 Debrecen, Hungary to habitats of different quality for wildlife is of increasing importance. A typical example of 4 MTA Centre for Ecological Research, Danube Research Institute, Jávorka S. utca 14, 2131 Göd, Hungary this process is the creation and maintenance of temporary and permanent mitigation meas- ures to help herpetofauna species under or over roads. Legally, it is dealt with in many Amphibian mitigation measures have been built on roads worldwide since 1969. In 2006 European Union countries, the practice, however is not so successful. Here an overview is a tunnel-barrier system was constructed between Hont and Parassapuszta, which is the site given on Hungary with a focus on characteristic problems and possible solutions. Temporary of the longest-running amphibian rescue operation in Hungary organized by Varangy Akció- mitigation measures are made by several organisations. They often focus on the breeding csoport Egyesület. Tunnels of different sizes were constructed, the diameter of new tunnels migration towards the breeding sites only, which may lead to situations when the erected were either 40 or 60 cm, already existing culverts included into the system were over 100 cm. fences become barriers for amphibians leaving the breeding sites. Besides positioning, the set However, the new system built by the local national park hardly reduced amphibian road kills of fences is often without following basic rules, e. g. how to make the edge of fences to avoid in the 2007 and 2008 migration period, its use was below 0.5%. Initiated by Varangy Akció- animals to get onto the roads in large numbers. However, these actions save well over an csoport Egyesület, maintenance was made by the local road authority in spring 2009 and by annual 100,000 amphibians and reptiles (mainly grass snakes), on average. The priorities of the Danube–Ipoly National Park Directorate in 2010 and 2011. As a result, at least 8.8–14.6% such activities should also be enlargened, besides focussing on common species dying on of the migrating amphibians crossed the road in the tunnels in 2009–2011. Still, more than roads in large numbers, rare species and small populations should also be given priority when ten thousand amphibians died on that road section during every spring migration between decisions are made in what areas to intervene. Permanent mitigation measures can be even 2009 and 2011 even though toad rescue was also carried out by volunteers. In 2012 migration more effective, the most successful, modern design in the Danube Tisza Interfluve created in was much less intensive than in the other years due to severe drought. We observed the 2011 saves more amphibians than temporary mitigation measures as it functions during the corpses of seven anuran and two newt species along the road, nearly 90% of them were com- whole year including the juvenile dispersal period, which is completely overlooked at tem- mon toads (Bufo bufo). In the first half of the migration period Pelobates fuscus, Rana dal- porarily protected road sections. However, most of these constructions do not work properly matina and Lissotriton vulgaris, in the middle and the final stages of the migration B. bufo due to unproper design and realisation or the lack of maintenance, which means a conserva- and Bombina bombina migration peaks were observed. Depending on the weather, the start tion failure as well as waste of money. and the peak of the migration varied among the studied years by 20 and 10 days, respectively. Unfortunately, no maintenance was made in 2012 that lead to the gradual disintegration of the plastic fences. Marked differences were found among the tunnels, common toads (Bufo bufo) significantly more frequently crossed the road in large culverts than in smaller tunnels.

70 71 Monitoring of amphibians and reptiles in the frame of the Habitats Directive Dozen years of herpetological monitoring in Hungarian Biodiversity Monitoring amphibians and reptiles of community interest Monitoring System in Romania

Botond Bakó Dan Coga˘lniceanu 1, Laurentiu Rozylowicz 2, Tibor Hartel 3

Ministry of Rural Development, Department of Nature Conservation, Kossuth Lajos tér 11, 1055 Budapest, Hungary; 1 University Ovidius Constant¸a, Faculty of Natural Sciences and Agricultural Sciences, Aleea Universita˘¸iit 1, corp B, [email protected] 900740 Constant¸a, Romania; [email protected] 2 University of Bucharest, Centre for Environmental Research (CCMESI), N. Balcescu Blvd. 1, 010041 Bucharest, Altogether 18 amphibian and 15 reptile species are known from Hungary, of which all are Romania 3 Sapienza University, Department of Environmental Sciences, Deva St. 19, 400375 Cluj Napoca, Romania included in the annexes of the Habitats Directive, meaning that they are protected or strictly protected. The standardised, long-term study of Hungary’s amphibian and reptile fauna Romania has a high diversity of habitats and landscapes and covers five biogeographical regions. began in 2001. Monitoring under the Hungarian Biodiversity Monitoring System (HBMS) Half of the 41 protected amphibians and reptiles occurring in Romania reach the limits of their initially covered various habitats in seven regions of Hungary. These surveys were carried out geographical range here, being more prone to range shifts (contractions or expansion). under guidance of specialised experts, and following HBMS protocols for amphibians and Romania joined the European Union (EU) in 2007 but started the process of implementing reptiles, using numerous methods. the environmental legislation and designating Natura 2000 sites in 2005. Despite the pro- gresses made in developing the Natura 2000 network, which now numbers 383 SCIs and 148 Sampling methods for monitoring amphibians SPAs and covers 22.66% of the land surface, little has been done in promoting a scientifically Night search by torch light, bottle-trapping: methods suitable for sampling Caudata. sound and coherent monitoring system, as required by article 17 of the Habitats Directive. Egg counts: a method suitable for the assessment of the number of reproducing females of We identified several major issues needed for complying with this requirement and for sup- toads, agile frogs and common frogs. porting the development of a comprehensive network of protected areas: Count of adult animals along shorelines (fire salamander assessment can be done on land as (1) Monitoring should be carried out at a biogeographical region scale, keeping in mind that well). the goal of monitoring is to detect range/population trends, and not just provide raw data Bottle trapping for Triturus species. for each Natura 2000 site. The average size (± SD) of 108.41 (± 314.98) km2 of a SCI, does not allow identification of trends in range changes; Sampling methods for monitoring reptiles (2) Agreeing upon regional scale favorable conditions for each species of Community interest One of the possible methods is transect sampling, during which the numbers of individuals in terms of acceptable limits for a minimum set of attributes is needed before the start of of the observed species are recorded. It is recommended to invest the same time effort during a monitoring program; each sampling. Another method is quadrat sampling, which is used when features of the (3) Agreeing upon a common working methodology, including data collecting, management sample site do not allow transect sampling to be performed. and analysis; (4) Undertaking a baseline survey that will provide a starting reference condition; A new monitoring method (5) Cooperating with neighboring countries for a regional scale assessment; The collected monitoring data were not always reliable (different weather, different observ- (6) Identifying the nature of threats and if/how these can be mitigated, by involving local ability, changing habitat or microhabitat, etc.). communities and other stakeholders. In 2011 for the better observability and gaining more information we designed a not habitat- based but a new species-based monitoring methodology for Caudata species, and some Anura and three reptile species, too.

74 75 Assessment of favourable conservation status for herpetofauna in Conventional field methods vs. eDNA-based species monitoring. the Netherlands A comparative assessment for three aquatic species under the EU Habitats Directive Edo Goverse, Jeroen van Delft, Ronald Zollinger Jos Kielgast 1, Lars L. Iversen 1,2, Philip Francis Thomsen 3, Michael Hastrup 4, RAVON Foundation, P.O. Box 1413, 6501BK, Nijmegen, The Netherlands; [email protected] Søren Oldenburg 4, Martin Hesselsøe 1

According to the EC Habitats Directive, article 17, Member States have to report in a regular 1 Amphi Consult Aps, Fruebjergvej 3, 2100 København Ø, Denmark; [email protected] six-yearly progress on the conservation status of Europe’s most vulnerable habitat types and 2 Freshwater Biology Section, University of Copenhagen, Helsingørgade 51, 3400 Hillerød, Denmark 3 species. As part of this assessment, Favourable Reference Values (FRVs) are crucial. In 2008, Center for GeoGenetics, Natural History Museum of Denmark, University of Copenhagen, Øster Voldgade 5–7, 1350 København, Denmark in the first Dutch report on this topic, the assessment of the conservation status of the species 4 The Danish Nature Agency, Nature and Water, Fejøgade 1, 4800 Nykøbing F., Denmark was based on expert judgements and a description of the species. In the next report, which will be published summer 2013, the Dutch government changed the methodology in describing Reliable species monitoring is a crucial foundation for the implementation of most conser- the Conservation status. Favourable Reference Population Size (FRPs) and Favourable Refe- vation projects. However, for many species it remains a challenge owing to field methods that rence Range (FRRs) are now the most important figures to assess the Conservation status of are hard to standardize and depend heavily on taxonomic expertise in decline. In recent years a species. The year 1994 is used as year of reference. Details of the methodology are presented a new DNA-based approach has been proposed to address this problem in aquatic animals. as well as the outcome for Annex II and IV species such as the great crested newt, based on It has been demonstrated for several rare species that even minute volumes of water from the large database collected largely by volunteers. their habitat contain environmental DNA (eDNA) that can be used to confirm their con- temporary presence. However, little is known about the comparative performance of this new monitoring strategy compared to conventional field surveys. We tested the eDNA detection approach in the context of the current Danish national monitoring regime for three aquatic species on the Habitats Directive Annex II and IV. Our results support previous studies on amphibians and validate a high level of sensitivity. The great creasted newt was detected by eDNA at 39% of the investigated localities corresponding with 97% of the those obtained by conventional monitoring. The common spadefoot toad was detected at 22% of the moni- tored localities by eDNA including eight where conventional monitoring failed to find the species. On the contrary, the large white-faced darter was only detected by eDNA at half of the localities (26%) compared to results of conventional monitoring and exhibited indications of specificity issues. These findings clearly illustrate that there are vast differences in the effi- ciency of eDNA monitoring across taxa and underline the importance of thorough com- parative documentation. The use of eDNA in monitoring is clearly in its infancy and there are many avenues of research that can be expected to improve the utility of this general prin- ciple. Nonetheless, these first comparative results do indicate that eDNA detection may have a promising future in species monitoring under the EU habitats directive.

76 77 First attempt to set up the national monitoring scheme for four target Assessing the occurrence of the great crested newt (Triturus cristatus) amphibian species in Slovenia using water samples: is the method more efficient than the direct observation? K. Poboljsˇaj, M. Cipot, A. Lesˇnik, M. Govedic´ Sylvain Ursenbacher 1,2, Benedikt R. Schmidt 2,3 Centre for Cartography of Fauna and Flora, Antolcˇicˇeva 1, Miklavzˇ na Dravskem polju, Slovenia; [email protected] 1 Institut für Natur-, Landschafts- und Umweltschutz (NLU), Universität Basel, St. Johanns-Vorstadt 10, 4056 Basel, Switzerland; [email protected] The Centre for Cartography of Fauna and Flora carried out the project “Setting up the mon- 2 Karch, Passage Maximilien-de-Meuron 6, 2000 Neuchâtel, Switzerland itoring of target amphibian species in Slovenia” between 2010 and 2011. Target species were 3 Institute of Evolutionary Biology and Environmental Studies, University of Zurich, Winterthurerstraße 190, Italian crested newt (Triturus carnifex), Italian agile frog (Rana latastei), yellow-bellied toad 8057 Zürich, Switzerland (Bombina variegata) and fire-bellied toad (Bombina bombina). This was the first attempt to set up a monitoring scheme for Habitats Directive Annex II species of amphibians in Slovenia During the last years, the development of environmental DNA (eDNA) allowed the detec- on the basis of the analysis of existing data and carrying out additional surveys. tion of species in aquatic environment only based on a very few number of cells of the target As a result there is a proposal for monitoring scheme for each species in terms of both types species. Recent publications showed the possibility to assess the occurrence of an amphibian of monitoring (species distribution and population size). Since there are significant differences or fish species in a pond or river based only on a single water sample (15 ml). In some exam- in species distribution and ecology, we took an individual approach in designing monitoring ples, the detection was even better based on the eDNA method compared to direct obser- schemes. vation. However, the efficiency of both eDNA and direct observation approaches has never For T. carnifex there are still a lot of gaps in known distribution in Slovenia, therefore the been properly tested. distribution monitoring should primarily aim at surveying the areas with potential newt In 2012, we sampled three times 29 ponds in the Basel region (Switzerland) and visually habitats. We also proposed population monitoring on 27 known sites and prepared the pro- detected the occurrence of the great crested newt (Triturus cristatus). The water samples were tocol that allow comparison over time within each site and between sites. later analyzed in an adequate laboratory with specific methodology used for ancient and low In 2009 there was also the first morphological evidence for the presence of Danube crested quantity DNA. Real time PCR with specific primers were conducted to evaluate the amount newt (Triturus dobrogicus) in the Prekmurje region, NE Slovenia. Therefore we need also addi- of eDNA belonging to the focus species and we compared this results with direct observation. tional research on the distribution of the species and potential hybrid zone with T. carnifex. Site occupancy models were used to evaluate the detection probability by both methods. The direct observation (detection probability = 0.73 ± 0.06) was slightly better in our study area compared to the eDNA (0.61 ± 0.07). When conducting the analyses for both methods separately, the eDNA method overestimated the detection and underestimated the occupancy. The probability of detection using eDNA is also strongly associated with the number of crested newts observed in the pond. In conclusion, eDNA is a good and simple method to detect organisms, even if not neces- sarily better than other ones. This method is not perfect (as other ones), should not replace direct observation, but can be used in complement, especially when the target species is difficult to detect.

78 79 Educative and participative monitoring for a wider and more effective Conservation status mapping of great crested newts in the UK amphibian conservation, applicable on a wider European scale? J. W. Wilkinson 1, A. P. Arnell 1, J. Foster 2, A. H. Gent 1 Wouter de Vries 1, Edo Goverse 2, Adolfo Marco 1,3 1 Amphibian & Reptile Conservation Trust, 655a Christchurch Road, Bournemouth BH1 4AP, United Kingdom; [email protected] 1 Asociación AMBOR, Ctra. Constantina, El Pedroso km 1, 41450 Constantina, Spain; [email protected] 2 Amphibian & Reptile Conservation Trust, The Witley Centre, Haslemere Road, Godalming, Surrey GU8 5QA, 2 RAVON Foundation, P.O. Box 1413, 6501BK Nijmegen, The Netherlands United Kingdom 3 Estación Biológica de Doñana, CSIC, c/ Americo Vespucio, s/n, 41092 Sevilla, Spain The great crested newt is the only Habitats Directive Annex II species of amphibian in the In 2011 a new monitoring design was developed by Association AMBOR and RAVON UK. Since the 1970s there have been various attempts to define status and to assess trends, Foundation with the aim of obtaining comparable data on the status of amphibian popula- with estimates given of 50% declines during the 1960s–1970s, and 2% changes every five tions and habitats in a Spanish natural park of 177,000 ha with low accessibility: Educative years in the 1990s. National population estimates have produced widely ranging estimates and Participative Monitoring for Amphibian Conservation (EPMAC). Since 2011 annual based on numbers of ‘sites’ or ‘breeding populations’ and the focus for recent work has been monitoring were executed twice in 110 preselected sites with involvement of professional and to provide data for the reporting requirements of the Habitats Directive. The data collected amateur herpetologists (national and international), local community, nature managers and are, however, not ideal for helping ‘on the ground conservation action’ or for providing infor- biology students. An auto-financed educative system was set-up and the results are of high mation that helps with issues where the species is affected by development. value to nature conservation as well as for understanding the natural dynamics and involving Amphibian and Reptile Conservation is developing fine-scale modelling tools to predict the local community by a constructive educational method. During the three years around distribution of great crested newts, target surveys and conservation action (e. g. habitat link- 150 persons were involved and a large spin-off effect was observed. For 2013 we have applied ages), and assess the effects of putative developments on local conservation status. These goals for various small grants to set-up additional areas with this monitoring system in other contribute to an integrated surveillance strategy for the species. We demonstrate these regions in Europe among others to evaluate the applicability on a wider European scale. The approaches with examples from the great crested newt ‘hotspot’ in North-east Wales. An direct involvement of locals together with professional participants make it possible to obtain understanding of the effects of development and changes in pondscape connectivity, and their a wider understanding of the natural dynamics and population dynamics, get in contact with impacts on conservation status, can inform sustainable development whilst delivering posi- local communities and create awareness. This stimulates more effective conservation activities, tive conservation benefits in the long term. increase knowledge and experience of professionals or nature managers. We would like to suggest the applied approach of monitoring for other areas with high nature values, low local involvement and low accessibility. We will present the methods, results, limitations and dif- ficulties of three year monitoring in southwestern Spain and the evaluation of the experiences for launching the method in other regions.

80 81 Oral Presentations PRACTICAL CONSERVATION SYSTEMATICS

Understanding nesting ecology and behaviour of green sea turtles: Cryptic speciation patterns in Iranian rock lizards uncovered by intensive monitoring and ground survey as a method of conservation integrative taxonomy

Aini Hasanah Binti Abd Mutalib 1, Nik Fadzly N Rosely 1,2, Nurolhuda Nasir 3 Faraham Ahmadzadeh 1,2, Morris Flecks 2, Miguel A. Carreteros 3, Omid Mozaffari 4, Wolfgang Böhme 2, D. James Harris 3, Susana Freitas 3, Dennis Rödder 2 1 School of Biological Sciences, Universiti Sains Malaysia, 11800 Penang, Malaysia; [email protected] 2 Centre for Marine and Coastal Studies (CEMACS), Universiti Sains Malaysia, 11800 Penang, Malaysia 1 Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti 3 World Wide Fund for Nature (WWF), 47400 Petaling Jaya, Malaysia University, G.C., Evin, Tehran, Iran; [email protected] 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany Monitoring and ground survey on the nesting beach provide sightings on the population 3 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de status and conservation of green sea turtles, Chelonia mydas in Setiu, Terengganu, Malaysia. Vairão, 4485-661 Vairão Portugal 4 This method is carried out especially to overcome anthropogenic threats in the area such Aria Herpetological Institute, No. 1, 4 Parastoo St., Shahrak-e-Homa, Ashraf Esfehani Highway, Tehran, Iran illegal poaching, excessive noise and light pollution, unplanned development near the coast While traditionally species recognition has been based solely on morphological differences and so on. We collected the distribution data from 2007 until 2012, and recorded that the either typological or quantitative, several newly developed methods can be used for a more highest number of nesting occurred in 2012 (mean=28.714). Nesting attempts are recorded objective and integrative approach on species delimitation. This may be especially relevant mostly in May (mean=33.333±4.539, p<0.001). We also mapped 98 nesting areas in reserved when dealing with cryptic species or species complexes, where high overall resemblance be- Telaga Papan Beach, Setiu within six months period (March until August) in 2012 by using tween species is coupled with comparatively high morphological variation within populations. Global Positioning System (GPS). By plotting the areas into ten plots, we observed the Rock lizards, genus Darevskia, are such an example, as many of its members offer few diag- nesting frequency in each plot and relate it to human activities and land-use. We recorded nostic morphological features. Herein, we use a combination of genetic (two nuclear and two that Plot 4 has the most frequent nesting activities with least human disturbance and land- mitochondrial loci), morphological (15 morphometric, 16 meristic and four categorical char- use. We recorded that most attempts occurred at dunes and emergence hour were most fre- acters) and ecological (11 newly calculated spatial environmental predictors) criteria to delimit quently at 12 am–1.59 am (23%). During this extensive survey in Telaga Papan, we measured cryptic species within two species complexes, D. chlorogaster and D. defilippii, both distributed the curved carapace length (mean=98.369 cm) curved carapace width (mean=85.595 cm) and in northern Iran. Phylogenetic analyses of the molecular data confirmed the monophyly of clutch size (mean=81.602 eggs) of the nesting marine turtles. In this study, no significant cor- D. chlorogaster, while D. defilippii is paraphyletic in respect to D. steineri. However, each of relation recorded between both clutch size with curved carapace length (rs=0.2339, p=0.1359, the complexes comprises several highly divergent clades, especially when compared to other p>0.05). Out of 151 nesting attempts, 35.1% false crawls are recorded and both variables (nest- congeners. We identified seven candidate species within each complex, of which three and ing attempts and false crawls) are strongly correlated (rs=0.8827, p=0.0198). We suggest four species are supported by Bayesian species delimitation within D. chlorogaster and D. that strict monitoring by adequate number of rangers is to be done to reduce illegal poach- defilippii (including D. steineri), respectively. Although the species within one complex lack ing and other disturbances from human activities. As trainings are continuously provided clear diagnostic features, they can be well separated based on morphological variables when to these rangers, these threats can be reduced and conservation status of green sea turtles sample size is appropriate. Ecological Niche Modeling provided additional support for the can be improved from time to time. identified species and niche overlaps between them are generally low, especially in the D. defilippii complex.

84 85 ECOLOGY DISTRIBUTION MODELLING

Rapid lizard radiation lacking niche conservatism: ecological diversification within a complex landscape Analysis of distribution of Acrodonta lizards in Vietnam as a tool for designation of Important Herpetofaunal Areas Faraham Ahmadzadeh 1,2, Morris Flecks 2, Miguel A. Carretero 3, Wolfgang Böhme 2, Çetin Ilgaz 4, Jan O. Engler 2,5, D. James Harris 3, Nazan Üzüm 6, Dennis Rödder 2 Natalia B. Ananjeva

1 Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti Division of Herpetology and Ornithology, Zoological Institute, Russian Academy of Sciences, Universitetskaya nab. 1, University, G.C., Evin, Tehran, Iran 99034 St.Petersburg, Russia; [email protected] 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; [email protected] 3 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Results of faunistic, taxonomic and phylogenetic study of agamid lizards in the fauna of Vairão, 4485-661 Vairão, Portugal 4 Dokuz Eylül University, Fauna and Flora Research and Application Center, 35150 Buca, Izmir, Turkey Vietnam and data on their distribution are summarized and analyzed. Taxonomic diversity 5 Biogeography Department, Trier University, 54296 Trier, Germany of agamids and representation of different evolutionary lineages in different regions of South- 6 Adnan Menderes University, Faculty of Science and Arts, Department of Biology, 09010 Aydin, Turkey east Asia are considered. Composition of the fauna of agamids in Vietnam and ratio of endemic and wide distributed species is discussed in biogeographic aspect. Based on data on Diversification and rapid radiation are well documented in lacertid lizards. Niche conservation distribution of endemic agamids within Vietnam territory we offer to designate Important is frequently observed among related taxa, whereby niches will not change much during spe- Herpetofaunal areas in mountain tropical forests in southern regions of Central Vietnam on ciation events. Here, we investigate the relationship between environmental niche divergence the Tay Nguyen Plateau. and phylogenetic relatedness in a widespread group of green lizards, the Lacerta trilineata group. This study is partially supported by RFBR 12-04-00057-a and RFBR-Viet 12-04-93005. A dated phylogeny based on three mitochondrial genes was contextualized using species dis- tribution models of all genetically identified lineages in the Lacerta trilineata group. Based on this analysis, ancestral climatic niche occupancy was reconstructed using niche occupancy pro- files. Niche divergence among lineages was quantified by computing multivariate niche over- laps. All taxa are associated with humid areas, but there is extensive variation in their climatic niche breadths and positions, which accord with the main phylogenetic split in the group. Our results suggest divergent niche evolution within subclades and convergent evolution among clades, which implies only a limited degree of niche conservatism regarding annual variations in temperature and precipitation. In contrast, niche axes—mainly reflecting precipitation patterns of the coldest quarter—show a greater difference among clades than within clades, and therefore a higher degree of niche conservatism. Based on estimated divergence times between taxa and geological events in Anatolia, our results can be explained by fragmentation of the range of a hypothetical ancestral species, resulting in different adaptations of subclades either to humid continental climates or to more Mediterranean climates. Our study highlights deviations from classical niche conservatism theory due to significant niche shifts among sister taxa.

86 87 SYSTEMATICS SYSTEMATICS

A re-evaluation of the taxonomic status of the foam-nest treefrogs, How complex is the Bufo bufo species group? Chiromantis in Thailand based on morphological, molecular and bioacustic evidence Jan W. Arntzen 1, Jacob McAtear 1, Ernesto Recuero 2, Janine M. Ziermann 1,4, Annemarie Ohler 3, Jacques van Alphen 1, Inigo Martinez-Solano 2 Anchalee Aowphol 1, Attapol Rujirawan 2, David S. McLeod 3 1 Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands; [email protected] 2 Instituto de Investigación en Recursos Cinegéticos (IREC), CSIC-UCLM-JCCM Ronda de Toledo, 1 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Bangkok, Thailand; [email protected] s/n 13071 Ciudad Real, Spain 2 Department of Zoology, Faculty of Science, Kasetsart University, 10900 Bangkok, Thailand 3 Muséum National d’Histoire Naturelle, Département Evolution et Systématique, UMR 7205 CNRS Origine, 3 University of Kansas Biodiversity Institute, 1345 Jayhawk Boulevard, Lawrence, KS 66045-7561, USA Structure et Évolution de la Biodiversité, 25 rue Cuvier, CP 30, 75005 Paris, France 4 Present address: Department of Anatomy, Howard University, College of Medicine, 520 W St. NW, Washington, With a wide distribution in two continents, some rhacophorid frogs of the genus Chiromantis DC 20059, USA exhibit phenotypic similarities. The Asian species C. hansenae has been considered a junior synonym of C. vittatus. We assessed the taxonomic status and phylogenetic relationships of The common toad (Bufo bufo) has an enormous range, from Tunesia to deep into Russia. congeneric species of Chiromantis frogs from Thailand based on three types of evidence. A range-wide recent study revealed a marked genetic differentiation, with one species in and Analyses of morphological, molecular and bioacoustic data sets support four evolutionarily adjacent to the Iberian Peninsula (Bufo spinosus), two species in the Caucasus and the—still very distinct and monophyletic clades: C. doriae, C. nongkhorensis, C. vittatus and C. hansenae. wide ranging—Bufo bufo. We expanded upon our work as follows. (1) With mitochondrial Genetic divergence between C. vittatus and C. hansenae is significantly greater than between and nuclear gene sequences, we confirmed the deep split and that the mutual range border of C. doriae and C. nongkhorensis. These four Chiromantis species show distinct call characteristics. B. bufo and B. spinosus is in France. (2) We searched for morphological characters that would Multivariate analyses showed morphological differentiation within Thai Chiromantis species. discriminate the species in France and we tested some potentially informative characters on Our results support the taxonomic validity of C. hansenae and suggest that there may be more museum material (Leiden, Paris and Madrid) covering a North to South transect in western diversity within Chiromantis in Thailand than is currently recognized. Europe, from the UK to southern Spain. (3) We identified the position of the contact zone in France, from the Atlantic coast to the upper Rhone valley near Lyon, which gave us some clues about the “why” of morphological species differentiation. Finally, (4) we scrutinized published nuclear data to investigate claims of hybridization and extensive introgression among species across the Mediterranean.

88 89 MORPHOLOGY, ANATOMY, PHYSIOLOGY AND DEVELOPMENT BEHAVIOUR

Variation of sexual dimorphism in snakes: the case of Male throat UV colouration affects maternal investment in the Echis coloratus complex (Ophidia: Viperidae) European green lizards

Gergely Babocsay 1,2, Hervé Seligmann 3 Katalin Bajer 1, Orsolya Molnár 2, János Török 3, Gábor Herczeg 3

1 Mátra Museum of the Hungarian Natural History Museum, Kossuth Lajos utca 40, 3200 Gyöngyös, Hungary; 1 Laboratório de Biogeografia e Macroecologia, Universidade Federal do Rio Grande do Norte, Centro de Biociências, [email protected] Departamento de Botânica, Ecologia e Zoologia, Campus Universitário, Lagoa Nova, Natal–RN 59078-900, Brazil; 2 Department of Ecology, Evolution and Behavior, The Hebrew University of Jerusalem, 91904 Jerusalem, Israel [email protected] 3 Herpetological Collection, National Natural History Collections, The Hebrew University of Jerusalem, 2 Department of Biological Sciences, Darthmouth College, Life Sciences Center, 78 College Street, Hanover, 91904 Jerusalem, Israel New Hampshire 03766, USA 3 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, Using museum specimens, we examined sexual differences in size, in 54 morphological char- Pázmány Péter sétány 1/c, 1117 Budapest, Hungary acters and in allometric growth in three populations of the Echis coloratus complex: E. c. col- According to life-history theory, individuals should increase their reproductive effort when oratus (n=50) from the southern Levant, E. c. terraesanctae (n=70) and E. omanensis (n=32). fitness benefits of reproduction are high. Differential allocation hypothesis predicts that it Significance of differences in meristic characters was tested using two-tailed T-test and Chi- can be advantageous for females to invest more heavily into offspring sired by males of high square test. Differences between sexes in the slope of allometric growth were tested by quality and decrease investment into future reproduction. However, there are species in which ANCOVA. In many characters conventionally regarded as sexually dimorphic, all populations even opposing reproductive strategies are present, and females compensate for low offspring showed sexual dimorphism. Differences were not tested for causation, but we propose possible quality by increased maternal investment. Animals have developed various costly sexual sig- functional explanations. The largest number of characters showing sexual dimorphism oc- nals to advertise their quality. Here, we investigated whether European green lizard females curred in E. omanensis (20) followed by E. c. coloratus (16) and E. c. terraesanctae (15). Males adjust their maternal investment according to males’ nuptial UV colouration, a trait that were larger than females, possibly due to their agonistic behaviour, had longer tales presum- plays an important role in mate choice. We experimentally manipulated the UV colour of the ably as a result of need for more room for hemipenes. Females had more ventrals, more dor- male pairs, where the members of the pair did not differ significantly in their morphological sals along the entire length of the body, and with a varying level of significance, tended to characters, or throat colouration measured prior to manipulation. After the mate choice have more scales in head characters. All these female biased differences may be attributed to experiments, half of the females mated with their preferred male, while the rest had only the fecundity related increased bulk and swallowing capacity. Numbers of body somites (ventrals option to mate with the male they rejected during the trials. Females showed preference +subcaudals) did not differ between sexes, but the longer males had more dorsal blotches toward males with higher relative UV intensity. Maternal investment was affected by the than females. We suggest that this helps to maintain a blotch frequency necessary to achieve males’ UV colouration, because males differed systematically only in this character, and a flicker fusion effect of the blotchy pattern to promote visual illusion. Echis coloratus females, clutches sired by preferred males included heavier eggs. Clutch size was affected by the male unlike males, showed an ontogenetic change in relative tail length. Snakes of this population body size. Our result support the predictions of positive differential allocation, as females tend to be arboreal at adult age, that necessitates long tails which may be a priori available invested differently into clutches sired by different males of quality, with higher investment for males but not for females. To substantiate our proposed functional explanations testing to those of sired by attractive males. Moreover, increased investment to these clutches was with possible environmental correlates or in experimental sets is necessary. based on the direct preference showed for the male throat UV colouration manipulated within the natural range. Taken together, our results suggest that UV colour signals can be a cue of male quality and affect the patterns of maternal investment.

90 91 BEHAVIOUR BEHAVIOUR

Green lizard (Lacerta viridis) personalities: behaviour and individual Geographic variation in the signaling behaviour of the Jacky dragon, quality at different ontogenetic stages Amphibolurus muricatus

Katalin Bajer 1, Gergely Horváth 2, Orsolya Molnár 3, János Török 2, Gábor Herczeg 2 Marco D. Barquero 1,2, Richard Peters 3, Martin Whiting 1

1 Laboratório de Biogeografia e Macroecologia, Universidade Federal do Rio Grande do Norte, Centro de Biociências, 1 Department of Biological Sciences, Faculty of Science, Macquarie University, Sydney, New South Wales 2122, Departamento de Botânica, Ecologia e Zoologia, Campus Universitário, Lagoa Nova, Natal–RN 59078-900, Brazil Australia; [email protected] 2 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, 2 Universidad de Costa Rica, Montes de Oca, 2060 San José, Costa Rica Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 3 Department of Zoology, La Trobe University, Bundoora, Victoria 3086, Australia 3 Department of Biological Sciences, Darthmouth College, Life Sciences Center, 78 College Street, Hanover, New Hampshire 03766, USA Animal communication systems are extremely complex and diverse. Signal diversification has been shown to be the result of several mechanisms, including genetic differentiation, natural The importance of animal personality became well recognised in behavioural ecology during selection and sexual selection. However, the influence and interaction of these mechanisms the past decade. Nevertheless, our knowledge about the mechanisms in the evolution and are still far from being understood, especially in a within-species context. We use an agamid development of animal personality is still incomplete. For instance, the link between animal lizard endemic to Australia, the Jacky dragon (Amphibolurus muricatus), to examine relation- personality and individual quality has rarely been established. In male European green lizards ships between genetic structure, morphology and signaling behavior. We also investigate the (Lacerta viridis), components of nuptial throat colorization has been linked to individual consequences of this variation on the ability to discriminate conspecifics. We used three pop- quality and fitness. In this present study, we first measured exploration-boldness in adult ulations separated by over 280 km, two of which belong to the same genetic clade. We found male European green lizards during the reproductive season, and second, we measured the that individuals from the more closely-related populations exhibited more similarities both same behaviour in their 2–3 weeks old laboratory-reared offspring. Our aims were to see if (i) in morphology and signaling behaviour than lizards from the third, more distant population. animal personality was present, (ii) there were correlations between behavioural type/be- In addition, animals from all populations showed different levels of aggression if confronted havioural consistency (the main components of individual personality) and individual quality, with individuals from the same or different populations. We discuss the significance of this and (iii) behavioural type/behavioural consistency were heritable. Both adults and juveniles concordant pattern between genetic structure, morphology and behaviour. showed strong personality. Males with brighter throat colour were more explorative-bold, and larger males were more consistent. However, we did not find any correlation in juveniles. Behavioural type differed between the F1 families, but behavioural consistency did not. Neither behavioural type, nor behavioural consistency showed heritability in our father- offspring regressions, however, we note that the assumption of equal age of fathers and off- spring was obviously violated. We conclude that European green lizards have strong ‘built in’ personalities, but its link to individual quality emerges only during ontogeny. Based on earlier studies, we suggest that better quality males are more explorative-bolder and more consistent than worse males.

92 93 ECOLOGY BEHAVIOUR

Population dynamics of the North African green water frog Observations of reproductive behaviour in Salamandra species Pelophylax saharicus in four lakes from northern Tunisia Sergé Bogaerts 1, Max Sparreboom 2, Wouter Beukema 3, Frank Pasmans 4 Meher Bellakhal, Mouna Fartouna Bellakhal 1 Lupinelaan 25, 5582CG Waalre, The Netherlands; [email protected] 2 Naturalis Biodiversity Centre, P.O. Box 9517, 2300RA Leiden, The Netherlands Exploitation of Aquatic Environments, High Institute of Fisheries and Aquaculture, Errimel, B. P. 15, 7080 Bizerta, 3 Cátedra Rui Nabeiro, Universidade de Évora, Casa Cordovil, Rua Dr. Joaquim Henrique da Fonseca, Tunisia; [email protected] 7000-890 Évora, Portugal 4 Laboratory of Veterinary Bacteriology and Mycology, Clinic for Exotic Animals, Faculty of Veterinary Medicine, Amphibians are declining more rapidly than other animals in many areas but the data on Ghent University, Salisburylaan 133, 9820 Merelbeke, Belgium amphibian populations are usually given as unadjusted counts, which explain why it is not easy to understand their decline. Most surveys are concerned only with the presence-absence Fire salamanders (Salamandra) mate at night, often in cold and rainy weather. They lead a of species and not with trends within populations. To determine relationships between the secretive life and for this reason observations of their reproductive behaviour in the field are density of tadpoles, froglets and adults of Pelophylax saharicus, many sessions of mark-release- rare. Much of the salamanders’ courtship behaviour has been documented from observations recapture were conducted in four lakes from north of Tunisia. Densities of tadpoles were re- in captivity. The behavioural repertoire of many fire salamanders is now well-known, although ducing between August and September, but did not differ between sites. In September and not every aspect has been explained. On a recent excursion to the Middle Atlas in Morocco October, the froglet densities in two of the lakes were always higher than in the other two. in December 2012, we had the chance to observe Salamandra algira during mating activity. Similarly, densities of adults remained higher in the two first lakes. These results could be ex- Thus far, courtship behaviour in this species was documented only fragmentary, but we plained by a smaller size of the two first water bodies, i. e. with a higher edge effect. However, managed to make a video of a full courtship sequence and will present it during our talk. for each site there was no difference between years. The yield of tadpoles ranged according Salamandra algira behaviour appears to be similar to that of other fire salamanders. Recently to sites from 31 to 65 ind. per mature female. Survival from the egg to the froglet stage ranged various videos of fire salamander behaviour have appeared on YouTube. We compare some from 4.9 to 8.5‰ and survival between froglets and subadults from 31 to 45%. of these observations and discuss the interesting aggressive male-male interactions, which have puzzled researchers since Werner Kästle’s first account of male interaction during the third SEH meeting in Prague in 1985.

94 95 BEHAVIOUR DISEASES AND PARASITES

Influence of termitophagy on prehension mode in cordylid lizards: Designing screening protocols for amphibian disease that account for a morphological and kinematic analysis imperfect and variable capture rates of individuals

Chris Broeckhoven, P. Le Fras N. Mouton Stefano Canessa 1, An Martel 2, Frank Pasmans 3

University of Stellenbosch, Private Bag X1, Matieland, 7602, South Africa; [email protected] 1 ARC Centre of Excellence for Environmental Decisions, School of Botany, University of Melbourne, 3010 Victoria, Australia; [email protected] Although there has been considerable interest in understanding the complexity of prehension 2 Faculty of Veterinary Medicine, Ghent University, Salisburylaan 133, 9820 Merelbeke, Belgium mode in lizards, little insight has been gained into the selective pressures underlying shifts between different prehension modes. The Cordylidae family offers ideal opportunities to Given the threat posed by the amphibian chytrid fungus Batrachochytrium dendrobatidis, evaluate whether a specialist diet may affect prehension mode, as a bimodal prehension mode accurate knowledge of its presence and prevalence in an area is needed to trigger conservation is well established in the sister families Scincidae and Gerrhosauridae, and dietary specialisa- actions. However, imperfect capture rates determine the number of individuals tested during tion is present in at least one species, Ouroborus cataphractus. The southern harvester termite field surveys and increase the uncertainty surrounding estimates of prevalence. We combined (Microhodotermes viator) is the main prey item of O. cataphractus and has played an impor- a model for estimation of capture rates of individuals and a beta-binomial model of preva- tant role in the evolutionary history of this species. We used high-speed filmography to test lence estimates to select the seasonal window of sampling that would provide the most precise the effect of prey species and prey size on prehension mode by comparing prey prehension estimates of prevalence. We applied the models to data from a screening program for three between O. cataphractus and its closest extant relative, Karusasaurus polyzonus. In addition, populations of the endangered Apennine yellow-bellied toad Bombina variegata pachypus in we included several cordylid lizards representing the major clades in the family into our northern Italy. We estimated population size and capture rates from mark-recapture data and analysis to investigate whether any functional specialisations observed in O. cataphractus rep- combined them with estimates of chytrid prevalence obtained from screening the populations resent autapomorphies. Finally, we investigated the morphology of the lingual structure in 2011. We evaluated the model by predicting the size and precision of prevalence estimates in Cordylidae, with emphasis on O. cataphractus and K. polyzonus, using light and scanning that would result from seasonal fluctuations in recapture rates, and compared predictions electron microscopy techniques. Our results show that termitophagy in O. cataphractus has with data collected during additional surveys in 2012. resulted in the evolution of a novel lingual prehension mode, accompanied by morphological No individuals tested positive for chytrid: in 2011, prevalence was estimated between 0.04 modifications of the tongue structure. Moreover, none of the other cordylid lizards tested and 0.07 using uninformative priors. Sampling during the period of maximum capture rate in this study used lingual prehension during prey capture, except K. polyzonus which used the was predicted to decrease uncertainty by a maximum of 36% compared with least suitable tongue in a very small percentage of feeding trials. This data suggest that dietary specialisation periods, with greater gains when using uninformative priors. Using repeated, consecutive might underlie the reacquisition of a specific prehension mode or evolution of novel pre- surveys was also predicted to decrease upper 95% confidence intervals by between 20% and hension mechanisms in lizards. 36%. The results of surveys carried out in 2012 closely matched the predicted figures for all populations, suggesting this method can be reliably used to plan cost-effective surveillance programs and improve estimates of pathogen prevalence. These can then be more reliably used to inform conservation actions, for example allowing managers to declare “disease-free” populations with greater confidence.

96 97 BEHAVIOUR BEHAVIOUR

Investigating the degree of behavioural plasticity in lacertids by Predation pressure versus antipredator behaviour: a comparative study comparing two populations of Podarcis carbonelli under different using a lacertid model disturbance regimes Alex Cortada 1, Enrique García-Muñoz 1,2,3, Miguel A. Carretero 1 Miguel A. Carretero 1, Elena Argaña 2, Ricardo Duarte 3 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto Campus Agrário Vairão, 4485-661 Vairão, Portugal; [email protected] 2 de Vairão, 4485-661 Vairão, Portugal; [email protected] CESAM, Centro de Estudios de Ambiente o do Mar, Universidade de Aveiro, Campus Universitário de Santiago, 2 Centro de Investigação em Ciências Geo-Espaciais (CICGE), Faculdade de Ciências da Universidade do Porto, 3810-193 Aveiro, Portugal 3 Rua do Campo Alegre 687, 4169-007 Porto, Portugal Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Campus de las Lagunillas s/n, 3 CESAM & Departmento de Biologia, Universidade de Aveiro, 3810-193 Aveiro, Portugal 23071 Jaén, Spain

Antipredator strategies in lacertids vary across and within species. Previous investigations have Predation pressure plays a determinant role on animal populations selecting antipredatory documented divergent strategies between syntopic species, conspecific populations inhabiting strategies in the putative prey. Among them, antipredator behaviours are of interest when different habitats and colour morphs and size classes within the same population. Here, we analysing species interactions since selection may favour animals taking advantage of previ- evaluate the degree of plasticity in the different components of the antipredatory behaviour ous experiences to prevent new attacks. Lacertid lizards use escape as the most common tactics by comparing two conspecific populations under divergent disturbance regimes but other- when detected by a predator. Both escape and recovery vary across and within species. Here wise environmentally similar. We examined the lizard responses under direct attack from a we analysed in parallel the predation intensity and the antipredator behaviour to assess the predator (simulated by the researcher) in two populations of Podarcis carbonelli from geo- degree of intraspecific variation and the nature of predator-prey interactions. Size and sex- graphically close sites from Doñana National Park (SW Spain). In both cases, lizards mainly ual variation were also considered to infer eventual interactions between natural and sexual occupied wooden passes on sandy soil allowing park visitors to reach fauna observatories but, selection and the possible contribution of experience in predator avoidance. while one (El Acebuche) was heavily frequented the other (Ribetehilos) was not. Field ex- We studied six populations of the generalist lacertid Podarcis bocagei from NW Portugal, periments were carried out during a period with optimal temperatures but out of the repro- three located in open landscapes of coastal dunes and other three inhabiting agricultural areas ductive period (November 2010) to exclude factors other than site, sex and size class. We with granite walls. During spring days with suitable conditions for lizards’ activity, we recorded recorded variables describing lizard traits (sex, size class), environment (microhabitat, refuge) lizard traits (sex, size class), environment (microhabitat, refuge) and behavioural responses and behavioural responses for 128 observations throughout random walking routes. Lizards when simulating a predator attack for 80–100 lizards/population. Simultaneously, areal predator from the most disturbed population were closer to their refuges, regardless lizard and habitat pressure was estimated by placing plasticine models (100/site/day) to record predatory attacks. features. While lizards from this site were also observed on hotter substrates and males in The variation in morphology and ecophysiology of the same populations was already known. general escaped into hotter refuges, thermal environment was insufficient to explain the be- Among populations, predation intensity rather than microhabitat use appeared to be the havioural differences found. By contrast with previous studies comparing species or colour main source of variation in antipredator behaviour. Within populations, interaction between morphs, no variation between sites for either escape distance, recovery time or recovery dis- natural and sexual selection was observed both in escape and recovery behaviours with tance was recovered. Results reinforce previous claims that individual lizards may respond to females tending to be shier than males. The complex variation observed appear to be in agree- different levels of disturbance in a directional way. However, they also indicate that some ment with the predictions of the escape theory. Further manipulative experiments are needed components of the antipredator behaviour are more plastic that others. to disentangle between the effects of plasticity and selection.

98 99 ECOLOGY PRACTICAL CONSERVATION

Size variation in Italian Testudo hermanni: does it follow Bergmann’s Grazing effects on population densities of viviparous lizard rule? (Zootoca vivipara) in heathlands

Claudia Corti 1, Lara Bassu 2, Marta Biaggini 1, Anna Rita Di Cerbo 1, Raymond C. M. Creemers, Jeroen C. W. van Delft, Ronald Zollinger Nicoletta Di Francesco 3, Luciano Di Tizio 3, Carmen Fresi 2, Fabio Mastropasqua 4, 2 1 2 5 Valeria Nulchis , Antonio Romano , Maria Grazia Satta , Giovanni Scillitani , RAVON Reptile, Amphibian and Fish Conservation the Netherlands, P.O. Box 1413, 6501BK Nijmegen, Marco Zuffi 5 The Netherlands; [email protected]

1 Museo di Storia Naturale dell’Università di Firenze, Sezione di Zoologia “La Specola”, Via Romana 17, The viviparous lizard (Zootoca vivipara) is the most widespread and still the most abundant 50125 Firenze, Italy; [email protected] reptile in large parts of northwestern Europe. This small lacertid is a highly adaptable species, 2 Sezione Sardegna SHI, Societas Herpetologica Italica, c/o ALEA, Via Canepa 3, 09170 Oristano, Italy inhabiting a broad range of habitats. However, monitoring results (National Monitoring 3 Sezione SHI Abruzzo-Molise, Societas Herpetologica Italica Network) showed a significant decline in population sizes in the Netherlands over the last 20 4 Sezione SHI Puglia, Societas Herpetologica Italica 5 Dipartimento di Biologia, Sezione di Biologia Animale e Ambientale, Università di Bari “Aldo Moro”, years. The species has almost completely disappeared from agricultural areas and populations Via Orabona 4a, 70125 Bari, Italy are now confined to protected nature reserves and National Parks. Even in these protected 6 Museo di Storia Naturale e del Territorio, Università di Pisa, Via Roma 79, 56011 Calci, Pisa, Italy areas the decline in population size continues. Generally, the effects of climate change, lowering of water tables and habitat fragmentation Many ecological and environmental factors, such as air temperature (T), can act on body size are accepted explanations causing the decline. We additionally consider ‘conservation grazing’ and shape variation within a species. Bergmann’s rule, stating that size tends to increase in to be an important factor causing population declines. RAVON studied viviparous lizard colder climates, has been tested on chelonians with positive results. Testudo hermanni matches populations on lowland heathlands under different grazing regimes from 2010–2012. The this rule in Greece, Balkan Peninsula and Italy where however the rule was tested comparing grazing intensity in different plots is statistically negatively correlated to lizard densities, where- northern and southern populations a priori distinguished. To verify if size variation of Italian as sympatric sand lizard populations (Lacerta agilis) seem to be less affected. We additionally T. h. hermanni is related to climate we correlated linear carapace length with latitude, mean compared the data from this study to data from long-term population studies in the 1970’s annual T, maximum T of the warmest month, minimum T of the coldest one, using Spear- in the same areas. Population declines over 40 years ranged from 30 up to 100%. Even low man correlation test. Data derived from field sampling (7 populations) and literature (4) for grazing intensities lead to changes in vegetation coverage, microstructural changes in heath- a total of 610 adult tortoises (303 mm, 302 ff) from 11 localities in peninsular Italy and Sardinia. land vegetation and degradation of lizard microhabitats. Climatic data were obtained from WorldClim dataset at a resolution of about 1 km2. Interestingly size of both sexes positively correlated with Tmin that likely is a limiting factor for growth while it increased with decreasing Tmax, significantly in females (R=–0.603, P=0.037) but showing just a trend in males (R=–0.547, P=0.065). Male size positively cor- related with Tmean (R=0.651,P=0.029), females not (P>0.05). Correlations with latitude were not significant for both sexes (P>0.05), challenging the use of latitude as approximation of climatic conditions. Our results seem to confirm Bergmann’s rule in T. hermanni con- sidering the highest T of the year, which are crucial for factors like length of growing season, water and food availability. However, in warmer climates, as in Mediterranean area, environ- mental temperatures may be not be the key force shaping variability of ectothermic mor- phology and additional factors can influence mechanisms of phenotype selection and animal survival. A complete discussion should also involve age classes analysis because results could be influenced by different survival and recruitment capacity of populations.

100 101 MORPHOLOGY, ANATOMY, PHYSIOLOGY AND DEVELOPMENT DISEASES AND PARASITES

Relation between righting time and carapace shape in adult Analyzing the impact of blood parasites in selected lizard species from Hermann’s tortoises Portugal and Morocco

Jelka Crnobrnja-Isailovic´ 1,2, Dragana Stojadinovic´ 1, Durad Milosevic´ 1 Isabel Damas-Moreira 1,2, João P.M.C. Maia 1,2,3, Beatriz Tomé 1, Daniele Salvi 1, Veronica Gomes 1,4, Antigoni Kaliontzopoulou 1,5, Miguel A. Carretero 1, 1 Department of Biology and Ecology, Faculty of Sciences and Mathematics, University of Nisˇ, Visˇegradsˇka 33, D. James Harris 1, Ana Perera1 18000 Nisˇ, Serbia; [email protected] 2 Institute for Biological Research “S. Stankovic´”, University of Belgrade, Despota Stefana 142, 11000 Belgrade, Serbia 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, 4485-661 Vairão, Portugal; [email protected] Righting ability is important for survival of adult tortoises and it was suggested that particular 2 Departament of Biology, Faculty of Sciences, University of Porto, Portugal determinants of carapace shape could be an important factor regarding their ability to recover 3 Institut de Biologia Evolutiva (CSIC-UPF), Barcelona, Spain prone position. 4 Cátedra “Rui Nabeiro”, Biodiversidade, Centro de Investigação em Biodiversidade e Recursos Genéticos (CIBIO), We measured duration of righting in 303 adult Hermann’s tortoises at two different habitat Universidade de Évora, 7004-516 Évora, Portugal 5 Department of Ecology, Evolution, and Organismal Biology, (515) 294-0133, Iowa State University, Ames, IA, USA types (forest and vineyard) in the southeastern Serbia. During sampling period, 5 parameters of carapace morphology (straight carapace length—SCL, curvilinear carapace length—CCL, Numerous studies place parasites in a central role for studying populations’ ecology and mean carapace width—MCW, maximum carapace width—MAXCW and maximum cara- their dynamics. The phylum Apicomplexa is an interesting and diverse group of obligate uni- pace height—MAXSH) and body mass—BM were measured. cellular blood parasites, with a vast distribution, that can pose serious health risks both The sexual size and shape dimorphism of carapace morphology emerged up in both pop- for animals and humans. Studies conducted in lizards have been mainly focused on the ulations. It was not the case with the righting time (RT) where only the population effect was analysis of parasite prevalence and intensity, using Haemogregarines as a model since it is statistically significant (p<0.0001, eta squared=0.08). Moreover, RT was also controlled by the most common apicomplexan blood parasites in lacertids. However information about ambiental temperature (p=0.002), relative BM (p=0.002), and, except CCL, by all morpho- the impact of these parasites on the host is still scarce, which can be of major concern for metric traits describing carapace shape. However, all covariates were poor at explaning the conservation efforts. overall variation in RT with a weak effect size (eta squared less than 0.06 in all tests). Canonical Therefore our main goal is to assess what are the impacts of a Haemogregarine parasite on correlation analysis revealed moderate and significant relation between carapace shape (“form”) the lizard host. Particularly, we analysed the impact on the immune system. We used the sim- and RT (“function”) (canonical r=0.30, df=5, p<0.0001). plified PHA skin-testing technique, a standard tool for estimating host immune condition. The results suggest that general carapace shape slightly contributes to righting ability This condition is then related with parasite intensity and circulating blood cells, examined in adult Hermann’s tortoises. However, it seems that some other morphological and/or physi- under the microscope. Identity of parasites was confirmed using 18S rRNA gene sequences. ological variables or more specific components of shell structure are also important drivers Two assays were carried out: one performed in the field at three Moroccan sites, using two of righting time. sympatric lizard species (Podarcis vaucheri and Scelarcis perspicillata); the second experiment was a laboratory based comparison, with performance tests incorporated, using Podarcis bocagei from four sites in Portugal. We found all populations parasitized with overall high prevalence and intensity. For both assays no correlations were found between the PHA response, parasitemia levels or any other factor. We thus suggest that these parasites might have limited impact on lizard health and fitness. This conclusion can be of conservation significance and we discuss this in the light of different possible scenarios, particularly that these parasites might have been coevolving with the lizards for a long time, resulting in a insignificant pathogenicity for the host.

102 103 POPULATION GENETICS ECOLOGY

Various DNA sequence analyses of amphibian larvae in the semi-arid Environmental correlates of paedomorphosis and metamorphosis zone in palmate newts

Gad Degani Mathieu Denoël 1, G. Francesco Ficetola 2

School of Science and Technology, Tel Hai Academic College, Upper Galilee 12210, Israel; [email protected] 1 Laboratory of Fish and Amphibian Ethology, Behavioural Biology Unit, University of Liège, 22 Quai van Beneden, 4020 Liège, Belgium; [email protected] 2 Department of Environmental and Earth Sciences, University of Milano-Bicocca, 1 Piazza della Scienza, The molecular DNA variation among Salamandra infraimmaculata and Triturus vittatus lar- 20126 Milano, Italy vae populations, representing diverse breeding sites in Israel, was analyzed by cytochrome b fragment, control region, DNA polymerase chain reaction (RAPD PCR) and amplified frag- Paedomorphosis and metamorphosis are alternative developmental pathways expressed in ment length polymorphism (AFLP) methods. Although the molecular polymorphisms in populations of several caudate species. They result respectively in the development of indi- both mitochondrial and nuclear DNA are small, they reflect a sharp ecological separation viduals that mature in the larval stage (the paedomorphs) and of larvae that metamorphose between DNA variation of seasonal breeding sites and permanent water sources. These are into a terrestrial morph that acquire maturity later (the metamorphs). The ontogenetic presumably adaptive changes caused by natural selection. Low genetic (cyt b and 12S, and trajectories rely on environmental cues but their effects across both space and time have RAPD PCR) variation was revealed by sequences from specimens of Hyla savignyi, Bufo not been examined yet. To determine the relationships between environmental variables viridis, Rana bedriagae and Pelobates syriacus populations, respectively, at each location, ana- and morph expression, we sampled more than 150 populations of palmate newts (Lissotriton lyzed by Arlequin software. We found that both H. savignyi and B. viridis are adapted to more helveticus) during a 10-yr study and we used an information-theoretic approach to test the unpredictable breeding places than R. bedriagae and P. syriacus. likelihood of multiple ecological processes. All environmental processes influenced the re- lative abundance of both morphs. Paedomorphs were particularly advantaged in deep and oxygenated wetlands, without fish, and surrounded by vertical slopes. Higher forest and lower aquatic vegetation cover were also associated with a higher prevalence of paedomor- phosis over metamorphosis. Analyses of paedomorph variation along time confirm these con- clusions on the three variables that showed inter-annual variations (water depth, oxygen content, and aquatic vegetation). All together these results show that paedomorphosis and metamorphosis depend on multiple environmental factors and can evolve rapidly. This could explain the absence and extirpation of paedomorphosis in natural populations, particularly when affected by anthropogenic pressures. The degradation of habitats inhabited by paedo- morphs and the rarity of these populations make them at a high risk of disappearance.

104 105 PHYLOGEOGRAPHY AND PHYLOGENY CLIMATE CHANGE

Ain’t no mountain high enough, ain’t no valley low enough? Occupancy modelling for predicting the distribution of Attaining biogeographical barriers using Cordylus cordylus as Philoria loveridgei in the face of climate change template taxon Mariel Familiar Lopez 1, Greg Lollback 1, David Newell 2, Jean-Marc Hero 1 Genevieve Diedericks, Savel R. Daniels 1 Environmental Futures Centre, School of Environment, Griffith University, Gold Coast Campus, QLD 4222, Australia; [email protected] Evolutionary Genomics Group, Department of Botany and Zoology, Stellenbosch University, Private Bag X1, 2 School of Environment, Science and Engineering, Southern Cross University, Lismore Campus, NSW 2480, Matieland 7602, South Africa; [email protected] Australia

South Africa, more specifically the Cape Floristic Region, is a known floral and faunal hot- Climate change and disease have been strongly linked to enigmatic global amphibian de- spot with high levels of herpetofaunal endemism. Previous studies conducted on southern clines, particularly for species and populations at higher elevations, as it can increase tempera- African taxa illustrated that rupicolous species show marked genetic structuring associated ture and cause precipitation fluctuations that may act as stressors. Global warming is there- with habitat type. This study seeks to determine if this holds true for Cordylus cordylus, a rupi- fore predicted to have a wide range of effects on amphibian species, one being the alteration colous sit-and-wait foraging lizard. Samples were collected across the species range from 55 of distribution and abundance, with the general trend that species will shift towards the poles localities, yielding a sample size of 175 specimens. Four genes, two nuclear and two mitochon- and to higher elevations. For mountain-top endemic frogs, global warming may have devasta- drial, were amplified using the polymerase chain reaction (PCR). Standard tree methods were ting effects, as it would force these species beyond the limit of their distribution into extinction. employed to test for species monophyly, followed by population structure analyses and niche Niche modelling algorithms are powerful tools for modelling species distribution and have modelling. Our results confirm the species’ monophyly and highlight the presence of two dis- great potential in species conservation. However, field surveys based on occupancy models tinct clades. The divergence time estimates revealed a Late Miocene (6.01 Ma), Early Pliocene may be used to evaluate the predicted distribution models improving there potential. We (4.31 Ma) divergence for each of the two clades retrieved. Furthermore, the niche modelling present a modelling framework which combines a species distribution approach (present and shows that C. cordylus occupies a wide range of unfavourable habitats. This is a very un- future) for Philoria loveridgei using Maxent and verification using occupancy modelling. The characteristic phylogeographic pattern for a rupicolous vertebrate species, since all the southern highest number of Philoria frogs recorded in one transect was 18, with an occupancy estimate African rupicolous taxa that have been subjected to phylogeographic study demonstrated of 76.1%, describing high occupancy within the middle upper range of the predicted distri- marked genetic structuring at small spatial scales. We believe the ecological pliability and bution. Detection decreased as survey season progressed with an estimate range from 0.69 to non-specialist nature of C. cordylus presumably contributed to the observed phylogeographic 0.07. These results reinforce the species distribution model and confirm the restricted distri- pattern and have facilitated the absence of within clade differentiation. Moreover, we suggest bution of this species. The validated model was used to predict the likely impacts of climate that microclimatic variables, rather than physical geographic barriers influence the genetic change on the species distribution and results suggest it will reduce in area (~10%). The structuring of C. cordylus. Because reptile taxa are reliant on climatic variables to govern their method used in this study provides a new approach to verify species distribution models that distribution, this pattern may be more prominent in squamate taxa when compared to other is suitable for other species. Extinction risk is exacerbated for high elevation Philoria species as taxa, such as small mammals. they are confined to cooler climates that could become more suitable for the fungal disease chytridiomycosis with the increasing temperatures.

106 107 ECOLOGY ECOLOGY

Multi-species occupancy modeling of natural and anthropogenic An evaluation of the robustness of global amphibian range maps habitats by Mediterranean amphibians: grim prospects for conservation in irrigated farmland Gentile Francesco Ficetola 1, Carlo Rondinini 2, Anna Bonardi 1, Vineet Katariya 3, Emilio Padoa-Schioppa 1, Ariadne Angulo 3 Mário Ferreira, Pedro Beja 1 Department of Earth and Environmental Sciences, University of Milano-Bicocca, Piazza della Scienza 1, 20126 Milano Italy, [email protected] EDP Biodiversity Chair, CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do 2 Global Mammal Assessment Program, Department of Biology and Biotechnologies, Sapienza University of Rome, Porto, Campus Agrário de Vairão, 4485-661 Vairão, Portugal; [email protected] Viale dell’Università 32, 00185 Roma, Italy 3 IUCN Global Species Programme, Rue Mauverney 28, 1196 Gland, Switzerland In farmland landscapes, amphibians often breed in artificial water bodies, which may offset at least partly the loss of natural wetlands associated with agricultural intensification. It is Maps of species ranges are among the most used distribution data in biodiversity studies. As possible, however, that artificial habitats provide conditions for a minor proportion of the with any biological datum, range maps have some level of measurement error, but this error is regional species pool, benefiting just a few widespread generalists. We assessed these alter- rarely quantified. The aim of this study was assessing the error associated with the amphibian native views by documenting the decline of temporary ponds in a Mediterranean farmland range maps published by the IUCN’s Global Amphibian Assessment. The amphibian range landscape between 1991 and 2009, and by estimating the occupancy of natural (temporary maps were assessed against two datasets of species point localities: the Global Biodiversity ponds and streams) and artificial water bodies (farm ponds, irrigation channels and drainage Information Facility (GBIF), and a refined dataset including recently published, high quality ditches) by amphibians across the breeding season (February–June). We used hierarchical presence data. Range fit was measured as the proportion of presence records falling within Bayesian dynamic multi-species occupancy modeling to control for differences in detectability the IUCN range polygon(s) for each species. Using the high quality point data provided better across species, sampling occasions and habitat types. Over two decades the farmland landscape fit measures than using the raw GBIF data. Range fit was highly variable among continents, lost 56% of its temporary ponds, of which 89.3% were destroyed through agricultural activi- being highest for North American and European species (fit=84–94%), and lowest for Asian ties such as cultivation, conversion to permanent farm ponds, and drainage. The survival rate and South American species (fit=57–64%). At the global scale, 95% of amphibian point of ponds was lowest within an irrigated perimeter, and there was no positive effect of protection records were inside published maps, or within 31 km from the range edge. However, differ- by a natural park. Estimated species richness per site was at least twice as high in temporary ences among continents were striking, and more points were found far from range edges for ponds as in the other habitat types. From the ten amphibian species recorded, seven had the South American and Asian species. IUCN range maps represent the known distribution of highest occupancy in temporary ponds, and were absent or occurred rarely in artificial habi- most amphibians well; this study provides measures of accuracy that can be useful for future tats. Only a single generalist species was widespread in farm ponds and irrigation channels. research using amphibian maps as baseline data. Nevertheless, there is a need for greater in- The results suggest that artificial water bodies are unlikely to sustain populations of most vestment in the continuous update and improvement of maps, particularly in the mega- amphibian species in Mediterranean intensive farmland. Conservation efforts should be diverse areas of tropical Asia and South America. directed at protecting the temporary ponds still remaining within the farmed landscape, and at restoring or creating new temporary ponds where these have been lost during the last decades.

108 109 BEHAVIOUR PRACTICAL CONSERVATION

A non-invasive water-borne hormone assay shows that tadpole Detection of olm (Proteus anguinus) using environmental DNA in populations of Alytes obstetricans infected with Batrachochytrium Croatia dendrobatidis have elevated corticosterone levels Júlia Tünde Gál 1, Dusˇan Jelic´ 2, Judit Vörös 3 Caitlin R. Gabor 1, Matthew C. Fisher 2, Jaime Bosch 3 1 Department of Ecology, Faculty of Veterinary Science, Szent István University, Rottenbiller u. 50, 1078 Budapest, Hungary; [email protected] 1 Department of Biology, Texas State University, San Marcos, TX 78666, USA; [email protected] 2 Croatian Herpetological Society—HYLA, Prva Breznicka 5a, 10000 Zagreb, Croatia 2 Department of Infectious Disease Epidemiology, Imperial College London, Norfolk Place, London W2 1PG, 3 Department of Zoology, Hungarian Natural History Museum, Baross u. 13, 1088 Budapest, Hungary United Kingdom 3 Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain The olm (Proteus anguinus) is the only cave-dwelling vertebrate in Europe, inhabiting the karst The fungus Batrachochytrium dendrobatidis (Bd) causes the disease chytridiomycosis and is cave systems of the Dinaric Alps. It is a troglomorphic species, having a hidden way of living. associated with amphibian declines. Populations vary in their susceptibility to Bd infections Since it inhabits the interiors of caves, it is hardly seen on the surface and we are lacking a and this variation may be influenced by immunosuppression caused by chronic stress im- lot of information about its exact distribution. posed by environmental factors. Using a non-invasive water-borne hormone technique we Recent studies have shown that by sampling the environment, we can get information assessed stress levels (corticosterone) of Bd infected and uninfected free-living tadpole pop- about the species living there. To gather further data about the olm, a new, non-invasive ulations. We found that corticosterone release rates were higher in infected populations of monitoring method is urgently needed. Alytes obstetricans than in an uninfected populations. The relationship between corticosterone The aim of this study was to develop a method by which we can detect olm DNA from and the intensity of infection was significantly positively correlated in some populations. The cave waters. In the first phase, we took 30 ml samples from aquariums of the Zoo of Zagreb next step is to determine whether higher corticosterone levels make individuals more sus- and from Miljacka cave in June 2012. In the second phase 10, 5 and 2 l filtered samples were ceptible to Bd or if Bd infections drive the higher corticosterone levels. taken from Rupecica cave system in September 2012. We developed specific primers to am- plify short DNA sequences from the mitochondrial D-loop region (87 bp) to determine the presence of the species in different environments. The reliability of the results was tested by sequencing and the sequences were compared to the DNA extracted from olm tissue. With this method, first time applied eDNA technique on a cave-dwelling organism, we were able to detect P. anguinus DNA from one sample from Rupecica (filtered), one from Miljacka (not filtered) and one from aquarium of the Zoo of Zagreb (not filtered). However, further improvement of the PCR reactions and the sequencing technique is needed.

110 111 ECOLOGY POPULATION GENETICS

Annual activity dynamics of terrestrial amphibians in the monsoon Is expert knowledge adapted to build friction map? A landscape lowland forest in southern Vietnam genetic study of the slow worm (Anguis fragilis)

Eduard A. Galoyan 1,3, Anna B. Vassilieva2,3, Nikolay A. Poyarkov, Jr. 2,3 Céline Geiser 1,2, Nicolas Ray 2, Anthony Lehmann 2, Sylvain Ursenbacher 1

1 Zoological Museum of Moscow University, Bolshaya Nikitskaya Street 6, 125009 Moscow, Russia; [email protected] 1 Department of Environmental Sciences, Section of Conservation Biology, University of Basel, St. Johanns-Vorstadt 10, 2 Department of Vertebrate Zoology, Biological Faculty, Lomonosov Moscow State University, Leninskiye Gory, GSP–1, 4056 Basel, Switzerland; [email protected] 119991 Moscow, Russia 2 Laboratory of Spatial Predictions and Analyses in Complex Environments, Institute for Environmental Sciences, 3 Joint Russian–Vietnamese Tropical Research and Technological Center of the A.N. Severtsov Institute of Ecology and University of Geneva, 7 route de Drize, 1227 Carouge, Switzerland Evolution, South Branch, 3, Street 3/2, 10 District, Ho Chi Minh City, Vietnam Landscape genetics was developed to detect landscape elements shaping genetic population Amphibian activity is highly dependent on temperature and humidity. As far as in Asian structure, including the effects of fragmentation. Multifarious environmental variables can tropical monsoon forests temperature changes slightly during the year, we may suggest that influence gene flow in different ways and expert knowledge is frequently used to construct frog activity must be regulated mostly by precipitation amount; more animals and species friction maps. However, the extent of the migration and the movement of single individuals should be active during the rainy season (May–October) than during the dry season. To test are frequently unknown, especially for non-model species, and friction maps only based on this suggestion we studied amphibian activity during the whole annual cycle from August expert knowledge can be misleading. In this study, we used three different methods: isolation 2011 to July 2012 using registration of calling activity and pit-fall traps placed in seven study by distance (IBD), least-cost modelling and a strip-based approach to disentangle the human sites in Cat Tien National park. 2644 individuals of 17 species were found in the traps dur- implication in the fragmentation process in the slow worm (Anguis fragilis), as well as the spe- ing 1152 trap-days. The number of catches was highest during the rainy season, however, some cific landscape elements shaping the genetic structure in a highly anthropized 16 km2 area species (Occidozyga martensii, Limnonectes sp., Kalophrynus interlineatus) demonstrated high in Switzerland. Friction maps were constructed using expert opinion, but also based on the terrestrial activity during the first half of the dry season (November–January). Frog activity combination of all possible weightings for all landscape elements. The IBD indicated a sig- reduced greatly in the driest period, in February–March. Adults prevailed in traps in the nificant effect of geographic distance on genetic differentiation. Further approaches demon- beginning of rainy season and subadults in the end of this period. strated that highways and railways were the most important elements impeding the gene flow The number of calling species positively correlates with the reduction of precipitation in in this area. Surprisingly, we also found that agricultural areas and dense forests seemed to the period August–February. In contrast, the number of calling species grows explosively up be used as dispersal corridors. These results confirmed that the slow worm has relatively un- to a maximum of 20 registered species with the first heavy rains in April. However, some specific habitat requirements. Finally, we showed that our models based on expert knowledge species, such as tree-hole breeding Theloderma stellatum, keep on calling and reproducing performed poorly compared to cautious analysis of each variable. This study demonstrated even during dry season. that landscape genetic analyses should take expert knowledge with caution and exhaustive Finally, we distinguish two main ecological groups of terrestrial amphibians: species active analyses of each landscape element without a priori knowledge and different methods can be all-year round (O. martensii, Limnonectes sp., K. interlineatus), surviving the dry season in recommended. small water bodies or in the humid leaf litter, we define them as active patients; frogs active during the rainy season (Caluella guttulata, Kaloula pulchra, Micryletta erythropoda) with ex- plosive growth of terrestrial activity at the onset of heavy rains and surviving the dry period in shelters are defined as aestivates.

112 113 PRACTICAL CONSERVATION BEHAVIOUR

Newts in the city: research and conservation of amphibians in Lobau Tortoise’s shell—a blessing and a curse. How tortoises cope with (Vienna, Donau-Auen National Park) various obstacles

Günter Gollmann 1, Heimo Schedl 2, Daniel Philipp 1, Walter Reckendorfer 3, Ana Golubovic´ 1, Marko Andjelkovic´ 2, Dragan Arsovski 3, Sonja Djordjevic´ 1, Andrea Waringer-Löschenkohl 4 Vuk Ikovic´ 4, Ana Vujovic´ 5, Xavier Bonnet 6, Ljiljana Tomovic´ 1,2

1 University of Vienna, Department of Theoretical Biology, Althanstraße 14, 1090 Wien, Austria; 1 Faculty of Biology, University of Belgrade, Studentski trg 16, 11000 Belgrade, Serbia; [email protected] [email protected] 2 Institute for Biological Research “S. Stankovic´”, University of Belgrade, Bulevar Despota Stefana 142, 2 University of Natural Resources and Life Sciences, Institute of Integrative Nature Conservation Research, 11000 Belgrade, Serbia Gregor Mendel Straße 33, 1180 Wien, Austria 3 Macedonian Ecological Society—herpetology group, Bulevar “Kuzman Josifovski-Pitu” 28/3-7, 1000 Skopje, 3 WasserCluster Lunz—Biologische Station GmbH, Dr. Carl Kupelwieser Promenade 5, 3293 Lunz am See, Austria FYR of Macedonia 4 University of Vienna, Department of Limnology, Althanstraße 14, 1090 Wien, Austria 4 Municipality of Danilovgrad, Trg 9. Decembra, 81410 Danilovgrad, Montenegro 5 Department of Biology, Faculty of Natural Sciences and Mathematics, University of Montenegro, Lobau is part of a floodplain of the Danube River in Vienna (Austria). Due to river engi- Dzˇordzˇa Vasˇingtona bb, 81000 Podgorica, Montenegro 6 CEBC-CNRS, 79360 Villiers en Bois, France neering, hydrodynamics are constrained and vary greatly among different sections. In Lobau at least ten amphibian species occur. During 200 million years of evolutionary history, all chelonians are characterized by a well We synthesize results of recent surveys, highlighting the challenges for research and con- developed, protective shell. Conserved during major evolutionary transitions (e.g. return to servation, with a focus on the species of highest conservation priority. These investigations the sea), shell likely provides substantial selective advantages. Conversely, heavy and rigid were carried out at different spatial and temporal scales. From 1997 to 2007 abundance of shell poses a set of serious motion limitations, especially in terrestrial species during displace- Bombina bombina declined to about a tenth in a study area. At another site, only five males ments in rugged and complex environments. We examined how terrestrial tortoises over- were heard calling in 2007, but in 2008, a year with a higher water table, population size was come various obstacles in their natural habitat. We compared several Balkan populations of estimated as above 500 (based on mark-recapture analysis of 392 individually registered toads). Hermann’s tortoises by placing individuals (app. 1100) in three types of challenging positions Intensive surveys in 2009, a wet year, showed that Triturus dobrogicus occurred in 39 water to measure their performances. bodies throughout Lobau, in 23 of which evidence for reproduction (eggs or larvae) was found. Righting ability: males were more successful in righting themselves than females. Body size In a few locations where the species was recorded in the 1990s no newts were observed; influenced righting abilities only in females: smaller ones were more efficient. Individuals formerly stagnant pools had been converted into flowing channels inhabited by fish. with markedly domed carapace (increased instability) and shorter bridges (enlarging space for Due to hydrodynamics in this floodplain area, habitat availability and abundances may leg movements) were more successful to right themselves. differ markedly between years. Population studies spanning several years are urgently needed. Boldness to cross a steep step: animals from rugged environment were more prone to jump Reproduction of amphibians often is highest in temporary water bodies. Extreme dynamics from 50 cm high obstacle then ones from the flat habitat. Genders had similar success, may be disadvantageous, however, as quickly drying ephemeral pools become reproductive but adults showed more boldness to leap off the hedge than juveniles. sinks and major floods may bring fish to formerly fish-free waters. Disentangling themselves form thick vegetation: residents of Mediterranean macchia dis- Lobau is still home of viable populations of these endangered amphibian species. Never- played moderate pulling force and released themselves quickly by backwards movements. theless, long-term trends towards terrestrialization will ultimately lead to a decline in these Tortoises living in herbaceous habitats moved forward, deployed strong pulling force, ripping habitats. Therefore, restoration measures sustaining overall habitat dynamics and diversity are on the substrate and were far less successful. urgently needed but must take great care to avoid deterioration of amphibian breeding sites. Individuals from different habitats, sex and age classes exhibited divergent and apparently appropriate traits and strategies to overcome each locomotory challenge. We further spec- ulate that behavioural responses, as a response to morphological constraints, are shaped by experience. These results may have conservation value and assist in improving translocation strategies for endangered tortoise populations.

114 115 MONITORING PRACTICAL CONSERVATION

Difference in the geographic overlap of the potential distribution of Tracking Hungarian meadow vipers (Vipera ursinii rakosiensis) the green and ocellated lizards at continental and regional extents in following reintroduction the Mediterranean Basin Bálint Halpern 1, Endre Sós 2, Chris Walzer 3, Tamás Péchy 1 M. X. González Mantilla 1, A. G. Toxopeus 1, T. Groen 1, A. K. Skidmore 1, 1 P. Lymberakis 2 MME BirdLife Hungary, Költo˝ u. 21, 1121 Budapest, Hungary; [email protected] 2 Budapest Zoo & Botanical Garden, Állatkerti krt. 6–12, 1146 Budapest, Hungary 3 Research Institute of Wildlife Ecology, University of Veterinary Medicine Vienna, Savoyenstraße 1, 1160 Wien, 1 Department of Natural Resources, ITC, University of Twente, Hengelosestraat 99, 7514AE Enschede, Austria The Netherlands; [email protected] 2 Natural History Museum Crete, University of Crete, Knosou Ave, P.O. Box 2208, 71409, Iraklion, Crete, Greece In order to stop the decline of Hungarian meadow viper (Vipera ursinii rakosiensis), MME BirdLife Hungary together with National Parks started a complex conservation program in The differences in the potential geographic overlap areas resulting from models fitted at con- 2004, supported by LIFE-Nature fund. tinental and at regional scale for seven related lizard species in the Mediterranean Basin have Hungarian Meadow Viper Conservation and Breeding Centre was started with 16 adult been investigated. The green lizards (Lacerta agilis, L. bilineata–viridis, L. trilineata, and L. individuals, collected from 6 different populations. By 2012 number of vipers bred reached schreiberi) and the ocellated lizards (Timon lepidus, T. tangitanus, and T. pater) were included nearly 1400. as target species. First reintroduction took place in March 2010, releasing 30 snakes into a reconstructed habi- The potential geographic overlap areas were derived from modelling the potential spatial tat in Kiskunság National Park. Additional 70 vipers were released there in February 2011, distribution of the target species at continental extent using only climate predictor variables. and further 42 snakes in July 2012. Snakes were released by removing them together with The compilation of distribution sample points for the species mentioned above were derived their artificial burrows. During regular monitoring we checked these burrows by using pipe- from various sources (existing atlases and other sources from web). Based on these results, four camera. zones were selected to analyse the differences with the outcomes modelled at regional extent. In 2010 we recorded vipers 54 times. Four of the observed females were gravid. Despite fur- In the selected zones, the potential geographic overlap areas were determined for pairs of ther releases we only spotted 40 snakes in 2011, but newborn vipers were seen. Altogether species based on models fitted at regional extent using only climate predictor variables and 15 individuals were identified by photos taken during these events. We found signs of pre- another adding variables related to land cover, topography and NDVI. By comparing the per- dation two times. formance of the models fitted, six out of eight of the models do not significantly improve the We installed camera traps in 2012 and recorded frequent presence of possible predators, area under the curve (AUC) values by adding variables related to land cover, topography and such as foxes, badgers, common buzzards and wild boars. NDVI. In order to develop a remote tracking method, pre-programmed radio-tags with a detection The resulting potential geographic overlap areas modelled at continental extent were zoomed range of 100–150 m were implanted surgically into the abdomen of vipers. These tags also in on the selected zones and compared with the results at regional extent keeping the same operate as temperature loggers, recording data every five minutes for a year-long operation spatial resolution. Based on this comparison, the potential geographic overlap areas do not period. After successful testing during 2010 and 2011, we released 6 tagged individuals in July significantly change depending on the type of predictor variables use to model the potential 2012. All 6 snakes survived until November, even though wild boars dig large part of the site. spatial distribution of the species at regional extent. On the other hand, the potential geo- We detected regular movements within the area, with tendencies to higher parts later in the graphic overlap areas significantly changed depending on the extent at which the potential season. On sunny spells of mid-November and late December, after emerging for basking, distribution models of the species were fitted. The potential geographic overlap areas derived 3 and 2 snakes disappeared, very likely due to predation by raptors. One tag was found on from models fitted at regional extent reduced considerably in comparison with the potential the site and logged temperature data proved our theory. geographic overlap areas from models fitted at continental extent. Therefore, results concerning modelling the potential distribution of species should be interpreted with caution and put into the right context.

116 117 SYSTEMATICS PHYLOGEOGRAPHY AND PHYLOGENY

Cryptic diversity in Calotes mystaceus: an integrative approach Phylogeny of the genus Salamandra to unravel a putative species complex J.S. Hauswaldt 1, D. Eikelmann 1, S. Steinfartz 2, M. Vences 1 Timo Hartmann 1, Peter Geissler 1, Flora Ihlow 1, Nikolay A. Poyarkov, Jr. 2, 1 Dennis Rödder 1, Andreas Schmitz 3, Wolfgang Böhme 1 TU Braunschweig, Mendelssohnstraße 4, 38106 Braunschweig, Germany; [email protected] 2 University of Bielefeld, Morgenbreede 45, 33615 Bielefeld, Germany

1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; [email protected] The genus Salamandra is widely distributed in the western Palearctic and comprises six main 2 Lomonosov Moscow State University, Biological Faculty, Department of Vertebrate Zoology, Leninskiye Gory, GSP-1, lineages, S. salamandra, S. algira, S. infraimmaculata, S. corsica, S. atra and S. lanzai, of which 119991 Moscow, Russia the first group has the widest distribution across Europe and at least 14 subspecies of it have 3 Muséum d’Histoire Naturelle, Department of Herpetology and Ichthyology, C. P. 6434, 1211 Geneva 6, Switzerland been described. Salamandra atra and S. lanzai are completely black, whereas the others show different amounts of yellow coloration. While the monophyly of each of these major groups Calotes mystaceus Duméril & Bibron, 1837 is regarded as being a widespread and relatively has been clearly demonstrated, the relationship among them has not yet been resolved. While common species, which occurs from north-eastern India in the West throughout Southeast previous phylogenetic analyses were based on mtDNA only, we have used sequence data from Asia to the South China Sea in the East, to the Isthmus of Kra in the South and up to 24°N four mitochondrial and ten nuclear gene fragments and included representatives of all species in the North. To assess variation within this agamid species, an extensive dataset including and most subspecies to further address this issue. Preliminary Bayesian analyses of the com- 23 mensural and 11 metric characters of over 90 specimens was obtained covering the whole bined dataset confirm the monophyly of the clade composed of the completely black S. atra area of distribution. Our dataset was analyzed by both classical multivariate analyzes (PCA) and the yellow-blotched S. corsica. Contrary to previous studies, our results show that this and spatial multivariate approaches. In addition, a selective set of specimens was sampled and group is the sister taxon to the clade containing all other species of Salamandra, with the analyzed genetically using the fast evolving cytochrome oxidase I (COI) to discover even small other completely black taxon, S. lanzai, at the basal position of that clade. While previous genetic distances. By our integrative approach, we were able to dismantle Calotes mystaceus studies showed—if any—only weak support for a monophyletic clade composed of S. algira as being composed of more than one species, by discovering a surprisingly high genetic and and S. salamandra, our results clearly support this relationship. Within the fire salamanders, morphological divergence between populations. S. s. longirostris is clearly placed at the basal position. We are currently conducting divergence dating of the taxa and will discuss the species and subspecies level relationships in light of these results, the geographic distribution and life-history traits.

118 119 DISEASES AND PARASITES DISEASES AND PARASITES

Wetland characteristics influence disease risk for a threatened Species composition and chytridiomycosis infection of the Pelophylax amphibian esculentus complex (Anura: Ranidae) in the Hortobágy National Park, Hungary Geoffrey W. Heard 1, Michael P. Scroggie 2, Nick Clemann 2, David S.L. Ramsey 2 Dávid Herczeg 1, Judit Vörös 2 1 School of Botany, University of Melbourne, Melbourne, Victoria 3010, Australia; [email protected] 2 Arthur Rylah Institute for Environmental Research, Department of Sustainability and Environment, P.O. Box 137, 1 University of Debrecen, Department of Evolutionary Zoology and Human Biology, Egyetem tér 1, 4032 Debrecen, Heidelberg, Victoria 3084, Australia Hungary; [email protected] 2 Hungarian Natural History Museum, Baross utca 13, 1088 Budapest, Hungary Identifying drivers of the probability and intensity of infections is crucial for understanding the epidemiology of wildlife diseases, and for managing their impact on threatened species. Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis (Bd), is an Chytridiomycosis, caused by the fungal pathogen Batrachochytrium dendrobatidis, has deci- emerging infectious disease implicated in declines of amphibian populations around the mated populations of some amphibians. However, recent studies have identified important globe. In Hungary the presence of the chytrid fungus from elevated habitats was documented risk factors for the disease, related to the pathogens’ physiological tolerances. In this study, before but little is known about the distribution in the lowlands. The Western Palearctic we identified several intrinsic and extrinsic drivers of the probability and intensity of chytrid water frogs of the genus Pelophylax include a series of about 12 morphologically similar species infections for the threatened growling grass frog (Litoria raniformis) in south-eastern Australia, in Europe. The Pelophylax species complex consists three species in Hungary: P. ridibundus, and used mark-recapture to estimate the effect of chytrid infections on the probability of sur- P. lessonae and the hybrid P. kl. esculentus. Our sampling area situated in the Hortobágy vival of these frogs. Water temperature and salinity had negative effects on both the proba- National Park and we collected 29 Pelophylax spp. from two localities associated with flowing bility and intensity of chytrid infections. We coupled models of the infection process with a water and 30 specimens from the area of fish ponds and from there additionaly non-invasive model of the effect of chytrid infections on the probability of survival to assess variation in samples (19) becoming from skin swabbing. We used newly developed primers for distin- the impact of chytridiomycosis between wetlands with differing temperature and salinity guishing among the water frog species, and real-time PCR for detecting the Bd zoospores. profiles. Our results suggest that warm, saline wetlands may be refuges from chytridiomycosis Based on our molecular taxonomy findings the species composition of the investigated pop- for L. raniformis, and should be priorities for protection. Our results also suggest that ulations of water frogs consists the P. ridibundus and the hybrid P. kl. esculentus. Our results management actions that increase water temperature (e. g. reducing canopy shading) and clearly showed that the population of water frogs connected with flowing water were infected salinity (e. g. complementing inflows with groundwater) could be trialled to reduce the im- with Bd, while the population inhabiting the fish ponds were uninfected. The prevalence was pacts of chytridiomycosis on this species. This study highlights the value of research on en- 18.9% for the whole population together, while 51.7% for the frogs from the flowing water. vironmental factors that influence chytridiomycosis, particularly those factors that can be Prevalence was 37.1% for P. ridibundus and 8.3% for P. kl esculentus. The mean intensity was manipulated in the field. 10.13 GE (genome equivalent zoospore quantity).

120 121 MONITORING CLIMATE CHANGE

The use of environmental DNA (eDNA) to monitor amphibians Amphibian disease triangle: investigating interactions between disease, physiology and climate change Jelger Herder 1, Wilbert Bosman 1, Tony Dejean 2, Alice Valentini 2 Jean-Marc Hero, Edward Narayan 1 RAVON, Toernooiveld 1, 6525ED Nijmegen, The Netherlands; [email protected] 2 SPYGEN, 12 allée du Lac de Garde, Bâtiment House Boat n°7, Savoie Technolac B. P. 274, Environmental Futures Centre, School of Environment, Griffith University, Gold Coast Campus, QLD 4222, 73375 Le Bourget du Lac, France Australia; [email protected]

Analysis of environmental DNA (eDNA) is a new approach for monitoring freshwater bio- Amphibian declines have occurred simultaneously around the world, and amphibians are the diversity. The method is based on the limited persistence of the DNA left behind by species most threatened vertebrate group in what is described as the 6th major extinction event in in their environment. This environmental DNA can be detected in water samples, there- history. Mass die-offs are associated with an emerging pathogenic fungus that causes the dis- by confirming a species’ presence. In this presentation we will briefly explain the method ease chytridiomycosis; however there is evidence that climate change could be the ultimate and show the potential of this method on the bases of some case studies carried out in the cause. The key elements of these two hypotheses have not been investigated due to the com- Netherlands and France. In 2012 we investigated 23 historical populations of the common plex interactions between frog physiology (thermal biology, stress physiology, and antifungal spadefoot (Pelobates fuscus) in the Netherlands with the use of eDNA. It concerned locations agents on the skin of amphibians), disease ecology and host ecology. Climate change threatens were the species was thought to have gone extinct, based on traditional monitoring. With the amphibians restricted to mountain tops, as the cooler environmental conditions at high ele- use of eDNA we found that in six locations common spadefoots were still present. The out- vation are optimal for the growth and development of mountain-top endemic frog species, come meant a 15% increase in the number of known populations in the Netherlands. In hence they are vulnerable to increasing temperatures. Cooler montane environments are also a study on the crested newt (Triturus cristatus) in nine ponds we found that detection with optimal for the growth of the pathogenic fungus Bd. Here we present the first physiological eDNA was higher (five ponds) than detection with traditional methods like dip nets and evidence suggesting (1) frogs with high intensity of Bd have higher baseline stress hormone amphibian traps (two ponds). Furthermore we show the results of studies on the invasive levels, and (2) frog populations at higher elevation have increased stress levels (elevated base- American bullfrog (Lithobates catesbeianus). In the Netherlands we detected bullfrogs in one line corticosterone levels in urine) compared to their lowland counterparts. Prevalence of Bd out of 16 investigated locations with the use of eDNA, while the species was thought to be zoospores from frog skin swabs was quantified using a real-time quantitative PCR technique. eradicated. In France a comparison between traditional methods (searching for eggs, adults Individual male frogs that were identified as positive for Bd infection had significantly higher and larvae) and eDNA was made. American bullfrogs were detected in 38 out of 49 ponds baseline urinary corticosterone concentrations in comparison to Bd negative male frogs. Base- with the use of eDNA while traditional methods only found the species in seven ponds. Due line urinary corticosterone concentrations were also significantly higher at high altitude sites to the higher chance of detection the eDNA approach offers a powerful tool in early warning (P<0.001). These results suggest that frogs at higher elevation are stressed and hence more systems for invasive species. Finally, we take a look into the future by (i) describing eDNA susceptible to the impacts of Bd than their lowland counterparts. So which factor is respon- metabarcoding by which a list of species is generated from an environmental sample and sible for the extinction of frogs at high elevation and who came first—the stress or the disease? (ii) suggesting areas where more research is needed.

122 123 BEHAVIOUR BEHAVIOUR

From simple field observations to a model species—confessions of Antipredatory behavior of Spanish terrapins depends on reproductive a taxon-oriented field biologist state and individual conspicuousness

Walter Hödl Alex Ibáñez, Pilar López, José Martín

Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; Dpto. de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, C. S. I. C., José Gutiérrez Abascal 2, [email protected] 28006 Madrid, Spain; [email protected]

An unexpected rapid phonotactic approach of a male Allobates (=Phyllobates= Dendrobates= Behavioral responses to predation risk are critical to survive but, because antipredator be- Epipedobates) femoralis to a control replay of its recording in the Yubineto region, northern havior is costly, animals should flexibly modulate their responses depending on threat level Peru, in 1978 started my curiosity-driven research on the bioacoustics of this species. Due to as well as on their own individual characteristics. Turtles have morphological structures its stereotypic phonotactic response to playbacks of conspecific and a large variety of synthetic (shell) that may use as a refuge for partial protection but hiding can impose very high costs. calls I introduced A. femoralis as a “handy fellow” to the herpetological and bioacoustic com- Thus, after an unsuccessful predator attack, turtles should optimize the trade off between munity at various scientific meetings. Rapidly, “my” handy fellow became the main focus remaining hiding into their shells and active escape to safer habitat refuges. Here, we exam- of an international research team forming a cohesive group out of a diverse collection of ined how reproductive state, coloration and body size influenced antipredator behavior of individual scientists. Important inputs from Adolfo Amézquita (evolutionary aspects), Peter Spanish terrapins under different predation risk levels. We measured the time spent hiding Narins (robotics and sensory physiology), Albertina Lima and Pedro Ivo Simões (biogeog- into their own shells (i. e., appearance times) by turtles before escaping in response to simu- raphy) as well as Eva Ringler and Max Ringler (population biology) clearly show that lated predatory attacks of different risk level. Results showed that gravid females spent more taxon-centered research can be a self-accelerating process. The valuable integration of various time hiding into their shells after a predator attack in comparison with non-gravid females aspects have led to more fruitful insights than would have been possible with a pure hypo- and males, suggesting that a detriment in body condition and/or thermoregulation require- thesis-based approach, which sometimes may narrow the angle of view on a given research ments constrain an active escape. Similarly, hiding response was size-dependent with larger question. In 30 years of trying to understand a living organism in the field I have gained turtles having longer appearance times. Conspicuousness of limb coloration was important increasing confidence in my scientific approach to let organisms rather than hypotheses lead for appearance times of males, but not for females. Thus, males with brighter coloration of me to the important questions. And last but not least: taxon-centered research is simply more the limb stripes had longer appearance times than duller males, suggesting that a greater visual fun! conspicuity increased risk. However, gravidity, coloration and size had different relative im- portance in hiding behavior depending on whether turtles were overturned or placed in prone position after the attack, suggesting that the main factors influencing boldness depend on the risk level assessed by turtles.

124 125 CLIMATE CHANGE PHYLOGEOGRAPHY AND PHYLOGENY

On the brink of extinction? How climate change may affect global The Vipera berus group: phylogeny, reticulate evolution and chelonian species richness and distribution species borders

Flora Ihlow 1, Johannes Dambach 1, Jan O. Engler 1,2, Morris Flecks 1, Timo Hartmann 1, Ulrich Joger 1, Oleksandr Zinenko 2 Sven Nekum 1, Hossein Rajaei 1, Dennis Rödder 1 1 State Natural History Museum, 38106 Braunschweig, Germany; [email protected] 2 1 Herpetological Department, Zoologisches Forschungsmuseum Alexander Koenig (ZFMK), Adenaueralle 160, Museum of Nature at V.N. Karazin University, 61022 Kharkiv, Ukraine 53113 Bonn, Germany; [email protected] 2 Biogeography Department, Trier University, Universitätsring 15, 54286 Trier, Germany Vipera berus has the widest geographical distribution of all snakes. Its tolerance to cool tem- peratures makes it exceptional among other European snakes and it may have had glacial Anthropogenic global climate change has already led to alterations in biodiversity patterns by refugia North of the Alpine–Black Sea–Caucasus belt. directly and indirectly affecting species distributions. It has been suggested that poikilothermic Molecular studies (both mitochondrial and nuclear) revealed that southern isolates such as animals, including reptiles, will be particularly affected by global change and large-scale rep- V. b. bosniensis, V. (b.) nikolskii, V. (b.) barani and an undescribed Alpine form are differenti- tile declines have already been observed. Currently, half of the world’s freshwater turtles and ated to a higher degree than even the Far Eastern V. b. sachalinensis. tortoises are considered threatened with extinction, and climate change may exacerbate these Heterozygous individuals carrying alleles of both V. berus and V. renardi were found in declines. In this study, we assess how global chelonian species richness will change in the near contact zones of both species. Such cases raise the question of secondary introgression and future. We use species distribution models developed under current climate conditions for 78% reticulate evolution in these snakes. of all extant species and project them onto different Intergovernmental Panel on Climate Change (IPCC) scenarios for 2080. We detect a strong dependence of temperature shaping most species ranges, which coincide with their general temperature-related physiological traits (i. e., temperature-dependent sex determination). Furthermore, the extent and distribution of the current bioclimatic niches of most chelonians may change remarkably in the near future, likely leading to a substantial decrease of local species abundance and ultimately a reduction in species richness. Future climatic changes may cause the ranges of 86% of the species to contract, and of these ranges, nearly 12% are predicted to be situated completely outside their currently realized niches. Hence, the interplay of increasing habitat fragmenta- tion and loss due to climatic stress may result in a serious threat for several chelonian species.

126 127 ECOLOGY PHYLOGEOGRAPHY AND PHYLOGENY

Influence of pesticides on the development of eggs of agile frog, Phylogeny and phylogeography of the Syrian spadefoot toad Rana dalmatina (Pelobates syriacus)

Olga Jovanovic´, Zeljka Loncaric´, Branimir Kutuzovic´ Hackenberger Sarah Kieren 1, Angelica Crottini 2, Philipp de Pous 3, Michael Veith 1

1 Department of Biogeography, Trier University, Universitätsring 15, 54296 Trier, Germany; [email protected] Department of Biology, University Josip Juraj Strossmayer, 31000 Osijek, Croatia; [email protected] 2 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal Amphibians, as the most threatened group of vertebrates, are very susceptible to various envi- 3 Institute of Evolutionary Biology (CSIC-UPF), Passeig Maritim de la Barceloneta 37–49, 08003 Barcelona, Spain ronmental influences. One of the factors surrounding them is the water quality and its pol- lution. Although some areas in Europe have intensive agriculture, the influence of various The phylogenetic relationships within the genus Pelobates, namely within Pelobates syriacus, are pesticides on amphibians is still very poorly studied in general, especially in Europe. In our still not sufficiently understood. There exist several different hypotheses on the systematics research, we examined the influence of three widely used pesticides dimethoate, pirimiphos- of the subspecies of Pelobates syriacus. Currently, four subspecies are recognized: P. s. syriacus, methyl and glyphosate on the development dynamic of eggs of agile frog, Rana dalmatina. P. s. transcaucasicus, P. s. balcanicus and P. s. boettgeri. Previous molecular analyses questioned Influence of each pesticide was examined in five different concentrations, namely the lowest the status of P. syriacus as a monophylum. We here used DNA sequence analyses on 519 bp was ten times lower than environmentally relevant, second was environmentally relevant, third of the mitochondrial 16S rRNA gene to resolve the intraspecific phylogenetic relationships of was ten times higher, fourth was 100 times and fifth was 1000 times higher than environmen- P. syriacus. Three different tree building methods were applied (maximum likelihood, neigh- tally relevant. Experiment was carried out on 30 eggs in each of three replicates. We measured bor-joining and Bayesian inference). Phylogeographic analyses and molecular clock dating time needed to hatching and the hatching rate/mortality. In the highest two concentrations were performed under a Bayesian framework. We used the split between P. cultripes and of all three pesticides, the eggs did not develop into tadpoles, while in the concentration ten P. varaldii and intraspecific separations within Scaphiopus and Spea as calibration points. times higher than the environmentally relevant, the hatching rate was significantly lower in In conclusion, P. syriacus turned out to be paraphyletic, with two major evolutionary lineages: glyphosate then in other two pesticides. The lowest two concentrations of all three pesticides a western lineage representing the subspecies P. s. balcanicus, and an eastern lineage with the did not show to affect the development of eggs. subspecies P. s. transcaucasicus and P. s. syriacus. We suggest a Miocene separation of the eastern lineage from the rest of the genus Pelobates in the northwest of the current distribution range. Pelobates s. boettgeri does not represent a valid subspecies.

128 129 MORPHOLOGY, ANATOMY, PHYSIOLOGY AND DEVELOPMENT PHYLOGEOGRAPHY AND PHYLOGENY

Sex colour and stress: investigating the role of stress and reproductive Mitochondrial phylogeography of grass snakes (Natrix natrix, hormones in amphibian colour N. megalocephala) conflicts with taxonomy and reveals an additional unexpected contact zone in Central Europe Christina Kindermann 1, Edward J. Narayan 1, Clyde H.Wild 2, Jean-Marc Hero 1 Carolin Kindler 1, Wolfgang Böhme 2, Claudia Corti 3, Václav Gvozˇdík 4, 1 Environmental Futures Centre, School of Environment, Griffith University, Gold Coast Campus, QLD 4222, Daniel Jablonski 5, David Jandzˇik5, Margarita Metallinou 6, Pavel Siroky´ 7, Uwe Fritz 1 Australia; [email protected] 2 Griffith School of Environment, Science, Environment, Engineering and Technology Group, Griffith University, Gold Coast Campus, QLD 4222, Australia 1 Senckenberg Natural History Collections Dresden—Museum of Zoology, Koenigsbruecker Landstraße 159, 01109 Dresden, Germany; [email protected] 2 Zoologisches Forschungsinstitut und Museum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany Some animals have the capacity to change their colour, sometimes dramatically. There are 3 Museo di Storia Naturale dell’Università di Firenze, Sezione di Zoologia “La Specola”, Via Romana 17, 50125 Firenze, two main questions that are asked when investigating such colour change: (1) how does this Italy animal change colour; and (2) why does it change colour? Understanding how colour change 4 Department of Zoology, National Museum, Cirkusová 1740, 19300 Prague, Czech Republic occurs will often help determine why. Here we demonstrate dynamic colour change in an 5 Department of Zoology, Faculty of Natural Sciences, Comenius University Bratislava, Mlynská dolina B-1, anuran and investigate regulatory mechanisms of how it changes colour. Litoria wilcoxii 84215 Bratislava, Slovakia 6 Institut de Biologia Evolutiva, Animal Phylogeny and Systematics, Passeig Marítim de la Barceloneta 37–49, rapidly changes from brown to yellow during amplexus; we show this by comparing dorsal 08003 Barcelona, Spain colour of unpaired and amplecting males. Colour change involves the movement of pigments 7 Department of Biology and Wildlife Diseases, Faculty of Veterinary Hygiene and Ecology, University of Veterinary and in chromatophores activated by hormones. We investigated whether the corticotropin (adreno- Pharmaceutical Sciences, Palackého 1/3, 61242 Brno, Czech Republic corticotrophic hormone [ACTH]) and the neurohormone (epinephrine) or the male repro- ductive hormone (testosterone) were triggering colour change. We injected males with epi- Grass snakes (Natrix natrix) represent one of the most widely distributed snake species of the nephrine (n=5), ACTH (n=6), testosterone (n=5), and saline (n=5) and sesame oil (n=5) as Palaearctic region. Within N. natrix, up to 14 distinct subspecies are regarded as valid. In controls. Colour was measured using digital photography from the images the Red Green addition, some authors recognize big-headed grass snakes from western Transcaucasia as a and Blue (RGB) colour values were calculated, and the major correlation axis computed by distinct species, N. megalocephala. Based on phylogenetic analyses of a 1984-bp-long align- a principal components analysis (PCA). Epinephrine injected frogs underwent a significant ment of mtDNA sequences (cyt b, ND4 + tRNAs) of 410 grass snakes, a nearly range-wide colour change to bright yellow within 10 minutes compared to ACTH and control frogs which phylogeography is presented for both species. Within N. natrix, 16 terminal mitochondrial did not change colour. Testosterone treated males underwent a much slower (1 hour) and less clades were identified, most of which conflict with morphologically defined subspecies. yellow change. These results suggest that even though the hypothalamo-pituitary gonadal These 16 clades correspond to three more inclusive clades from (i) the Iberian Peninsula (HPG) axis is important for the expression of sexual morphological traits in male frogs, it is plus North Africa, (ii) East Europe and Asia and (iii) West Europe including Corso-Sardinia, the stress-axis, particularly neurotransmitters, that mediates the link between physiological the Apennine Peninsula and Sicily. Hypotheses regarding glacial refugia and postglacial range and morphological traits (rapid color change) The results have opened up opportunities for expansions are presented. Refugia were most likely located in each of the southern European future research to unravel the functions of physiological systems in amphibian colour change peninsulas, Corso-Sardinia, northern Africa, Anatolia and the neighbouring Near and Middle in breeding. East, where the greatest extant genetic diversity occurs. Holocene range expansions led to the colonization of more northerly regions and the formation of secondary contact zones. Western Europe was invaded from a refuge within southern France, while Central Europe was reached by two distinct range expansions from the Balkan Peninsula. In Central Europe, there are two contact zones of three distinct mitochondrial clades, one of these contact zones was theretofore completely unknown. Another contact zone is hypothesized for Eastern Europe, which was colonized, like northwestern Asia, from the Caucasus Region. Further contact zones were identified for southern Italy, the Balkans and Transcaucasia. In agreement with previous studies using morphological characters and allozymes, there is no evidence for the distinc- tiveness of N. megalocephala. Therefore, N. megalocephala is synonymized with N. natrix.

130 131 MORPHOLOGY, ANATOMY, PHYSIOLOGY AND DEVELOPMENT ECOLOGY

Relative contribution of dietary carotenoids and vitamin E to Assessing the costs of plasticity in chemical defenses in common toad visual and chemical sexual signals of male Schreiber’s green lizards (Bufo bufo) tadpoles (Lacerta schreiberi): an experimental test Anikó Kurali, Katalin Pásztor, Zoltán Tóth Renata Kopena, Pilar López, José Martín “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of Sciences, Herman Ottó Street 15, 1022 Budapest, Hungary; [email protected] Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain; [email protected] Various organisms can exhibit phenotypic plasticity, i. e. change their phenotype according to Carotenoid-based sexual ornaments are widespread, but the role of carotenoids as honest sig- the surrounding environmental conditions; however, the expression of induced phenotypes is nallers of the trade-off between coloration and antioxidant protection remains controversial. predicted to be costly. Chemical defence is thought to be a common and diverse phenomenon It was suggested that the role of carotenoids might not be as antioxidants per se, but that in amphibians, but very little is known about the plasticity and fitness costs related to this colourful carotenoids would just reflect the actual contents of nonpigmentary antioxidants, type of defensive response. Our aim was to assess the probable cost of toxin production as a such as melatonin or vitamin E. We experimentally fed male lizards Lacerta schreiberi (n=52) function of the frequency of depletion in common toad (Bufo bufo) tadpoles. For inducing with supplementary carotenoids or vitamin E alone, or carotenoids plus vitamin E together, toxin release in tadpoles in vivo, we used either electric or hormonal stimulation (norepi- and had a non-supplemented control group. We examined the effects of this dietary supple- nephrine). Tadpoles were raised in presence or absence of predator cues and exposed to low mentation on sexual coloration and the chemical profile of femoral gland secretions of lizards. or high food levels in a full factorial design, using randomized blocks. To assess potential costs Results showed that different sexual traits were differentially affected. Thus, both carotenoids of toxin production, we measured survival, time until metamorphosis and body mass at alone and vitamin E alone increased brightness of throat coloration respect to control lizards, metamorphosis. We found that the frequency of toxin depletion had a significant effect on but brightness was greater in lizards supplemented with vitamin E plus carotenoids together. all investigated traits on those tadpoles that were raised together with predator cues: individ- Also, all supplemented lizards had more saturated “yellowish” chests than control lizards, but uals that were forced to release toxins every five days had lower body mass, decreased survival lizards supplemented with vitamin E had more saturated and yellowish chest coloration. In and reached metamorphosis later, than those forced to release toxins only once. Low amount contrast, the supplementation did not affect the dorsal green coloration. Femoral secretions of food also had a negative effect on survival in all treatment groups; in addition, the type of also varied between treatments; lizards supplemented with vitamin E alone or combined with stimulation itself also influenced costs. Our results suggest that there are considerable fitness carotenoids had higher proportion of vitamin E in secretions than lizards supplemented with costs related to toxin production during the larval period in common toads, and may con- carotenoids alone and control lizards. Finally, lizards supplemented with vitamin E had tribute to a better understanding of the evolution and the adaptive value of chemical defences greater PHA-induced immune responses, while the supplementation of carotenoids did not in general. have any effect. Our results support that other nonpigmentary antioxidants, such as vitamin E, are needed in addition to carotenoids to increase the expression of sexual coloration of L. schreiberi lizards. In contrast, an increase in dietary nonpigmentary antioxidants may be directly reflected in the chemical signals.

132 133 PRACTICAL CONSERVATION MORPHOLOGY, ANATOMY, PHYSIOLOGY AND DEVELOPMENT

Distribution, habitat requirements and conservation of the Morphological insights into pulmonary evolution in reptiles Aesculapian snake Zamenis longissimus in south-eastern Poland Markus Lambertz, Steven F. Perry K. Kurek 1, W. Król 1, K. Najberek 1, S. Bury 1,2, B. Najbar 3, R. Babiasz 1, G. Bas 1, M. Potoczek 1, A. Ziecik 1, A. M. Cmiel 1, P. Wierzbanowski 1 Institut für Zoologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Poppelsdorfer Schloss, 53115 Bonn, Germany; [email protected]

1 Institute of Nature Conservation Polish Academy of Sciences, al. Mickiewicza 33, 31-120 Cracow, Poland, [email protected] The so-called reptiles were the first amniotes that achieved full terrestriality as they became 2 Institute of Environmental Sciences, Jagiellonian University, Gronostajowa 7, 30-387 Cracow, Poland independent of aquatic habitats for reproduction. This involved the evolution of numerous 3 University of Zielona Góra, Licealna 9, 65-417 Zielona Góra, Poland specialized structures such as extraembryonic cavities during development, absence of a larval stage, impermeable skin in adults and a urogenital system modified for obligatory internal Polish population of Aesculapian snake (Zamenis longissimus) occurs at its edge of distribu- fertilization. Lungs already evolved in their piscine ancestors, and became the exclusive site tion and it is probably isolated from the main range of its distribution. The largest local pop- for respiratory gas exchange. Probably driven by grade/clade logic, the current textbook ulation occurs in the San River Valley at foothill of Otryt (Western Bieszczady Mountains). opinion is that the ancestral type of lung in amniotes was similar to that found in amphibians: The main threat for this endangered population is a shortage of natural and anthropogenic a single-chambered and sac-like organ. Putative support of this hypothesis is the apparent nests to successful incubation and low number. presence of single-chambered lungs in the majority of lepidosaurs, including the basal Rhyncho- Distribution studies were conducted in Western Bieszczady Mountains (southeastern Poland) cephalia. We performed a large-scale comparative anatomical study of pulmonary structure in 2009–2012. Population structure was studied using capture-mark-recapture method. Habitat in all major amniote taxa. These were complemented by new developmental data from the requirements analyses were based of detailed habitat maps and monitoring of 38 locations of geckonid Paroedura picta and a review of the 19th century embryological literature. We sub- the Aesculapian snake. In 73 plots (100 m radius) around each observation 20 types of habi- mit that the so-called single-chambered lungs of lepidosaurs in fact represent a simplification tat were distinguished. Mounds with sawdust and branches (as supplementary egg-lying sites) of a plesiomorphically branched and multichambered organ, and provide functional, onto- have been built in current and historic localities and between these places along San Valley. genetic and phylogenetic morphological support. The fundamental consequence is that in- Current distribution of Aesculapian snake is not limited to Otryt region. Snakes were also stead of a single-chambered lung, one must postulate a multichambered lung at the dawn observed in Komancza, Czarna, Wetlina and coast of Solina Lake. of amniotes. The simplifications found in the extant lepidosaurs are explained by an evolu- The results show that Aesculapian snake inhabits mostly forested (deciduous forest domi- tionary scenario based on the physical properties of different pulmonary baupläne and on the nated) and open areas with anthropogenic structures. The mean number of habitat types fossil record of the lepidosauromorphan lineage. in plots was 11, average cover of particular habitats was: forest (50%; dominant species A. incana), open areas (35%), other habitats were e. g. riverbanks (7%), roads (3%), anthropo- genic structures (2%). Average length of ecotonal zone was 1.15 km. 131 individuals were caught including 18 females, 85 males and 28 juveniles. Small number of females found might be a result of their lower detectability. The most of captured indivi- duals including juveniles (>20) were caught after 2010, probably as a result of active protection. We noticed population growth connected to active conservation. Artificial egg-lying sites were rapidly occupied, especially in locations with higher number of snakes. During four years of active protection, snakes used 15 out of 25 egg-lying sites.

134 135 MONITORING ECOLOGY

A camera trap study of mass-hibernation in Salamandra salamandra The effects of competition in two temperate gecko species in the Vienna Woods, Austria Duje Lisicic 1,2, Sanja Drakulic 1, Ela Milcic 1, Anthony Herrel 3, Zeljko Mihaljevic 4, 1 1 1 Christoph Leeb 1, Eva Ringler 1, Günter Gollmann 2, Max Ringler 1 Domagoj Dikic , Vesna Benkovic , Zoran Tadic

1 1 Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; Department of Animal Physiology, Division of Biology, Faculty of Science, University of Zagreb, Rooseveltov trg 6, [email protected] 10000 Zagreb, Croatia 2 2 Department of Theoretical Biology, University of Vienna, Althanstraße 14, 1090 Wien, Austria Current address: University of Applied Health Studies, Mlinarska cesta 38, 10000 Zagreb, Croatia; [email protected] 3 UMR 7179 C.N.R.S./M.N.H.N., Département d’Ecologie et de Gestion de la Biodiversité, 57 rue Cuvier, Camera traps are commonly used in animal ecology. The fact that most systems are triggered Case postale 55, 75231 Paris Cedex 5, France by motion detection in the infrared spectrum limits their use for poikilothermic species. This 4 Department of Pathological Morphology, Croatian Veterinary Institute, Savska cesta 143, 10000 Zagreb, Croatia is one of the reasons why camera trapping is underrepresented in herpetology, compared to studies in birds and mammals. We tested a self-constructed camera trap that was triggered by Complex interspecific interactions may be important structuring agents in biological com- a light barrier to monitor the activity of the fire salamander Salamandra salamandra in the munities. Competition over spatial niche utilisation is one of most common competitive Vienna Woods (Austria) from mid-October 2011 to mid-June 2012. The camera was installed interactions between species in sympatry. We investigated the interspecific interactions at the entrance of a burrow that had been previously identified as a mass-hibernation site of between two, ecologically similar, temperate climate, gekkonid species, the Turkish gecko S. salamandra. Out of more than 20,000 photos that were taken during eight months, over (Hemidactylus turcicus) and the Moorish gecko (Tarentola mauritanica). We compared popu- 7,340 showed fire salamanders, representing a total of 214 different individuals. 168 of them lations in sympatry and allopatry on two geographically close Adriatic islands, Hvar (both were known from a capture-recapture-study, so it was possible do determine the catchment species present) and Vis (only H. turcicus). We also compared populations of H. turcicus on area of this burrow and to compare the sex ratio of the fire salamanders that were found with- Hvar occurring with T. mauritanica (in syntopy) and without T. mauritanica (in allotopy). in this area and the sex ratio of the individuals that hibernate together. Based on the photos We monitored species during their complete diurnal and annual cycle. The relative pop- taken by the camera trap we were able to generate an activity profile for each individual and ulation density of H. turcicus differs depending on the presence/absence of T. mauritanica, so we could estimate the number of individuals inside the burrow (up to 189 at the same time). but relative population densities of T. mauritanica are not affected by the presence of H. tur- Other vertebrate species were captured on these pictures as well, indicating a syntopic hiber- cicus. Additionally, the two species showed differences in habitat use in sympatry. Moreover, nation of Salamandra salamandra, Ichthyosaura alpestris, Bombina variegata, Bufo bufo, Rana H. turcicus showed ecological release in allopatry. Differences in densities between allopatric dalmatina, Rana temporaria, Anguis fragilis, Natrix natrix and Zamenis longissimus. Recently, (Vis) and allotopic (Hvar) populations of H. turcicus point to direct competition between we tested the suitability of this camera for monitoring of an amphibian tunnel. The findings H. turcicus and T. mauritanica, but this competition does not occur over spatial niche since of our study point out the importance of richly structured habitats for amphibians and this species exhibits evasive strategies for habitat utilisation when found in sympatry. Observed demonstrate the significance of detailed knowledge on specific characteristics of populations interspecies interaction patterns may contribute to a better understanding of complex rela- for effective conservation management. Furthermore we show the usefulness of camera traps tions in natural communities. for herpetological studies.

136 137 BEHAVIOUR ECOLOGY

Effects of climatic factors on the evolution of sexual chemical signals Fall migration behaviour and hibernation site selection in alpine of the Iberian wall lizard, Podarcis hispanica common frogs (Rana temporaria)

Pilar López, Jesús Ortega, José Martín Gerda Ludwig 1, Ulrich Sinsch 2, Bernd Pelster 1

Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 1 Institute of Zoology, University of Innsbruck, Technikerstraße 25, 6020 Innsbruck, Austria; [email protected] 28006 Madrid, Spain; [email protected] 2 Institute of Sciences, Department of Biology, University of Koblenz-Landau, Universitätsstraße 1, 56070 Koblenz, Germany Signals used in intraspecific communication are expected to evolve to maximize efficacy under a given climatic condition. For example, many lizards scent mark territories with chemical The common frog (Rana temporaria) is a widely distributed, European amphibian species. In secretions that are important in intrasexual relationships between males and in female mate large parts of its distribution, temperatures drop below zero during winter. This bares the risk choice. Chemical secretions might change in the evolutionary time to ensure that signals are of the formation of lethal ice crystals inside cells. However, little is known about the hiber- perfectly tuned to local humidity and temperature affecting their volatility and therefore their nation behaviour of R. temporaria. Across its distribution range, both terrestrial and aquatic persistence and transmission through the environment. Thus, interspecific differences in hibernation have been reported. Previous studies have shown that common frogs are able to chemical signals of lizards might partly reflect selection for the efficacy of signals in different tolerate mild subzero temperatures for short periods by using glucose as a cryoprotectant. climatic conditions. We used gas chromatography-mass spectrometry (GC-MS) to examine Further, they are able to meet their metabolic oxygen demand during hibernation through differences in chemical composition of femoral secretions between two nearby populations cutaneous respiration. Therefore, for common frogs at high altitude three hibernation strategies of Iberian wall lizards, Podarcis hispanica, separated along a 500 m altitudinal gradient with seem possible: aquatic hibernation in ponds or streams or over-wintering in snow-covered contrasting environmental conditions. Then, we tested experimentally, using tongue-flick terrestrial hibernacula. essays, whether female lizards were able to detect by chemosensory cues alone the males’ scent To get a first insight in the hibernation behaviour of common frogs in alpine regions we marks that were experimentally maintained under different conditions of temperature and marked 15 adult common frogs with radio transmitters in fall 2011 and followed them over humidity. We also examined whether the temporal attenuation of the chemical stimuli a 10-week period to their hibernation sites. Migratory behaviour was assessed by calculating depended on the simulated climatic conditions. Results showed that the efficacy (i. e. detect- the line-of-sight distance between relocations. The mean distance animals moved between ability and persistence) of scent marks for each population is maximal under the original envi- two relocations was 19.4 m (SE=4.1). Migratory behaviour did not differ between males and ronmental conditions of each population, which may depend on their chemical composition. females. Among the radio tracked frog aquatic hibernation prevailed. One frog hibernated in We suggest that the abiotic environment may infer a selective pressure on the form and a pond, seven in streams or springs and none in terrestrial habitat. Overall mortality rate was expression of sexual signals, which might have consequences for interpopulation recognition 40%. and result in population divergence and speciation. Our study also suggests that rapid climate The distance travelled between relocations was significantly influenced by season, suggesting warming could lead to negative changes in the efficacy of sexual signals with potential detri- that factors such as photoperiod or availability of food might play a major role in triggering mental consequences for the sexual selection and conservation of lizards. the onset of hibernation in R. temporaria. Hibernation in lotic water might be beneficial due to constant microclimatic conditions, allowing frogs to maintain a reduced level of metabolic rate while guaranteeing a constant supply of oxygen. Both might be crucial for their overall energy balance.

138 139 SYSTEMATICS PHYLOGEOGRAPHY AND PHYLOGENY

A putative cryptic species of Galápagos marine iguana on the brink Phylogeographic relationships of green toads (Bufo v. viridis) from of extinction isolated alpine populations

Amy MacLeod 1, Volker Koch 2, Carolina García-Parra 3, Fritz Trillmich 1, Andreas Maletzky 1,*, Christophe Dufresnes 2,*, Nicola Novarini 3, Lucio Bonato 4, Sebastian Steinfartz 1 Matthias Stöck 2,5

1 Department of Animal Behavior, Unit of Molecular Ecology and Behavior, University of Bielefeld, Morgenbreede 45, 1 University Salzburg, Department of Organismal Biology, Hellbrunnerstraße 34, 5020 Salzburg, Austria; 33619 Bielefeld, Germany; [email protected] [email protected], 2 Investigación para la Conservación y el Desarrollo, La Paz, BCS, México 2 University of Lausanne, Department of Ecology and Evolution, Biophore, 1015 Lausanne, Switzerland 3 Charles Darwin Research Station, Puerto Ayora, Santa Cruz Island, Galápagos, Ecuador 3 Museo di Storia Naturale di Venezia, Santa Croce 1730, 30125 Venezia, Italy 4 Università di Padova, Department of Biology, via Ugo Bassi 58b, 35131 Padova, Italy The recent discovery of strong population structuring in the Galápagos marine iguana indi- 5 Leibniz-Institute of Freshwater Ecol. and Inland Fisheries (IGB), Müggelseedamm 301, 12587 Berlin, Germany cates the potential existence of a cryptic species, the Punta Pitt iguana, represented by a single * equal contribution small discrete population on San Cristobál Island. Though data from both mitochondrial sequences and 13 microsatellite loci indicate this population to be highly distinct and probably Green toads (Bufo viridis subgroup) inhabit an enormous Palearctic range (at least 12 mtDNA ancestral, the small number of sampled individuals had until recently severely limited analysis. clades, including polyploids: 16 forms). In Austria, the nominotypical B. v. viridis occurs in During an expedition to San Cristobál in 2012, we collected 130 new blood samples, many the eastern lowlands (type locality: Vienna), while the Alps delimit the range to the West, with from previously unknown locations. Utilising 19 highly polymorphic microsatellite loci, we only three isolated patches in Salzburg, Tyrol and adjacent Bavaria, located up to 1,400 m analysed population structure on the island using STRUCTURE, calculated levels of genetic a.s.l. In northeastern Italy, B. v. viridis inhabits pre-Alpine regions and the lowland, reach- differentiation (Reynolds FST) in ARELEQUIN, and estimated the effective population size ing parapatric B. balearicus at the Po River. To understand colonization and relationships of (Ne) using three methods: ONESAMP, MLNE and TM3. STRUCTURE analysis supported remote populations, we studied 77 individuals from isolated and neighboring localities using the existence of two strongly divergent and reproductively isolated populations on San Cristóbal, mtDNA and seven microsatellites. mtDNA shows toads at some localities from the interior separated by an FST of 0.15, the highest level of intra-island genetic distance found in this Alps (S Bavaria, Tyrol) to share identical haplotypes with southern pre-Alpine populations (N species. Though the geographical distance between the two populations is less than 12 km Venetia), suggesting a genetic contribution to Alpine ones. However, microsatellite genotypes of coast, the level of genetic divergence is equal to that found between island populations indicate strong differentiation between populations from the South (Venetia) and the North separated by more than 140 km of ocean. We therefore suggest that the marine iguanas of the Alps (W Austria, Bavaria). In addition, despite geographic proximity, N Alpine green from Punta Pitt represent a currently undetected, i. e. cryptic, species of marine iguanas. Three toads form several distinct genetic groups (W Tyrol, E Tyrol, S Bavaria), testifying of strong separate methods estimated the Ne of the Punta Pitt population to be 50 or fewer, indi- genetic drift and population disconnection in these remote parts of the range. We also find cating that urgent conservation management may be required. In particular, predation by inva- indications for potential human introductions in at least one of these populations. sive feral cats is a serious concern that warrants further attention. Considering the small pop- ulation size and the fact that all analyses to date indicate reproductive isolation and a deep split between this population and all others, we pose the question of whether Punta Pitt iguanas should now be considered a cryptic species on the brink of extinction.

140 141 ECOLOGY SYSTEMATICS

Does foraging plasticity favours adaptation to new habitats in fire A history of species delimitation methods applied in the Sceloporus salamanders? Preliminary data grammicus species complex and what it tells us about the future

Raoul Manenti 1, Mathieu Denoël 2, Gentile Francesco Ficetola 3 Jonathon C. Marshall

1 Dipartimento di Bioscienze, Università degli Studi di Milano, Via Celoria 26, 20133 Milano, Italy; Department of Zoology, Weber State University, Ogden, UT 84408, USA; [email protected] [email protected] 2 Laboratory of Fish and Amphibian Ethology, Behavioural Biology Unit, Department of Biology, Ecology, and Evolution, University of Liège, Belgium The historical developments of species identification and delimitation methods have been 3 Dipartimento di Scienze dell’Ambiente e del Territorio, e di Scienze della Terra Università degli Studi di Milano- played out and reflected in the Sceloporus grammicus species complex of México. Early mor- Bicocca, Piazza della Scienza 1, 20126 Milano, Italy phological studies were followed by allozyme studies and soon scores chromosomal races were being described based on cytogenetic analyses in the 1970s and 1980s. These gave way to Predators often show strong plasticity of optimal foraging strategies. A major difference in restriction fragment digest analysis in both mitochondrial and ribosomal DNA and then to foraging strategies occurs between sit-and-wait and active predators. Environmental condi- DNA gene sequencing, ecological niche modeling, and now thoughts of next generation tions affect their efficiency, with active foraging behaviour being favoured when prey is scarce sequencing. The history of attempts to identify species boundaries in S. grammicus illustrates or its detection is difficult. Both phenotypic plasticity and local adaptations may cause a shift the complex nature and intrinsic difficulty of delimiting species even as our technology and between the two strategies. Here we studied larvae of Salamandra salamandra originating from analytic prowess has exploded over the last half century. In many cases there is no ‘silver bullet’ either typical epigeous streams and from caves. We evaluated whether local adaptations or to delimit species boundaries and the S. grammicus complex reminds us of the necessity of an phenotypic plasticity determine the shift of foraging strategy between stream and cave pop- integrative approach in understanding this nebulous entity we call a species. ulations. The foraging behaviour of larvae was evaluated using a full-factorial design, taking into account three test conditions: light, prey availability, and starvation (larvae with and with- out an available prey in darkness and light, sate or starved larvae tested in light and in dark- ness). Behaviour was recorded both visually and through video-tracking. Salamander larvae modified their behaviour in response to environmental conditions. When in the darkness, salamanders moved longer distances. Movements also increased in starving larvae, and with prey occurrence. Furthermore, larvae from cave populations showed higher behavioural plas- ticity than stream larvae, as they changed more their foraging strategy according to light con- ditions. Cave larvae also better exploited the space available in test environments. Variation of foraging behaviour was strong, and involved complex interactions between plasticity and local adaptations. When larvae were in conditions similar to the ones encountered in caves, plasticity enabled behavioural shifts toward an active foraging strategy. The higher behavioral plasticity showed by cave larvae supports the importance of this trait for the exploitation of novel environments, as caves are for epigeous fauna.

142 143 PHYLOGEOGRAPHY AND PHYLOGENY PRACTICAL CONSERVATION

Trapped in the Iberian Peninsula: reconstructing the evolutionary An integrative approach for inferring spatial and temporal patterns history and predicting suitable refugia under climate change for the for snakes fatalities on electricity distribution networks: the case of endemic Vipera seoanei Rhinechis scalaris in Spain

Fernando Martínez-Freiría 1, Guillermo Velo-Antón 1, José C. Brito 1,2 Fernando Martínez-Freiría 1, Marcial Lorenzo 2, Pablo García-Díaz 2,3, Miguel Lizana 2

1 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto, 1 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto, Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal; Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal [email protected] 2 Departamento de Biología Animal, Universidad de Salamanca, Campus Miguel de Unamuno, 37007 Salamanca, 2 Departamento de Biologia da Faculdade de Ciências da Universidade do Porto, Rua Campo Alegre, 4169-007 Porto, Spain; [email protected] Portugal 3 The School of Earth and Environmental Sciences, University of Adelaide, Adelaide, South Australia 5005, Australia

Pleistocene climatic oscillations are major factors related to current species’ biogeographic Electricity facilities contribute to habitat fragmentation, causing mortality mainly in birds patterns and climate change is predicted to strongly affect species distribution patterns. and bats. Since 1998, the Spanish electricity company Iberdrola has registered fatalities by Integrative approaches identifying genetic and ecological variability are needed to better electrocution of the terrestrial fauna, resulting in 70 cases for the ladder snake (Rhinechis understand the effect of climate on the species evolutionary and biogeographic histories, and scalaris) but this number is suspected to be underestimated. Fatalities occur as a consequence for coherent conservation planning. The Iberian adder Vipera seoanei is a Euro-Siberian species of snakes climbing to medium voltage power towers (MVPT) and could be related to their restricted to northern Iberia, for which available evolutionary scenarios are solely inferred hunting activity (active foraging). In order to identify areas and times of high electrocution through morphological studies and suitable climatic space was predicted to disappear in 2020. risk and also testing the hypothesis that fatalities are related to the number of MVPT in a This work aims to reconstruct its evolutionary history and to identify suitable areas for the given area, multivariate analyses, Geographic Information Systems, Ecological Niche-based future. Fifty-eight tissue samples from specimens covering the species distributional range were Models and circular statistics were combined with the environmental variability associated sequenced for two mitochondrial gene fragments (cytb, ND4) and sequences were analysed to the fatalities and the geographic distribution of MVPT. Two environmentally coherent using Bayesian approaches. Ten different ecological modelling algorithms were used over 352 groups have been recognized, each one presenting specific spatial and temporal patterns. distributional records at 1 x 1 km and climatic variables for current, past (LIG, Last Inter- Ecological models identified probable areas of occurrence and environmental conditions Glacial; LGM, Last Glacial Maximum) and future (2050 and 2080) conditions. Vipera seoanei for each group within the species ecological requirements. No strong correlations were found was found to have split from V. berus likely in the Pliocene and its shallow phylogeographic between the density of MVPT and density of fatalities or potential areas for groups, sug- structure indicates a single refugial population followed by population expansion. Projections gesting that snakes could be attracted to specific MVPT. Different seasonal patterns for both to past conditions identified probable suitable areas that may have acted as refugia for the groups were related to climatic factors varying on a monthly scale, reflecting snakes’ activity species in Iberia, being smaller in the LIG than in the LGM and present time. Predictions periods. Common corrective policies among Iberdrola and other Spanish electric companies according to future climatic conditions suggested dramatic reductions of distributional ranges should be performed, since areas of high risk of mortality for snakes are likely to occur inside and population persistence in north-western Iberia. Contrarily to other European viper species and outside the studied electricity network. Moreover, experimental studies of snake’s attrac- for which population isolation and refugia are suggested during Pleistocene cold periods, this tion to MVPT and monitoring studies of populations in areas of high probability for fatalities study indicates that V. seoanei was deeply affected by warm periods. Consequently, populations should be made for a fine identification of the factors triggering the species electrocution and monitoring and local management actions are likely to be needed under ongoing climate change. for their prevention. Current methodological approach is revealed as useful for further inves- tigations on fatalities in other terrestrial species, even with low sample sizes.

144 145 BEHAVIOUR ECOLOGY

Phonotaxis field experiments on Peruvian poison frogs—do male Occurrence of amphibian deformities in the Egyek-Pusztakócs marsh frogs respond to modified calls of syntopic congenerics? and grassland system (Hortobágy)

Michael Mayer, Stefan Lötters Béla Mester 1, Szabolcs Lengyel 1, Miklós Puky 2

Department of Biogeography, Trier University, 54286 Trier, Germany; [email protected] 1 University of Debrecen, Faculty of Science and Technology, Department of Ecology, Conservation Ecology Research Group, Egyetem tér 1, 4032 Debrecen, Hungary; [email protected] We studied acoustic communication in the Peruvian poison dart frog Ranitomeya imitator, 2 MTA Centre for Ecological Research, Danube Research Institute, Jávorka S. utca 14, 2131 Göd, Hungary which performs Müllerian mimicy with Ranitomeya variabilis. In order to avoid the risk of ‘wasting’ energy for territorial defense we suggest that aggressive and territorial R. imitator The occurrence of amphibian deformities is well known since the 18th century but reports males are able to discriminate between conspecifics and its co-model species by differently have become more frequent in the last two decades. Several factors, e. g. parasites, were proved structured advertisement calls. We conducted phonotaxis field experiments attracting wild R. to cause such changes, in most cases it is unclear why they developed. Above the 2% back- imitator males towards (i) the R. imitator advertisement call, (ii) the R. variabilis advertise- ground frequency deformities may be considered as an indicator of environmental distur- ment call, (iii) four modified calls with the temporal call parameters note length and inter- bance. Though in Hungary deformities had been detected in at least 13 amphibian taxa, only note interval substituted between the two species and (iv) two R. imitator calls at different limited information is available from the Hortobágy region. Our main aim was to monitor dominant frequencies beyond the species’ range. Experiments were videotaped and the male the malformation frequency of the Egyek-Pusztakócs marsh and grassland system. We have reaction was measured in approach time, number of jumps, jump distance, jump angles and collected data since 2010 by using several field methods e. g. pitfall and bottle traps, dip net- path straightness. The response intensity of R. imitator towards its original advertisement call ting and catching individuals by hands during several surveys. We collected more than 5,000 was used as reference for ‘normal’ territorial behavior. Males of R. imitator neither showed specimens in 2010–2012. Amphibian deformities occurred in 2010, an extremely wet year with responses towards R. variabilis advertisement call, nor to R. variabilis call when modified in good reproductive success, even if their frequencies were below the background value. Ectro- temporal call parameters to match the R. imitator call. In contrast, when playing imitator- mely was present in 60% of the cases, mainly in juvenile anurans (Bombina bombina, Pelo- playbacks with properties of R. variabilis, frogs approached the sound source. Furthermore, phylax esculentus complex). Besides anurans, we also collected four deformed newts (Triturus approaches could be provoked when only modified in dominant frequency. Results suggested dobrogicus), one of them had a duplicated tail, which is a very rare morphological anomaly. that our modified call parameters did not have significant effects on R. imitator males’ ability In all other cases limb or tail reductions were recognized. The observed malformations may to discriminate their own advertisement call from that of R. variabilis. Nevertheless, we could be caused by predators, such as birds or fish, which can be rather abundant in the Egyek- show that R. imitator is able to discriminate between conspecifics and its syntopic co-model Pusztakócs marsh system. species by its advertisement call. This leaves to conclude that the call parameters note length, internote interval or dominant frequency are not essential in terms of call discrimination in R. imitator.

146 147 ECOLOGY DISTRIBUTION MODELLING

The impacts of glyphosate and predation threat on tadpoles of Use of distribution modelling for exploring new localities of an the agile frog (Rana dalmatina) in different experimental venues endangered European viperid, Vipera ursinii graeca

Zsanett Mikó, János Ujszegi, Zoltán Gál, Attila Hettyey Edvárd Mizsei 1, Bálint Üveges 2, János Pál Tóth 1

“Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, 1 Department of Evolutionary Zoology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary; Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] [email protected] 2 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, The widespread application of pesticides emphasises the importance of understanding the Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary impacts of these chemicals on natural communities. Toxicologists traditionally use laboratory- based tests to determine lethal and sublethal concentrations of pesticides. However, these tests Vipera ursinii is one of the most endangered snake species of Europe and provoked strong may over- or underestimate the impacts of pesticides by neglecting possible indirect and inter- conservation approaches, and most of its subspecies are subject to elaborate conservation active effects. For example, it is known that the presence of predators can alter the lethality programmes. The only European exception is the Greek meadow viper (V. u. graeca), which of chemical contaminants. Consequently, analysing pesticide impact under more natural is the least studied subspecies in Europe. Furthermore a recent study suggest that it may be conditions allows for estimating realistic effects of chemicals on non-target organisms. The a separate, basal species to the V. ursinii complex, thus it may receive a novel conservation most commonly applied broad-spectrum herbicide in the world is glyphosate, which is usually status in the future. We have modelled the potential distribution of V. u. graeca with MaxEnt, applied together with surfactants. We exposed agile frog (Rana dalmatina) tadpoles to three derived from four ecogeographical layers and 28 presence data in an attempt, to update the concentrations (zero, low and high) of glyphosate, combined with three predator treatments list of habitats for this subspecies. We tested the model at six localities, five sites in Albania (no predator, Aeshna cyanea or Lissotriton vulgaris) in a randomized block design. To allow for and one in Greece. We identified the observed vipers based on morphological characteristics a direct comparison of results obtained in standard ecotoxicology tests typically done in the and mitochondrial ND4 sequences. We found V. u. graeca at three mountains in Albania and laboratory with those performed in outdoors mesocosms, we performed the experiment in one in Greece, from which two were undocumented localities for this viper until now. Our both venues. We surveyed tadpole survival, behaviour, growth, and morphology. Our results results show that distribution models are very useful and accurate for designing field research. showed that the type of venue influenced almost all investigated traits. Under laboratory con- Furthermore, the model prediction shows further 24 potential, but yet undiscovered habitats ditions predator-stress increased the effect of glyphosate on tadpole mortality, but in the for V. u. graeca in addition to 16 already known localities. mesocosms it made the herbicide less lethal, most probably because of the interactive effects of the stratification of the chemical and of the antipredator-response shown by tadpoles. Our study suggests that the type of experimental venue and the inclusion of further stress factors can have large effects on results of ecotoxicological experiments.

148 149 DISTRIBUTION MODELLING BEHAVIOUR

Evidence for niche conservativism in the Vipera ursinii complex Experimental evidence for environmental factors affecting colouration in the European green lizard (Lacerta viridis) Edvárd Mizsei 1, Márton Szabolcs 2, Patrik Katona 1, Szabolcs Lengyel 3 Orsolya Molnár 1, Katalin Bajer 2, János Török 3, Gábor Herczeg 3 1 Department of Evolutionary Zoology and Human Biology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary 1 Dartmouth College, Hanover, New Hampshire 03755, USA; [email protected] 2 Nature Conservation Association of Tokaj, Liget köz 1, 3910 Tokaj, Hungary; [email protected] 2 Universidade Federal do Rio Grande do Norte, Campus Universitário–Lagoa Nova, Natal-RN 59078-900, Brazil 3 Department of Tisza River Research, Hungarian Academy of Sciences, Centre for Ecological Research, Bem tér 18/c, 3 Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary 4026 Debrecen, Hungary Carotenoid colouration has been previously demonstrated to act as an honest sexually selected The Vipera ursinii species complex is a group of highly endangered small-sized snakes of steppe signal in numerous species due to costs raised by the synthesis of pigments and precursors. and alpine grasslands in the Palearctic region. The systematics of this complex, comprising Since the quality of an individual determines the amount macromolecules it can consume 10 to 17 taxa (species and subspecies), differs by whether reconstruction is based on morpho- from the environment, carotenoid signals should raise high costs for the owner, thus provid- logical or on molecular data. Here we use phylogenetic comparisons to study the relation- ing reliable information about owners’ genetic quality. In previous studies, we demonstrated ships between environmental properties (mainly climate) and niche overlap among members the role and described the information content of an structural (ultraviolet) colour patch of the species complex. We modelled the ecological niche of 17 taxa (species and subspecies) in male European green lizards (Lacerta viridis), but importance of colouration on other derived from 11 ecogeographical (climate and topography) and 216 “presence-only” data. The body parts remains unknown. Our aim in the current study was to investigate, whether the ecological niche models were then used to compute pairwise niche overlap values between all carotenoid (yellow) and structural (UV) colouration of the belly had any costs associated taxa. Our results showed that morphological similarity was positively correlated with the ex- with development or maintenance, and thus could possibly play a role in sexual selectional tent of niche overlap. We detected a higher level of niche conservativism among the molecular processes. During the breeding season of 2009, we captured 60 adult male lizards and created sister taxa than among non-sister taxa. Furthermore, we found high niche overlap between four groups experiencing optimal or sub-optimal environmental conditions. Temperature lowland taxa and greater niche divergence among mountain taxa, which may be related to and food treatments were applied in factorial design, and colour and morphological traits the greater local adaptation to altitudinally changing and locally more diverse environmental were measured before and after experimental sessions. Results showed that both carotenoid factors in mountain habitats compared to lowland habitats. Our study provides evidence that and structural components of belly colouration were affected by temperature and amount of (i) morphological similarity is related to niche overlap, (ii) that niche conservativism may be available food, with animals experiencing sub-optimal environmental conditions elaborating strong and (iii) that niche overlap can be related to variation in environmental factors (e.g., duller colour. This suggests that belly colouration is highly costly to develop and maintain, altitude, climate) in the studied group of vipers. therefore can possibly act as an honest sexual signal. Considering that throat colouration has been shown to function as a multiple signaling system, it is possible that other colour traits, like belly colour, are also taken into account when estimating individual quality.

150 151 SYTEMATICS ECOLOGY

The study of correlation between age, body size and pattern Effects of prolonged summer drought and simulated complete water (number and size of spots) in some Neurergus derjugini and food deprivation on body condition and osmotic responses in (Nesterov, 1916) population from North-West of Iran Souss Valley tortoises (Testudo graeca soussensis) from an arid area of West-Central Morocco Elnaz Najafi-Majd, Elif Yildirim, Ug˘ur Kaya Hichami Nawal, Mohammed Znari, Mohamed Naimi, Salwa Namous Ege University, Faculty of Science, Biology Section, Zoology Department, 35100 Bornova-Izmir, Turkey; [email protected] Laboratory “Biodiversity and Ecosystem Dynamics”, Department of Biology, Faculty of Science, Semlalia, Cadi Ayyad University, Bd Prince Moulay Abdellah, P.O. Box 2390, 40000 Marrakech, Morocco; and Number of spots and spots size has been used as a distinguishing taxonomical character between Natural History Museum of Marrakech, Cadi Ayyad University, Avenue Allal El Fassi, Marrakech; [email protected] Neurergus derjugini (Nesterov, 1916) and its close relatives. As a result of the present confu- sion in the taxonomy of the closely related taxa, N. derjugini microspilotus and N. derjugini Because of the unpredictable availability of resources in arid environments, adult Souss Valley derjugini based on number and size of spots on the dorsum of specimens, we investigated the tortoises are able to survive long periods without food and water. We investigated the effects age structure and its relationship to the number of spots and body size in some populations of summer drought and simulated complete water and food deprivation (at 25 °C for 20 of N. derjugini from North-West of Iran. weeks) on body condition (g body mass/cm3 body volume) and osmotic responses, respec- Age structure was studied by using skeletochronology performed on the phalanges. The tively, in free-ranging tortoises in an arid steppe-land of West-Central Morocco, and captive analysis of the age structure in five populations (45 males, 35 females) based on continuing tortoises during the summer period 2012 (June–October). Body condition in aestivating tor- lines of arrested growth (LAGs) demonstrated that age in males rages from 9 to 22 years toises, decreased markedly from 0.211±0.005 in early June to 0.166±0.009 in early October. (mean=12.14±2.8) and from 10 to 20 years (mean=14.17±2.5) in females. The mean Snout- Osmolality initially dilute bladder urine in early June increased until it was isosmotic to plasma Vent Length (SVL) was 129.23±8.27 mm in males and 150.23±10.6 mm in females and the by mid-summer, after which osmolality of both fluids increased to a highest level at the end SVL difference between the sexes size was statistically significant. The mean spots numbers of drought period. The increase in plasma osmolality comprised increases in plasma sodium, was 32.07±9.5 in males and 39.69±9.7 in females. The number of spots and age were signifi- chloride, and especially urea concentrations. Bladder urine osmolality increased due primarily cantly correlated in both males and females (Pearson’s correlation, males: r=0.790, P<0.01; to increased soluble potassium from their diet apparently osmotically stressful. When rainfall females: r=0.743, P<0.01). The number of spots and body size were also significantly correlated occurred, tortoises drank copiously, voided concentrated bladder urine, and stored dilute urine; in both males and females (Pearson’s correlation, males: r=0.644, P<0.01; females: r=0.607, body mass, plasma and urine concentrations returned back to hydrated levels. Tortoises P<0.01). exhibited a tolerance of temporary “anhomeostasis” and that allows them to osmoregulate Our results show that N. derjugini is a long-lived species exhibiting sexual size dimorphism opportunistically. Body mass loss during the deprivation period was only 17.28±4.83% of ini- and younger animals have fewer and larger spots than the older ones, therefore number of tial body mass with an average body mass loss rate of 0.00196±0.00055 d–1. Body condition spots and spots size cannot be considered a reliable taxonomic character in these taxa. index decreased from 0.1762±0.0336 to 0.1379±0.0263. Tortoises completely deprived of water and food, drank an average of 11.16% of their body mass in a mean time interval of 14±4.43 mn. The lack of, or only small, disturbances in tortoises naturally aestivating and experimentally deprived of food and water for a prolonged period is indicative of their high degree of physio- logical adaptation to the summer extreme dry conditions.

152 153 ECOLOGY BEHAVIOUR

Divergent altitudinal phenotypes in Iberian wall lizards are not driven The homing frog: orientation and homing behavior in a territorial by different life-histories dendrobatid frog Allobates femoralis

Jesús Ortega, Pilar López, José Martín Andrius Pasˇukonis 1, Walter Hödl 2

Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 1 Department of Cognitive Biology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; 28006 Madrid, Spain; [email protected] [email protected] 2 Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria The interplay between ecological conditions and life histories has been widely acknowledged in vertebrates, particularly in lizards. Temperature, food availability and humidity may exert Dendrobatidae (dart-poison frogs) exhibit some of the most complex spatial behaviors among different selective pressures, and, hence, produce divergent phenotypes even in geographically amphibians, such as territoriality and tadpole transport from terrestrial clutches to widely close populations. In this context the Iberian wall lizard, Podarcis hispanica, constitutes a per- distributed deposition sites. High homing performance of tadpole transporting adults can fect organism model as it is considered a species complex with a complicated evolutionary be often assumed but the experimental evidence is lacking and the orientation mechanisms are history. Here we report the results of a common garden experiment to test for life-history dif- unknown. ferences of two nearby P. hispanica populations separated along a 500 m altitudinal gradient Our study is aimed at understanding and quantifying the orientation ability of male with contrasting environmental conditions, where adults show marked morphological and Allobates femoralis, a dendrobatid frog with paternal extraterritorial tadpole transport. First, pheromonal differences in instead of being closely related genetically. We specifically focus on we conducted a translocation experiment to quantify the homing success and speed under reproductive investment, incubation time, morphology at hatching and growth rate differences. natural conditions. Additionally, we tested the initial orientation of the translocated males in Results showed differences in clutch size and egg morphology mediated by a larger body size an arena assay. Finally, we obtained homing trajectories by tracking individuals equipped with in highland females and size independent differences in clutch weigh and volume. However, miniature transponders. hatchling lizards only differed in head height even after controlling for body size. A mixed We found that translocated A. femoralis return successfully from up to 400 m and show a model ANOVA (with clutch as a random factor and population and sex as fixed factors) did very high homing success for distances up to 200 m. Individuals also show significant home- not show significant differences between populations, nor between sexes, in mass-specific ward orientation in the arena essay. Tracking experiment revealed that homing trajectories are growth, size specific growth, head growth or femoral growth of hatchling lizards. Overall, characterized by long periods of immobility (up to several days) and short periods (several we provide evidence that phenotypic differences are not driven by different life-history traits. hours) of rapid movement closely fitting a straight line towards the home territory. Also we infer that this pattern of variation is caused by environmental differences in produc- We observed that distances at which homing performance is high correspond to the longest tivity linked to contrasting climatic conditions. Thus, we suggest that an important role of distances covered by males during tadpole transport. Consequently, our results demonstrate phenotypic plasticity during ontogeny could be responsible of adult morphological and life strongly developed and very precise orientation ability within a potential area of familiarity. history differences between both populations. We discuss possible orientation mechanism involved and our future plans to test them.

154 155 SYSTEMATICS PHYLOGEOGRAPHY AND PHYLOGENY

Resolving the publishing bottleneck and increasing data Phylogeny and phylogeography of the Podarcis tauricus group: interoperability in biodiversity science insights from mitochondrial and nuclear data

Lyubomir D. Penev 1,3, Teodor A. Georgiev 3, Pavel E. Stoev 2,3, David M. Roberts 4, Nikos Psonis 1,2, Oleg V. Kukushkin 3, Boyan Petrov 4, Jelka Crnobrnja-Isailovic´ 5,6, Vincent S. Smith 4 Iulian Gherghel 7, Petros Lymberakis 1, Nikos Poulakakis 1,2

1 Institute for Biodiversity and Ecosystem Research, Bulgarian Academy of Sciences, Gagarin St. 2, 1113 Sofia, Bulgaria 1 Natural History Museum of Crete, University of Crete, Knosos Avenue, 71409 Irakleio, Greece; 2National Natural History Museum, Bulgarian Academy of Sciences, Tsar Osvobiditel Blvd. 1, 1000 Sofia, Bulgaria [email protected] 3 Pensoft Publishers, Prof. Georgi Zlatarski St. 12, Sofia, Bulgaria; [email protected] 2 Department of Biology, University of Crete, Vassilika Vouton, 71409 Irakleio, Greece 4 The Natural History Museum, Cromwell Road, London, United Kingdom 3 Karadagh Nature Reserve of Ukrainian National Academy of Sciences, Nauki St. 24, 98188 Theodosiya, Republic of the Crimea, Ukraine “Publishing bottleneck” is caused by: (1) increasing amount of data due to the intensification 4 National Museum of Natural History, 1000 Sofia, Bulgaria 5 of scientific exploration increases; (2) publishing in non-machine-readable formats, e. g. paper/ Department of Biology and Ecology, Faculty of Sciences and Mathematics, University of Nisˇ, Visˇegradsˇka 33, 18000 Nisˇ, Serbia PDF; (3) low uptake and inconsistent policies for data publishing; (4) pressure of adminis- 6 Department of Evolutionary Biology, Institute for Biological Research “S. Stankovic´”, Despota Stefana 142, trators to publish in “high-impact” journals; (5) increasing difficulties with peer-review. 11000 Beograd, Serbia In an attempt to solve some of the problems, Pensoft has launched the Biodiversity Data 7 Department of Zoology, Oklahoma State University, Stillwater, USA Journal (BDJ) (www.pensoft.net/journals/bdj), the first journal ever to complete the cycle from writing a manuscript, through submission, via community peer-review, to publication and The wall lizard genus Podarcis (Sauria, Lacertidae) comprises 20 currently recognized species dissemination within a single, fully XML-based, online collaborative platform. The journal in southern Europe, where they are the predominant reptile group. The taxonomy of Podarcis changes thoroughly the way how biodiversity information, irrespective of whether it is taxo- is complex and unstable. Based on DNA sequence data the species of Podarcis fall into four nomic, floristic/faunistic, morphological, genomic, phylogenetic, ecological or environmental main groups that have substantial geographic coherence (Western Island group, south- data, is published. western group, Italian group and Balkan group). The Balkan species are divided in two The scientific quality and importance of the papers and data submitted to BDJ will be subgroups: the subgroup of P. tauricus (P. tauricus, P. milensis, P. gaigeae and P. melisellensis), judged by the scientific community, through a novel community-based pre-publication and and the subgroup of P. erhardii (P. erhardii and P. peloponnesiacus). Here, we explored tauricus’ post-publication peer-review. Furthermore, the journal will provide automated registration taxonomy and investigated the evolutionary history of the species by employing phylogenetic of nomenclature acts, such as new taxa and new synonymies, in ZooBank, thus facilitating and phylogeographic approaches and using both mitochondrial (mtDNA) and nuclear (msats) authors. To mobilize taxonomic information and increase the interoperability of biodiversity markers. The phylogenetic relationships and the genetic distances retrieved, supported the data, a technologically advanced platform called “Pensoft Writing Tool” was developed. The monophyly of Podarcis tauricus group and suggest that P. gaigeae, P. milensis and P. melisellensis tool provides a set of pre-defined but flexible article templates, as well as a search and import form a clade, which thereinafter connects to P. tauricus. Within the previous clade, P. gaigeae function from external databases, electronic registries, occurrence data in Darwin Core for- is more closely related to P. milensis than to P. melisellensis. However, P. tauricus is subdivided mat, reference bibliographies, track change and comments tools, revision history and version in several lineages with high genetic differentiation, leaving insinuations whether it is a single control, etc. To keep costs affordable, all manuscripts submitted to BDJ must be either writ- species or species complex. The phylogeographical scenario emerging from the genetic data ten in the Pensoft Writing Tool (PWT), or submitted from integrated external platforms, suggests that the present distribution of this group was determined by a combination of dis- such as Scratchpads or GBIF’s Integrated Publishing Toolkit (IPT). persal and vicariance events in the Balkan Peninsula dating back to Miocene and continuing BDJ is being launched within the EU-funded project ViBRANT (www.vbrant.eu). up to Pleistocene. Thus, our data stress the need for a reconsideration of the evolutionary history of the P. tauricus group and help overcome difficulties that classical taxonomy has encountered at species level.

156 157 ECOLOGY ECOLOGY

Habitat use of the common spadefoot toad (Pelobates fuscus) in A combination of divergence and conservatism in the niche evolution Estonia—is forest essential? of Tarentola mauritanica (Gekkota: Phyllodactylidae)

Riinu Rannap 1, Maris Markus 1, Tanel Kaart 2 C. Rato 1, D. J. Harris 1, A. Perera 1, S. B. Carvalho 1, M. A. Carretero 1, D. Rödder 2

1 Institute of Ecology and Earth Sciences, University of Tartu, Vanemuise 46, 51014 Tartu, Estonia; 1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de [email protected] Vairão, 4485-661 Vairão, Portugal; [email protected] 2 Estonian University of Life Sciences, Kreutzwaldi 48, 51006 Tartu, Estonia 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany

Habitat loss and degradation are the most obvious and acute factors which influence amphib- The quantification of ecological niche overlap and the integration of species distribution ian decline. Fortunately, however, habitat deterioration is potentially reversible. Recognizing models (SDMs) with calibrated phylogenies to study niche evolution are becoming not only the factors critical for the species is therefore essential for successful habitat restoration and powerful tools to understand speciation events, but also as proxies regarding the delimitation of maintenance. We explored the habitat requirements of the common spadefoot toad (Pelobates cryptic species. In this study, we applied these techniques in order to unravel how ecological fuscus), a severely declining species in Europe. The toad is considered a typical representative niche evolved during cladogenesis within the Tarentola mauritanica species-complex. of herpetofauna in open agricultural landscapes but at its northern and western range edge Ordination methods as well as SDMs are used to estimate niche differences and similarities the species occurs also in forested areas. So far, studies on the habitat requirements of the between the lineages of T. mauritanica. In order to unravel the evolution of environmental common spadefoot toad have only focused on agricultural areas and the significance of forest niche tolerances, the SDMs are merged with a known calibrated phylogeny for the genus habitats has remained indistinct. In order to determine the habitat characteristics vital for the Tarentola. species as well as the function of forests as possible sink or source habitats, we explored Diversification of the Tarentola mauritanica complex involved both niche divergence and 34 water bodies and their surroundings comparatively in forested and open landscapes in niche conservatism, with a pattern that changes depending on whether the variables involved Estonia. The fieldwork was carried out from June to August 2008 and the method used for are related to mean temperature and humidity or with the seasonality of these variables. detecting the species was dip-netting of larvae. Our study demonstrated that land cover type Between the Miocene and Pleistocene, the diversification of the lineages encompassed by had no significant influence on the presence of the species or its larval abundance—the lim- T. mauritanica resulted from a combination of niche conservatism and niche shift, with some iting habitat factors were related to soil type (sandy soils preferred) and quality of the repro- clades still maintaining a realized niche distinct from their relatives. Moreover, our results duction site only. Respectively, the species preferred large fish free sun exposed water bodies suggest that climatic variables related to humidity and temperature seasonality were involved in areas with sandy soil. In our study sites such conditions were best met in old growth coni- in the niche shift and genetic diversification of the European/North African clade during the ferous forest areas. Thus we suggest that these types of forests holding viable populations of Pleistocene and to its maintenance in a realized niche distinct from that of the remaining the common spadefoot toad should be protected with large scale clear cuttings as well members of the group. This study further highlights the need for a taxonomic revision of this as altering of hydrology avoided. species-complex.

158 159 BEHAVIOUR ECOLOGY

Sometimes bigger is better: the role of body size, nuptial colouration New home, new life: linking the evolution of salamander larval and parental relatedness for paternity success in moor frogs habitat choice to ecosystem processes (Rana arvalis) Timm Reinhardt 1, Sebastian Steifartz 2, Markus Weitere 1 Anna Rausch 1, Marc Sztatecsny 1, Eva Ringler 1, Robert Jehle 2, Walter Hödl 1 1 Helmholtz-Zentrum für Umweltforschung (UFZ), Department Fließgewässerökologie (FLOEK), Brueckstraße 3a, 39114 Magdeburg, Germany; [email protected] 1 Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; [email protected] 2 Department of Animal Behaviour, University of Bielefeld, Morgenbreede 45, 33615 Bielefeld, Germany 2 School of Environment and Life Sciences, Centre for Environmental Systems and Wildlife Research, University of Salford, M5 4WT Salford, United Kingdom Changing environments are the main driver of evolutionary changes. Corresponding shifts Sexual selection is expected to favour the evolution of large male body size when there is in the behaviour and life history traits of species might in turn also alter ecosystem attributes. direct male-male competition, and the development of male nuptial colouration in case of The reproduction of central European fire salamanders (Salamandra salamandra), in small female mate choice. In the moor frog (Rana arvalis), an explosively breeding species, males pond habitats instead of first order streams, is one example of a recent local adaptation. Since actively search and scramble for access to females during a short reproductive period. Males fire salamander larvae are important top-predators in these fish free habitats, their presence express a blue nuptial body colouration and some studies revealed a large male mating advan- changes various aspects of ecosystem functioning. Here we analysed how the ecological per- tage, but how these traits influence paternity success remains unknown. We performed formance of salamander larvae in ponds changes in comparison to stream populations. mating experiments in which we exposed three females to three small and large males each, Further we analysed how the presence of salamander larvae in ponds influences key ecosystem and measured male colour using a spectrometer. Four polymorphic microsatellite loci were attributes such as prey biodiversity and aquatic-terrestrial linkage. To assess the impact of used to determine the paternity of a subset of offspring, and to quantify genetic relatedness salamander larvae on the pond functioning, we combined detailed investigations of both between successful mating partners. Males fertilizing the majority of a female’s eggs were larger salamander larvae and invertebrates in ponds of a population in western Germany with ex- but not more intensely coloured than their competitors. As the males observed in amplexus not perimental manipulations of the salamander presence. Our calculations are based on biomass always achieved the highest siring success, and polyandry occurred in 67% of the analysed data of the pond fauna, as well as on the analysis of stomach content data, growth rates and egg clutches, we assume stray sperm or clutch piracy to cause multiple paternity. Successful population dynamics of the salamander larvae. We demonstrate that the larval behaviour and mating pairs were characterised by high genetic divergence, which was linked to the number performance can have a high interannual variability. The presence of salamander larvae dis- of offspring sired. tinctly influenced the macroinvertebrate community composition and food web structure in their aquatic habitat. Moreover we could show that the adaptation of fire salamanders to breed in pools leads to strong net increases of animal-mediated import of terrestrial matter into the aquatic habitats. The aquatic terrestrial subsidy exchange is altered through (1) a high net-biomass import of salamanders and (2) indirect biomass import through feeding on emerging insects and terrestrial prey.

160 161 BEHAVIOUR MONITORING

Tadpole transport logistics in the Neotropical frog Allobates femoralis Individual genetic tracking to assess amphibian dispersal: an island experiment in the Neotropical frog Allobates femoralis Eva Ringler 1, Andrius Pasˇukonis 2, Magdalena Erich 1, Walter Hödl 1, Max Ringler 1 Max Ringler, Walter Hödl, Eva Ringler 1 Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; [email protected] 2 Department of Cognitive Biology, University of Vienna, Althanstraße 14, 1090 Wien, Austria Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; [email protected] The evolutionary transition from aquatic to terrestrial egg deposition has enforced the devel- opment of various parental behaviours, in order to protect the eggs from external threats and The challenge to reliably mark amphibian larvae and to re-identify marked individuals after to ensure the final development of aquatic larvae. Poison-dart frogs (Dendrobatidae) have metamorphosis has hampered studies on larval and juvenile dispersal of amphibians for a evolved a remarkable variation in courtship and parental behaviours, and forms of parental long time. Current molecular methods allow establishing unique genetic fingerprints on an care are present in almost the entire family. Beside the knowledge that tadpole transport is individual basis. While these marker-based genotypes have been repeatedly used to establish obligatory in almost all dendrobatid frogs, very little is known about the underlying move- pedigrees and assess relatedness amongst individuals, they were not used in genetic mark- ment patterns and distribution strategies. We investigated the tadpole deposition behaviour recapture studies in amphibians so far. In the present study, in March 2012 we released 1800 in a natural population of Allobates femoralis in a lowland rainforest in the nature reserve “Les tadpoles of the Neotropical dendrobatid frog Allobates femoralis into 20 artificial pools on a Nouragues”, French Guiana, during five years from 2008 to 2012. We observed in total 129 5-ha river island in the Nature Reserve “Les Nouragues” in French Guiana, that was previously tadpole transport events. While in the vast majority of cases it were males (92.25%; n=119) uninhabited by this species. All tadpoles were genotyped at 15 highly variable microsatellite who were performing this task, we also observed 10 females (7.75%) with larvae on their loci with 19 alleles per locus on average, yielding unique genetic fingerprints for all individuals. back. Frogs carried on average 8.43 tadpoles on their back (range=1–25). Given the average In two recapture sessions, first we encountered 43 juvenile frogs (September–October 2012) clutch size of approximately 20 eggs, this indicates that frogs do not transport entire clutches and later 36 males and 31 females (January–March 2013) that had survived on the island. at once and/or that they distribute their larvae across several water bodies. When encountered Repeated genetic sampling and genotyping across all life history stages allowed us to match during tadpole transport, males were on average 38.58 m (range=1.64–185.12) away from their the microsatellite genotypes of tadpoles, juveniles, and adult frogs. The unique ventral pat- home territories. This distance significantly correlated with the number of tadpoles on the terns of A. femoralis thereby acted as a control for matched juveniles and adults. This enabled males’ back. For 114 tadpole-pairs, paternity was assigned to the male that was the respective us to reconstruct dispersal trajectories from the natal pool of tadpole development over juve- carrying individual, while for 5 pairs paternity was assigned to either neighbouring males or nile encounter locations to the final territory and perching sites of males and females. The unidentified individuals. For the 10 tadpole pairs that were transported by females, maternity same dataset of individually unique microsatellite genotypes will be used in further analyses was in all cases assigned to the transporting female. This study provides a comprehensive to assess factors that determine fitness, in terms of survival and reproductive success, in this analysis of movement patterns and discusses costs and benefits of different tadpole distribution semi-natural closed study population of A. femoralis. strategies in A. femoralis.

162 163 CLIMATE CHANGE DISTRIBUTION MODELLING

Learning from the past to predict the future: linking palaeophylographic Updating the distribution range of the narrow endemites models with future climate change risk assessments in Nearctic Salamandra atra aurorae and S. atra pasubiensis chelonians Enrico Romanazzi 1, Lucio Bonato 2 Dennis Rödder 1, A. Michelle Lawing 2,3, Morris Flecks 1, Faraham Ahmadzadeh 1, Johannes Dambach 1, Jan O. Engler 1, Jan Christian Habel 3, Timo Hartmann 1, 1 Department of Organismic Biology, University of Salzburg, Hellbrunnerstraße 34, 5020 Salzburg, Austria; [email protected] 1 1 1 1 2 David Hörnes , Flora Ihlow , Kathrin Schidelko , Darius Stiels , P. David Polly 2 Department of Biology, University of Padova, via Bassi 58b, 35131 Padova, Italy

1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; Salamandra atra aurorae and S. atra pasubiensis are limited to narrow areas along the Southern [email protected] Prealps, but the precise distribution of their population in known very incompletely. To 2 Department of Geological Sciences, Indiana University, Bloomington, Indiana, USA 3 Department of Biology, Indiana University, Bloomington, Indiana, USA improve knowledge, during 2012 we carried on 48 days of active exploration in putatively suitable areas, and gathered and evaluated occasional reports from people visiting the areas. Global climate is changing and species must respond by tracking suitable habitat, adapting Salamandra a. aurorae was recorded in the northern Altopiano dei Sette Comuni and to the changing conditions, or becoming extinct. To better understand how species will Vezzena, at altitude 1200–1800 m. Salamandra a. pasubiensis was confirmed with a single respond to future climate change, we have investigated the historic patterns of response to population in Vajo del Ponte–Val di Fontana d’Oro, 1450–1800 m, in the Pasubio Massif. fluctuating climates of the past 320 ka in North American chelonians. We further modeled the Undocumented reports of alpine salamanders from the Carega Massif remained unconfirmed. response to future climates of these species using different Intergovernmental Panel on Climate Change (IPCC) scenarios for 2080. Palaeophylogeographic models for the past 320 ka reveal different patterns of distributional changes in Nearctic chelonians, ranging from latitudinal range shifts and range contractions to a complete modeled ‘extinction’ during glacial maxima. Looking into the future, species distribution models projected on different IPCC climate change scenarios also reveal significant changes in future potential ranges resulting in range shifts. Especially in the South, large areas are facing completely novel climatic conditions from the species’ point of view, which might lead to increased environmental stress and even a high extinction risk. In combination of both approaches, this study shows that estimates on extinction risk pro- vided by assessments on prospective climate change have to proceed with caution, since even past climates lead to modeled ‘extinctions’ in certain species. This might happen for different reasons: (1) relevant processes are acting at much finer scale as typically assessed in macroeco- logical studies, and (2) other parts of the species’ fundamental niche which are not represented today become available during changing conditions, keeping the range still viable for the species.

164 165 PHYLOGEOGRAPHY AND PHYLOGENY POPULATION GENETICS

Molecular phylogeography of the long-nosed viper Conservation genetics tools for the spectacled salamander, (Vipera ammodytes) in the Cyclades, Greece Salamandrina perspicillata

Stephanos A. Roussos, Llewellyn D. Densmore III Valentina Rovelli 1, Ettore Randi 2, Francesca Davoli 2, Daniele Macale 3, Marco Alberto Bologna 1, Leonardo Vignoli 1 Department of Biological Sciences, Texas Tech University, MS 43131, Lubbock, Texas 79409-3131, USA; [email protected] 1 Dipartimento di Scienze, Università degli studi Roma Tre, Viale Guglielmo Marconi 446, 00146 Roma, Italy; [email protected] Dwarf populations of long-nosed vipers (Vipera ammodytes) are found on 15 islands of the 2 Istituto Superiore per la Protezione e la Ricerca Ambientale (ISPRA), Via Cà Fornacetta 9, Cyclades in the Aegean Sea. These populations were geographically isolated about 200 thousand 40064 Ozzano dell’Emilia, Bologna, Italy 3 Fondazione Bioparco di Roma, Viale del Giardino Zoologico 20, 00197 Roma, Italy years ago (KYA) when a large, ancient island (Cycladia) separated from mainland Greece because of rising sea-levels. Cycladia began to fragment into several islands about 20 KYA as The spectacled salamander (Salamandrina perspicillata) is an endemic species of peninsular the last glaciation of Europe was ending and the Aegean started to rise (120 m), leaving only Italy, listed as Least Concern by IUCN. Salamandrina perspicillata is widespread in the the higher elevations exposed creating the Cycladic archipelago. Recent molecular studies in Apennine Mountains but, due to its elusive habits and scattered distribution, more information the Cyclades have begun to uncover phylogeographic patterns and factors promoting diver- is needed to assess its actual conservation status. We tested the resolution power of 10 micro- sification of various herpetofauna, yet much is unknown of the biota in this Mediterranean satellite loci in order to verify their suitability as tool for conservation genetics studies, aiming biodiversity hotspot. Here we aimed to understand how isolation and fragmentation has and to estimate the genetic diversity within and among populations, as well as to investigate for continues to affect the genetic diversity and evolution of one of the top predators in the the first time patterns of paternity. We amplified the 10 loci in two multiplex PCR reactions Cyclades, V. ammodytes. Using phylogenetic analyses of mtDNA sequences we were able to (with five primer pairs each). Thereafter, we genotyped 129 adult individuals from Vejo unveil the extent of genetic diversity within the archipelago, evolutionary relationships to Regional Park (Latium, central Italy), collected between 2007 and 2012. Observed and expected mainland conspecifics as well as correlate those relationships to the complex paleogeography heterozygosity, deviations from Hardy-Weinberg equilibrium, probability of identity and of the region. Molecular data corroborated that extensive insular isolation of these viper pop- allele frequencies at each locus were calculated in GenAlEx 6.5. We found an average of 6.2 ulations has formed a divergent group/clade in the Cyclades that possess unique haplotypes alleles per locus. The values of observed and expected heterozygosity were Ho=0.617 and to almost each island. Our results also depict that two lineages are found within the clade He=0.622. All loci met the expectations of Hardy-Weinberg equilibrium, except one (Sala- that correspond to the initial split of Cycladia which created northern and southern islands. NC7), which was excluded. The probabilities of identity at 9 loci were PI=1.0E-07 and Additional support for this correlation is the clustering of haplotypes of individual islands PIsibs=1.03E-3. To test for the ability of the microsatellite markers to detect patterns of pater- within these northern and southern lineages as the islands continued to fragment. In conclusion, nity, we genotyped eight pregnant females (collected from their natural habitat) and their off- our results offer novel insights into the diversity that exists among Cycladic V. ammodytes and spring (278 larvae in total; 30 larvae on average for female). The reconstruction of paternal provides information for future taxonomic assessments and conservation management. genotypes was performed in Colony 2.0.3.5, revealing that each female mated with 3 males, on average (min=2, max=4). Our results showed that this set of markers can be efficiently used to estimate the extent of genetic diversity and to reveal patterns of multiple paternity in S. perspicillata.

166 167 SYSTEMATICS ECOLOGY

Morphology and advertisement call variation in the common tree frog Long-term dynamics of a Speleomantes strinatii population living Polypedates leucomystax complex in Thailand underground

Attapol Rujirawan, Anchalee Aowphol Sebastiano Salvidio 1,2, Fabrizio Oneto 1,2, Dario Ottonello 1, Mauro Valerio Pastorino 2

Department of Zoology, Faculty of Science, Kasetsart University, 10900 Bangkok, Thailand; [email protected] 1 DISTAV, Università di Genova, Corso Europa 26, 16132 Genova, Italy; [email protected] 2 Gruppo Speleologico Ligure “Arturo Issel”, Villa Comunale ex Borzino, Casella Postale 21, 16012 Busalla, Italy The common tree frog Polypedates leucomystax is widely distributed in Southeast Asia and has Long-term ecological studies are important because they produce data on slow evolving or been assumed to be species complex. We analyzed and described quantitative differences in complex ecological processes, such as climate change and prey-predator interactions. In this morphology and advertisement calls using univariate and multivariate statistics. We analyzed study, a population of the cave salamander Speleomantes strinatii was studied for 23 consecu- seven call parameters and twenty morphological characters from three localities in Thailand. tive years inside an artificial cave. In the period 1990–2012, salamanders active on the cave Two morphotypes of P. leucomystax were found in a southern population (Song Khla) as walls were counted in July to obtain a relative population index. Moreover, from 1996 this morphotypes A and B. These two morphotypes showed high difference in call types and call index was validated by absolute population abundance estimation obtained by temporary characters. Morphotype A has two call types whereas Morphotype B has three call types. The removals. The population time series was analysed by means of autoregressive moving average northeastern population has high call diversity with six different call types. Call properties of (ARMA) modelling. During the study, the salamander population showed large fluctuations, the northeastern population are distinct from southern populations. Discriminant function but its overall dynamics appeared constant. The selected ARMA (2,1) model indicated that analysis revealed three morphologically distinct groups and correctly classified 84.8% of indi- the salamander ecological dynamics was regulated by a complex interplay between delayed vidual to populations that is concordant with the bioacoustic results. Thus, our acoustic and density dependence and some external factor, possibly another species embedded in the same morphological data provide evidence for recognizing species within this complex in Thailand. food web. Moreover, this study confirmed that the cave salamander population showed weak evidence of periodicity, this phenomenon being rarely described in underground environments.

168 169 PRACTICAL CONSERVATION DISEASES AND PARASITES

Eleven years observation on the breeding season of the green turtle, Malformations in Neotropical viperids: qualitative and quantitative Chelonia mydas at Penang Island, Peninsular Malaysia and future analysis recommendations Sávio S. Sant’Anna 1,2, Kathleen F. Grego 2, Carla A. B. Lorigados 3, Ana Carolina B. C. Fonseca-Pinto 3, Marcelo P.N. Carvalho 1, Wilson Fernandes 2, Sarahaizad Mohd Salleh 1, Shahrul Anuar Mohd Sah 1, Mansor Yobe 2 Luís Carlos Sá-Rocha 1, José Luíz Catão-Dias 1 1 School of Biological Sciences, Universiti Sains Malaysia, 11800 Minden, Penang, Malaysia; [email protected] 1 Department of Pathology, School of Veterinary Medicine and Animal Sciences, University of São Paulo, 2 Penang Department of Fisheries, 11700 Gelugor, Penang, Malaysia 87 Prof. Dr. Orlando Marques de Paiva Avenue, 05508-270, São Paulo, Brazil; [email protected] 2 Laboratory of Herpetology, Butantan Institute, 1500 Vital Brasil Avenue, 05503-900, São Paulo, Brazil 3 Kerachut Beach and Kampi Beach are located on Penang Island, at the northern part of Department of Surgery, School of Veterinary Medicine and Animal Sciences, University of São Paulo, 87 Prof. Dr. Orlando Marques de Paiva Avenue, 05508-270, São Paulo, Brazil Peninsular Malaysia. The coordinates of Kerachut Beach are 5°27'4"N; 100°10'58"E and 5°26'20"N; 100°10'46"E for Kampi Beach. The Pantai Kerachut Turtle Conservation Centre Organogenesis, the most complex stage in embryonic development, is particularly sensitive was built in 1995 under the government organization, Penang Department of Fisheries. It to exogenous agents, being therefore an important period for the occurrence of malforma- represents the only turtle sanctuary on Penang Island, providing the information and as a tions, which may lead to newborn or premature death due to predation and/or diseases. This research unit. The objective is to study the pattern of breeding season during 11 years (2001– study describes the frequency and type of newborn malformations occurring in jararacas 2011). Two types of surveys were used to verify the green turtle nests which are; the inten- (Bothrops jararaca) and rattlesnakes (Crotalus durissus) at the Butantan Institute (IB), Brazil, sive nocturnal surveys (conducted at night from 1900–0600 h) and morning track counts from 2007 to 2013. Free-ranging pregnant specimens of these species delivered to IB were (a morning survey from 0900–0930 h). Result showed that the nests’ graph is fluctuated; kept in captivity until the birth of their offspring. Newborns were macroscopically and radio- the total nests detected per year ranged from 39 to 73 (mean=54.7, n=11) and the peak months logically analyzed for the presence of visible malformations. After natural death or euthanasia, are different every year. The total number of eggs collected during 11 years were 66,217 and complete necropsy was performed. Species B. jararaca showed a total of 95 malformed indi- ranged from 3985 to 7974 eggs per year. An independent-sample t-test was conducted and a viduals from 4087 births (2.3%). Species C. durissus showed 36 malformed snakes from 324 significant difference of the number of nest collected for both beaches was found. During our births (11.1%). The most common malformations were those associated with the spine, observation, the management of Pantai Kerachut Turtle Conservation Centre was able to whereas 74.8% (98/131) of snakes showed at least one malformation in the axial skeleton. The improvise more in order to upgrade it. We recommend to plant more Scaevola taccada trees in fusion of scales, leading to consolidation of body parts was found in 22.1% (29/131) of tera- order to attract turtles to land and to provide nesting sites; reducing and restricted lighting togenic snakes, and only two snakes, one of each species, showed fusion with no relation with from fishermen community and staff near the nesting beach; restricting visitors activities spinal deformities. There was no preferential side for eye malformation (microphthalmia, such as walking, fire camping, grouping near the beach; and fishermen activities should be anophthalmia and buphtalmos) which was observed in one or both eyes of the same snake. far away from beach due to trawling, fishing, lighting and engine gear that are disturbing Brachygnathia was observed in rattlesnakes and jararacas (2 and 9, respectively), as well as nesting activities. In conclusion, Penang Island has the least number of nest distribution upper lip malformation (2 and 1, respectively). All other congenital head malformations only compared to other states of Malaysia. occurred in jararacas. Three jararacas showed bicephaly. Schistosomia occurred in the tho- racic or abdominal region of teratogenic animals and, depending on the extent, the internal organs were exteriorized. Complementary studies are necessary to properly understand the etiology and significance of these malformations. Acknowledgments: FAPESP, CNPQ, INCTTOX.

170 171 ECOLOGY BEHAVIOUR

Back to black: the role of melanin in thermoregulation of The smell of a killer: what specific cues allow poison frogs to protect polymorphic lizards their offspring from cannibalistic conspecifics?

Stefano Scali 1, Marco Sannolo 1, Chiara Agata Giannoccari 1, Roberto Sacchi 2, Lisa M. Schulte Marco Mangiacotti 1 Department of Biogeography, University of Trier, Universitätsring 15, 54286 Trier, Germany; [email protected] 1 Museo di Storia Naturale di Milano, Corso Venezia 55, 20121 Milano, Italy; [email protected] 2 DSTA—Dipartimento di Scienze della Terra e dell’Ambiente, Università degli studi di Pavia, 27100 Pavia, Italy Chemical communication in amphibians is especially well documented in salamanders and newts, but several studies have also identified chemical communication in anurans. Here Many authors hypothesized that coloration and dorsal pattern could be important for reptile chemical communication between adult frogs and their tadpoles in the poison dart frog thermoregulation, because skin pigmentation causes significant differences in light reflectance Ranitomeya variabilis was investigated. As known for many poison dart frogs, the males of and some suggested that melanistic individuals could thermoregulate more efficiently. Podarcis this species deposit tadpoles singly into different phytotelmata. The exploitation of these muralis shows a dorsal polymorphism, with three different patterns, so the aim of our work small pools is advantageous as it lowers the risk of predation, but it is more costly because of was to test if different dorsal patterns were correlated to different thermal behaviours. We limited resource availability. Additionally, poison frog larvae are often cannibalistic, so the used two complementary approaches: a field study of a wild population (63 males and 30 identification and avoidance of conspecifics represents an adaptive behaviour for these females, ExpA) and an experimental study of heating rates in relation to dark pigmentation amphibians. (22 males and 29 females, ExpB). ExpA: all lizards were sexed, measured, photographed, marked By conducting pool-choice experiments in the Peruvian rainforest, it could be shown that with dorsal paintings and released;each time a marked lizard was seen, body temperature (Tb) parental R. variabilis are able to recognise the presence of conspecific tadpoles in phytotelmata was measured using an infrared thermometer and air temperature (Ta) was measured con- without invoking visual or physical stimuli, but are able to accurately determine the presence temporarily with a digital thermometer. ExpB: lizards were kept in terraria and heated pro- of tadpoles via chemical cues. Furthermore, it appears that these frogs can distinguish between gressively, measuring Tb and Ta at standard intervals with the same instruments. Data were different kinds of cues, produced by tadpoles of different species. While some tadpoles are analysed by means of a mixed model, using Tb as dependent variable, lizard identity as a ran- avoided as well (e. g. Hyloxalus azureiventris), others are not considered as dangerous for the dom factor to account for repeated measures, dorsal morph (reticulated, striped and mixed), own tadpoles (e. g. Rhinella poeppigii) or are even preferred (presumably recognized as a food SVL, sex, Ta and reproductive period (reproductive: April–June; post-reproductive: July– resource, e. g. Osteocephalus mimeticus). October) as fixed factors. Both experiments gave the same results, highlighting the depend- Current research is focused on the identification of the specific chemical cue(s) produced ence of Tb on Ta, but excluding any effect of other predictors. The lack of effects of dorsal by the tadpoles that elicit responses in adults. Cues of R. variabilis and H. azureiventris were patterns is particularly interesting, since it demonstrates that darker colorations do not con- extracted and fractionated and the fractions were tested by presenting them to the frogs in fer any advantage in thermoregulation. This result seems to confirm data collected for other the field. After one fraction of each species was found to be avoided, those fractions were frac- reptiles that underlined the effect of melanism on thermoregulation only in large species. tionated again into individual substances that are currently tested in the field, in order to find the cue that is responsible for the frogs’ reactions towards their tadpoles.

172 173 DISTRIBUTION MODELLING

NA2RE Project—The new atlas of amphibians and reptiles: The first Atlas of Amphibians and Reptiles in Europe was published in 1997, and since then, numerous regional and national organisations have collected new chorological data from new compilation and online system large parts of Europe. Furthermore, the taxonomic progress have contributed to multiple changes and new species descriptions. The Mapping Committee of the SEH presents here Neftalí Sillero 1, João Campos 1, Marco Amaro Oliveira 2, Luís Gonçalves-Seco 1,3, the New Atlas of Amphibians and Reptiles in Europe (NA2RE), a system of distributed data- Anna Bonardi 4, Claudia Corti 5, Raymond Creemers 6, Pierre-André Crochet 7, bases. Each distributed database has a national focus and is implemented in an online net- Jelka Crnobrnja-Isailovic´ 8,9, Mathieu Denoël 10, Gentile Francesco Ficetola 4, work, accessible through standard interfaces, thus sharing spatial-temporal data. A Web João Gonçalves 11, Sergei Kuzmin 12, Petros Lymberakis 13, Philip de Pous 14,15, interface facilitates the access to distributed data as if it was one virtual integrated database. Ariel Rodríguez 16, Roberto Sindaco 17, Jeroen Speybroeck 18, Pedro Sousa 3, Upon user request, the Web interface searches all distributed databases for the requested data, Fátima Sousa 3, Bert Toxopeus 19, David R. Vieites 20,21, Miguel Vences 16 integrating the answers in an interactive map. This methodology can easily actualise the records, as well as the taxonomy and systematics. Under this approach, data duplication is 1 Centro de Investigação em Ciências Geo-Espaciais, Universidade do Porto, Observatório Astronómico Prof. Manuel avoided and national databases are maintained in its own country. This system is implemented de Barros, Alameda do Monte da Virgem, 4430-146 Vila Nova de Gaia, Portugal; [email protected] using only open source software, for this purpose: (a) PostgreSQL database (with PostGIS 2 INESC Technology and Science (formerly INESC Porto), Rua Dr. Roberto Frias 378, 4200-465 Porto, Portugal extension to support spatial data and operations) to store observation data, (b) the Geoserver 3 Instituto Superior da Maia, Av. Carlos Oliveira Campos, 4475-690 Avioso S. Pedro, Portugal 4 Department of Earth and Environmental Sciences, Università di Milano-Bicocca, Piazza della Scienza 1, tool for accessing data, and (c) the OpenLayers framework to present dynamic and inter- 20126 Milano, Italy active maps in NA2RE web page (http://atlas.ismai.pt). The data on the NA2RE website is 5 Museo di Storia Naturale dell’Università di Firenze, Sezione di Zoologia “La Specola”, Via Romana 17, based in a compilation from different data sources, namely published (in books or websites) 50125 Firenze, Italia or ongoing national atlases, personal data kindly provided to the SEH, the 1997 European 6 RAVON, Postbus 1413, 6501BK Nijmegen, The Netherlands 7 Atlas, and the Global Information Facility. Databases were homogenised, projected to the CNRS-UMR5175 CEFE, Centre d’Ecologie Fonctionnelle et Evolutive, 1919 route de Mende, 34293 Montpellier, x France same coordinate system (WGS84) and transformed into a 50 50 km grid. Species richness 8 Department of Biology and Ecology, Faculty of Sciences and Mathematics University of Nisˇ, Visˇegradsˇka 33, maps were also calculated by the addition of all group species present in each square. The 18000 Nisˇ, Serbia obtained database comprises more than 380,000 records distributed across 40 countries. 9 Department of Evolutionary Biology, Institute for Biological Research “S. Stankovic´” University of Belgrade, Despota Stefana 142, 11000 Beograd, Serbia 10 F.R.S.—FNRS Research Associate, Behavioural Biology Unit, University of Liège, 22 Quai van Beneden, 4020 Liège, Belgium 11 CIBIO, University of Porto, R. Campo Alegre 687, 4169-007 Porto, Portugal 12 Institute of Ecology and Evolution, Russian Academy of Sciences, 117071 Moscow, Russia 13 Natural History Museum of Crete, University of Crete, Knossou Ave, P.O. Box 2208, 71409 Heraklion, Crete, Greece 14 Faculty of Life Sciences and Engineering, University of Lleida, Av. Rovira Roura 191, 25197 Lleida, Spain 15 Institut de Biologia Evolutiva (CSIC-UPF), Animal Phylogeny and Systematics, Passeig Marítim de la Barceloneta 37–49, 08003 Barcelona, Spain 16 Technische Universität Braunschweig, Division of Evolutionary Biology, Zoological Institute, Mendelssohnstraße 4, 38108 Braunschweig, Germany 17 Museo Civico di Storia Naturale, via San Francesco di Sales 88, 10022 Carmagnola, Italy 18 Research Institute for Nature and Forest, Kliniekstraat 25, 1070 Brussels, Belgium 19 University of Twente, Faculty of Geo-Information Science and Earth Observation (ITC), P.O. Box 217, 7500AA Enschede, The Netherlands 20 Museo Nacional de Ciencias Naturales, Consejo Superior de Investigaciones Científicas, José Gutiérrez Abascal 2, 28006 Madrid, Spain 21 REFER Biodiversity Chair, University of Porto, CIBIO, Campus Agrário de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal

174 175 DISEASES AND PARASITES POPULATION GENETICS

How evident are the co-evolutionary patterns between Phylogeography and population genetics of a Mediterranean tortoise haemogregarines and their turtle hosts? Testudo hermanni (Gmelin, 1789)

Pavel Siroky´ 1, Nela Dvoráková 1, Jana Kvicerová 2 Cäcilia Spitzweg, Melita Vamberger, Uwe Fritz

1 Department of Biology and Wildlife Diseases, Faculty of Veterinary Hygiene and Ecology, University of Veterinary Museum of Zoology (Museum für Tierkunde), Senckenberg Dresden, A. B. Meyer Building, 01109 Dresden, and Pharmaceutical Sciences Brno, Palackého tr. 1/3, 61242 Brno, Czech Republic; [email protected] Germany; [email protected] 2 Biology Centre, Institute of Parasitology, Academy of Sciences of the Czech Republic, v.v.i., Branisovská 31, 37005 Ceské Budejovice, Czech Republic Hermann’s tortoise, Testudo hermanni is a medium-sized, terrestrial species spread over the Mediterranean area of the Iberian, Apennine and Balkan Peninsulas. Until now the pop- Freshwater turtles belong to the most typical hosts of intracellular apicomplexan parasites of ulation differentiation within Testudo hermanni boettgeri is unclear and some authors have the genus Haemogregarina. The majority of Haemogregarina spp. were formerly described suggested that the western Croatian populations might represent a distinct taxon. Population based on the two criteria: (i) morphology of their gamonts in peripheral blood; and (ii) pre- differentiation is studied using 22 polymorphic microsatellite markers of more than 150 sumed strict host specificity, when each new studied host species denoted description of individuals from the Balkan Peninsula. a new parasite. Hundreds of Haemogregarina spp. described in this way thus represent a suit- able model for genealogical study and taxonomical revisions. Number of recent phylogenetic studies elucidates evolutionary relationships among turtle hosts and thus makes the model turtle-haemogregarine very attractive for studies on host-parasite co-evolution. To investigate the relevance of host specificity in phylogenetic/taxonomic framework, we examined haemo- gregarines from two groups of turtles. The first group contains 89 pelomedusid turtles, belonging to 10 species, namely 9 Pelomedusa subrufa, 25 Pelusios nanus, 14 P. upembae, 12 P. subniger, 9 P. marani, 6 P. rhodesianus, 6 P. gabonensis, 6 P. williamsi and one individual each of P. sinuatus and P. castaneus. These animals were freshly imported from Angola, Central African Republic, Democratic Republic of the Congo, Gabon, Kenya and Mozambique. The second group comprised 74 turtles from the western Palaearctic, namely 15 Emys orbicularis, 17 Mauremys caspica and 41 M. rivulata. All isolates were subjected to phylogenetic analyses of 1500 bp long sequences of 18S rDNA. Our results confirmed a low host specificity and considerable conspecificity, when branching pattern of trees did not follow host phylogeny. Moreover, studied Emys and Mauremys were parasitized by the same Haemogregarina. Evolution of haemogregarines thus probably did not follow evolution of vertebrate hosts, but evolution of their definitive hosts (leeches). This work was supported by the grant IGA VFU (11/2012/FVHE) and by the Grant Agency of the Czech Republic (P506/11/1738).

176 177 BEHAVIOUR PRACTICAL CONSERVATION

Take time to smell the frogs—Vocal sac glands of reed frogs contain Identifying conservation priorities for herpetofauna in Europe chemical cues on species identity Anton Stumpel, Ronald Zollinger Iris Starnberger 1, Dennis Poth 2, Stefan Schulz 2, Miguel Vences 3, Jette Knudsen 4, Michael F. Barej 5, Mark-Oliver Rödel 5, Manfred Walzl 1, Walter Hödl 1 Foundation RAVON, P.O. Box 1413, 6501BK Nijmegen, The Netherlands; [email protected]

1 University of Vienna, Department of Integrative Zoology, Althanstraße 14, 1090 Wien, Austria; SEH’s Conservation Committee assessed the status of European reptiles and amphibians in [email protected] the 1980s and categorized all species according to their degree of threat using defined criteria. 2 Technical University of Braunschweig, Institute for Organic Chemistry, Hagenring 30, 38106 Braunschweig, As a result, the most threatened species have been the subject of habitat studies and action Germany plans since. Generally, the outcome was published in reports to the Council of Europe and 3 Technical University of Braunschweig, Zoological Institute, Mendelssohnstraße 4, 38106 Braunschweig, Germany in many cases concrete management measures were taken in the field. 4 Department of Ecology, Lund University, 22362 Lund, Sweden 5 Museum für Naturkunde, Leibniz Institute for Research on Evolution and Biodiversity, Invalidenstraße 43, The status of many threatened species, habitats and herpetofaunal communities has changed 10115 Berlin, Germany (worsened) during the last decades by which an update is needed. Based on a new taxonomic species list, inquiries must be made in all European countries in order to identify at least the Males of many reed frog species (Anura: Hyperoliidae) have a prominent colorful gular patch most threatened species and habitats. This huge work can be seen as a continuation of the on their vocal sac, which is particularly conspicuous once the vocal sac is inflated. Although project ‘Important Herpetofaunal Areas in Europe’ that got stuck about ten years ago, but presence, shape and form of the gular patches are well-known diagnostic characters for these did by no means lose its relevance. After completion, a comparison can be made with the frogs, its function remained unknown. By integrating biochemical and histological methods, annexes of the Habitat Directive and of the Bern Convention, which were established using we found strong evidence that the gular patch is a gland producing volatile compounds, both ecological and political criteria. Conservation policy as well as practical conservation which might be emitted while calling. Volatile compounds were confirmed by gas chromato- management can benefit from an updated and possibly improved review. graphy-mass spectrometry in the gular glands in 11 species of the hyperoliid genera Afrixalus, We discuss the selection of criteria from today’s viewpoint and an outline is given of the Heterixalus, Hyperolius, and Phlyctimantis. Comparing the gular gland contents of 17 speci- thoughts how to run such a project. Also on behalf of the SEH Conservation Committee, mens of four sympatric Hyperolius species yielded a large variety of 65 compounds in species- collaboration is sought for this project. After having assessed the actual status of all species specific combinations. We suggest that reed frogs use a complex combination of at least and habitats, priorities can be set for actual and future conservation work. acoustic and chemical signals in species recognition and mate choice.

178 179 ECOLOGY PHYLOGEOGRAPHY AND PHYLOGENY

Impact of colour polymorphism in interaction with thermal Drang nach Süden: missing codons reveal unorthodox routes of conditions on reproductive success in asp viper (Vipera aspis) post-glacial expansion and secondary contacts in the fire-bellied toad, Bombina bombina Dubey Sylvain Jacek M. Szymura 1, Martin Sandera 2, Ufuk Bülbül 3, Nurs¸en Alpagut-Kerskin 4, Department of Ecology and Evolution, University of Lausanne, Switzerland; [email protected] Lukásˇ Choleva 5, Anna Wandycz 1, Sebastian Hofman 1 Ectotherms’ colouration plays a crucial role in many aspect of an animal life and can impact 1 Jagiellonian University, Kraków, Poland; [email protected] thermoregulation, metabolic pathway, animal behaviour and prey-predator interaction. 2 Polabské Muzeum, Podeˇbrady, Czech Republic Theoretically, in ectothermic animals the colouration is expected to have an important influ- 3 Karadeniz Technical University, Trabzon, Turkey ence on thermoregulation processes. Darker individuals should heat up faster and be able to 4 Biology Department, Ege University, Bornova-Izmir, Turkey reach and keep optimal temperature longer than would lighter individuals and therefore, 5 Institute of Animal Physiology & Genetics of the Academy of Sciences, Libechov, Czech Republic especially in cool areas, darker animals should exhibit longer activity periods, higher growth rate, survival, fecundity and reproductive output. The present study focused on the impact Dispersal from multiple glacial refugia resulted in a mosaic of parapatric clades in Bombina. of the dorsal colouration on the reproduction of asp viper (Vipera aspis) in interaction with In a lowland bound B. bombina AMOVA identified 3 geographic groups, the Northern, thermal conditions. In spring 2012, we collected a total of 24 gravid females of two distinct Southern and Anatolian, originating from presumed peri-Black Sea refugia. Upon glacial colour morphs (normal [light brownish-grey back colouration with a dark pattern] and retreat the N and S lineages experienced spectacular expansions to the West circumnavigating melanistic) from a colour polymorphic population of the Swiss Prealps. In order to test for the Carpathians and Sudety Mts. The two branches met in Central Bohemia, in a secondary an effect of colour phenotype (melanistic vs. normal) on the response of individuals to dif- contact, as revealed by lineage specific codon deletion in their divergent mtDNAs. Another ferent climatic conditions, these females were kept in captivity until parturition under two area of potential contact in European Turkey, where peripheral populations of the Southern treatments simulating “cold” versus “warm” conditions during their pregnancy (by modify- and Anatolian lineages occur in close proximity. Peculiar dispersal routes of the fire-bellied ing basking-time availability). The reproductive output of females (parturition time, clutch toad seem related to its specialized aquatic life style and post-glacial remodeling of the river size and mass) and the impact on the phenotype of neonates (body size, mass, dorsal and ventral systems resulting from global and local sea level rise. The toads may have colonized the colouration, and scalation) were analysed in relation to the temperature treatment and pheno- southernmost distribution margin in the Evros Delta only after influx of the saltwater into type of the mother. Our study revealed that females from the warm treatment gave birth on the lacustrine Black Sea from the Mediterranean. In contrast to widespread Northern and average earlier than did females from the cold treatment. Interestingly, melanistic females Southern mtDNA clades, the Anatolian clade seems restricted to the Sakarya River Valley gave birth on average earlier and produced more offspring than did normal females. As a con- and a few localities in European Turkey. Distinctness of these marginal populations, known clusion, our study demonstrated that colour polymorphism plays an important role in the as B. b. arifiyensis, deserves not only attention but quick conservation measures. ecology and evolution of the asp viper.

180 181 ECOLOGY PRACTICAL CONSERVATION

Age limit on plastic responses? Modelling plasticity in a species with Linking habitat suitability to demography in great crested newts complex life history (Triturus cristatus)

Zoltán Tóth 1, Josh Van Buskirk 2, Wieslaw Babik 3 Bianca Unglaub 1,2, Axel Drechsler 1, Benedikt Schmidt 3,4, Sebastian Steinfartz 1

1 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, 1 University of Bielefeld, Morgenbreede 45, 33615 Bielefeld, Germany; [email protected] Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] 2 University of Hamburg, Martin-Luther-King-Platz 3, 20146 Hamburg, Germany 2 Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Winterthurerstraße 190, 3 University of Zurich, Winterthurerstraße 190, 8057 Zurich, Switzerland 8057 Zürich, Switzerland 4 Karch, Passage Maximilien-de-Meuron 6, 2000 Neuchâtel, Switzerland 3 Molecular and Behavioural Ecology Group, Institute of Environmental Sciences, Jagiellonian University, Gronostajowa 7, 30-387 Kraków, Poland Understanding the relationship between population parameters and habitat suitability is cen- tral to monitoring, management and conservation of endangered species. Habitat suitability Various organisms can adjust their phenotype according to the environment they experience, index (HSI) models are often used to predict species occurrence and are valuable and con- i. e. exhibit phenotypic plasticity. Previous models suggest that plasticity is favoured by venient tools for wildlife management and conservation practice. The HSI is a numerical greater environmental variability and gene flow. However, temporal and spatial environmental index, ranging from 0 (unsuitable habitat) to 1.0 (optimal habitat). In the application of HSI challenges can be met with a variety of responses, and there is limited knowledge about which models for management purposes, habitat suitability is often used as a general indicator for traits are the most likely to become plastic and in which life stages. Previous theoretical works demographic responses. However, whether habitat suitability is actually related to demo- on the adaptive significance of phenotypic plasticity in species with complex life history focus graphic parameters such as survival and reproduction has rarely been evaluated. Great crested on predator-induced changes in larval traits or on optimal time for metamorphosis. In this newts (Triturus cristatus) are widespread in Europe but have suffered severe declines in many study we used an optimality model to investigate how plasticity in adult (affecting reproduc- countries within their natural range and are therefore strictly protected under the Habitats tion) and larval (affecting survival) stages contribute to the prevalence of a plastic phenotype Directive in the European Union. The HSI for great crested newts incorporates ten key habi- in a heterogeneous environment where two specialist phenotypes co-occur. We ran simulations tat variables, all of which are thought to have an effect on the species. Based on a capture- for 1000 generations and analyzed the change in the proportion of 9 phenotypes (all possible mark-recapture survey over three years, we evaluated the effects of habitat suitability on combinations of 3 larval and 3 adult strategies) in the larval and adult habitats. Generations apparent survival and reproduction for great crested newts, using model selection with were separate, and the proportions of the phenotypes at a site in each generation changed Akaike’s Information Criterion (AIC). In our study site in Krefeld, Germany, 24 ponds were according to their relative fitness. We found that phenotypes with adult and larval plasticity surveyed for presence and reproduction of great crested newts and 1919 individual capture were similarly successful in the heterogeneous environment, but specialists’ ability to choose histories were recorded. The candidate model sets for both demographic parameters included the right habitat for their offspring diminished the net advantage of plasticity. These results models with and without HSI values. Models containing HSI values received more support help explain why adult plasticity affecting reproductive success is less common than larval than intercept only models for reproduction but not for survival probability. Our results sug- plastic responses in species with complex life history. gest that the HSI can be a valuable predictor in conservation planning not only for occur- rence but also for reproduction probability in great crested newts.

182 183 ECOLOGY ECOLOGY

The adaptive value of predator-induced defences in Bufo bufo tadpoles Age and some growth parameters in two insular populations of Lyciasalamandra fazilae (Caudata: Salamandridae) from southwestern Bálint Üveges 1,2, Márk Szederkényi 1, Katharina Mahr 2, Herbert Hoi 2, Attila Hettyey 1 Turkey

1 “Lendület” Evolutionary Ecology Group, Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] Nazan Üzüm, Çiçek Gümüs¸ 2 Konrad Lorenz Institute of Ethology, University of Veterinary Medicine, Savoyenstraße 1A, 1160 Wien, Austria Adnan Menderes University, Faculty of Arts and Sciences, Department of Biology, Aydin, Turkey; [email protected] Phenotypic plasticity, the ability of a genotype to produce various phenotypes depending on the environment, is a well-studied phenomenon in anuran larvae. However, the adaptive In this study, we aim to describe some life-history traits (body size, age at maturity, longevity, value of such induced responses often remains unknown, especially so in species lacking spec- and growth) of Lyciasalamandra fazilae from two insular populations living in the Medi- tacular responses. It has been shown that tadpoles of the common toad (Bufo bufo) show re- terranean environment of southwestern Turkey. Samples were collected from the Tersane (30 latively weak morphological and behavioural responses to the presence of predators, while males, 30 females and 9 juveniles) and Domuz Islands (30 males, 30 females and 23 juveniles) probably relying on chemical defences. We conducted two experiments to evaluate if B. bufo located in the Göcek Bay. Body sizes of salamanders were measured with a plastic ruler and tadpoles exhibiting predator-induced phenotypes have elevated survival compared to naive the ages of juveniles and adults were estimated using phalangeal skeletochronology by counting conspecifics in the presence of various aquatic predators (Nepa sp., Aeshna cyanea, Lissotriton the number of resting lines (LAG) in cross-sections. The mean SVL of males and females was vulgaris, Esox lucius) and alternative prey (tadpoles of Rana temporaria). We reared tadpoles calculated as 65.92±0.80 mm and 65.30±1.40 mm on Tersane, and 62.47±0.62 mm and in outdoors mesocosms in the presence or absence of caged predators fed with tadpoles every 57.40±1.24 mm on Domuz Island, respectively. The mean SVL was significantly higher in other day. Three weeks after start we removed tadpoles from rearing tanks and exposed mixed females than in males on Domuz (p<0.05) whereas the sexes did not differ significantly on species tadpole assemblages to free ranging predators. We conducted bioassays in large and Tersane Island (p>0.05). This character was also significantly differed between the popula- in small plastic boxes. Large boxes contained leaf litter and aquatic vegetation and represented tions (p<0.05). In this study, one hundred and forty bones were scored and 92.10% were suit- natural conditions, where the full suite of induced defences was likely to affect survival prob- able for an estimation of the individual’s age. The mean age was 6.50±0.28 years in males, abilities. Small boxes only contained water, so that the effects of morphological and be- 6.69±0.37 years in females and 1.86±0.26 years in juveniles on Tersane Island. These values havioural defences could only have a little effect and variation in palatability may be assumed were recorded as 5.17±0.18 years, 4.31±0.22 years and 2.30±0.19 years for Domuz Island, to have played a major role in shaping survival probabilities. Induced tadpoles enjoyed ele- respectively. No significant differences were found between the age distributions of the two vated survival rates in both experimental venues compared to non-induced conspecifics. sexes in the Tersane population ( p>0.05) but there was a significant difference in the Domuz Furthermore, while predators varied in their effectiveness and degree of preference, they all population (p<0.05). The age structure of males and females varied significantly between pop- preferred Rana over Bufo. The latter result was more evident in small experimental boxes, ulations, with older mean ages to be observed on Tersane Island (p<0.05). Age at sexual where palatability was presumably more important. These results suggest that chemical maturity was determined as 4 years for males and 3 years for females in both populations. defences efficiently elevate the likelihood of survival in B. bufo tadpoles. Positive correlation between body size (SVL) and age was determined for both males and females in both populations. Growth was investigated according to the von Bertalanffy’s growth equation. The asymptotic size of males was lower (88.99±0.85 mm) than that of females

(101.16±1.11 mm) on Tersane, whereas the opposite was detected for Domuz Island (SVLasym(male)= 90.39±1.55 mm; SVLasym(female)=94.49±2.28). Our data show that males were bigger than females in both populations. There were significant differences between two insular L. fazilae popu- lations (Tersane and Domuz) for both sexes in terms of both SVL and age. This difference is thought to be resulted from the islands’ own ecological factors such as predation, aggres- sive competitors, food etc.

184 185 ECOLOGY POPULATION GENETICS

Reproductive interference between Rana dalmatina and R. temporaria Speciation under gene flow? The case of Mauremys caspica and affects reproductive success in natural populations M. rivulata

Balázs Vági 1,2, Tibor Kovács 2, Patrik Katona 3, János Ujszegi 1,2, Ákos Csillag 2, Melita Vamberger, Heiko Stuckas , Uwe Fritz Márk Szederkényi 1, Matteo Griggio 4, Herbert Hoi 4, Attila Hettyey 1,4 Museum of Zoology (Museum für Tierkunde), Senckenberg Dresden, A. B. Meyer Building, 01109 Dresden, Germany; 1 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, [email protected] Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] 2 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, The stripe-necked terrapins Mauremys caspica and M. rivulata are sister species occupying Pázmány Péter sétány 1C, 1117 Budapest, Hungary similar ecological niches and occurring in more or less mutually exclusive parapatric distri- 3 Department of Evolutionary Zoology and Human Biology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary bution ranges. The ranges of the two species are forming a narrow contact zone over hundreds 4 Konrad Lorenz Institute of Ethology, University of Veterinary Medicine, Vienna, Savoyenstraße 1A, 1160 Wien, of kilometres. We used 13 quickly evolving microsatellite loci and five nuclear genes for gain- Austria ing insights in gene flow and hybridization of 13 populations of M. caspica and 18 populations of M. rivulata from the contact zone and compared these to randomly chosen individuals from Experimental evidence suggests that reproductive interference between heterospecifics can the whole distribution range of each species. seriously affect individual fitness and population dynamics. In mating systems ruled by indis- criminate, coercive males, sexual isolation may collapse when the number of conspecifics drops below that of heterospecifics, leading to a decrease in the average reproductive success of individuals of the rarer species. Support from field studies for such an Allee effect on repro- ductive success has, however, remained scarce. The agile frog (Rana dalmatina) and the com- mon frog (Rana temporaria) are sympatric in the Pilis Hills, Hungary. Their breeding season overlaps and males of both species are often found in amplexus with heterospecific females, even though matings between heterospecifics produce no viable offspring. We monitored 25 breeding ponds during the breeding season. We estimated species abundance ratios based on the number of clutches laid by the two species and evaluated fertilization success in clutches. We assumed that zero or decreased fertilization success was caused by heterospecific males being involved in matings. We found that in populations where the species ratio was more biased towards R. temporaria, the average fertilization success of R. dalmatina eggs decreased, while fertilization success of R. temporaria did not decrease with increasing relative numbers of heterospecifics. This asymmetry in fitness-effects could be caused by the difference in the competitive ability of males of the two species: while males of the more competitive R. tem- poraria males are able to displace heterospecific males amplexing conspecific females, the smaller R. dalmatina males are not able to do so. Our results support the hypothesis that repro- ductive interference can lead to substantial fitness-loss and may affect population dynamics not only in experimental populations, but under natural conditions as well.

186 187 ECOLOGY SYSTEMATICS

The role of fire in structuring reptile communities in dry sclerophyll Big blues & small browns—systematics of African Agaminae forests in eastern Australia Philipp Wagner Diana Virkki 1, Guy Castley 2, Cuong Tran 3, Tom Lewis 4 Zoologisches Forschungsmuseum A. Koenig, Adenauerallee 160, 53113 Bonn, Germany, and Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; 1 Griffith School of Environment, Griffith University, Gold Coast Campus, Queensland 4222, Australia; [email protected] [email protected] 2 International Centre for Ecotourism Research, Griffith School of Environment, Griffith University, Gold Coast Campus, Queensland 4222, Australia The African agamas include several genera: Acanthocercus is a genus of mainly arboreal lizards 3 Transfield Services, 52 Merivale Street, South Brisbane, Queensland 4101, Australia widespread in savannas and woodland of eastern and southern Africa; Agama mainly includes 4 Agri-Science Queensland, Department of Agriculture, Fisheries and Forestry, Queensland Government, rupicolous and terrestrial lizards across the entire continent; Pseudotrapelus is only present University of the Sunshine Coast, Sippy Downs, Queensland 4456, Australia along the Red Sea coast, while Trapelus additionally occurs in northern Africa and the southern Sahara belt; and Xenagama is restricted to arid lowlands and mid-elevation at the Horn of Faunal responses to fire are generally less researched than for vegetation, but are no less Africa. For a long time the systematics of African agamas were poorly known but in the last important. This is particularly true for reptiles that are rarely studied in fire ecology, but likely years taxonomic research in these groups has intensified. to be impacted by fires due to their reliance on microhabitat structures. Previous studies In this talk I will present the new taxonomic concept of African Agaminae, based on the largely investigate local-scale effects in response to isolated fires, but few consider landscape results of several studies and I will show their relationships to other members of the subfamily. level spatio-temporal responses despite fire regime parameters, mainly frequency and extent, Moreover, I will demonstrate that this data can also be used to address more general questions. playing an important role in shaping landscape fire mosaics. Therefore, this study compared The African tropical rain forests harbour a huge number of reptile species and have been reptile assemblages in dry sclerophyll forests with differing fire mosaic histories in southeast well inventoried. On the other hand, the arid biomes of the continent are the worst explored Queensland to determine how reptiles respond to spatio-temporal fire regimes. Unique spatial habitats of the world and biodiversity and/or zoogeography is often examined on a defined fire mosaics were identified at two forests based on: total number of fires, mean fire interval geographic scale only. Therefore, agamid lizards were chosen as model group to get insights and time-since-fire. Reptile active searches were conducted at five plots within each fire his- into the zoogeographic “arid history” of the continent, into migration routes and colonization tory mosaic unit during two sampling periods, equating to 55 plots across the two forests. events, into early stage speciation processes and the evolution of a continent wide lizard group. Landscape and vegetation parameters were quantified at all plots. Results show that reptile abundance, richness and composition were influenced by fires. The best models revealed a negative response to the total number of fires, as well as fire type, where top disposal (forestry management) burns had higher species richness with a different composition to that of wild- fire and prescribed burns. Habitat variables were generally not strongly correlated with reptile parameters. However some fine-scale variables appeared within models alongside fire para- meters, where reptile abundance of common species was positively correlated with ground vegetation height, litter cover and log abundance and decay. The findings demonstrate a stronger relationship with fire as opposed to habitat structure, in contrast to previous work. This suggests that where land managers reduce the fire frequency, this may lead to a higher reptile abundance and diversity.

188 189 ECOLOGY ECOLOGY

Prey and microhabitat preferences of Natrix tessellata in a diverse Determinants of parapatric range limits in contact zones of European floodplain along the Hungarian Danube section at Göd (1669– salamanders 1671 rkm) Philine Werner András Weiperth 1, Tibor Gaebele 2, Imre Potyó 1, Gábor Guti 1, Miklós Puky 1 Trier University, Department of Biogeography, Universitätsring 15, 54296 Trier, Germany; [email protected] 1 MTA Centre for Ecological Research, Danube Research Institute, Jávorka Sándor u. 14, 2131 Göd, Hungary; [email protected] Abrupt range limits of parapatric species are interesting systems for studying the factors that 2 Szent István University, Faculty of Veterinary Science, István u. 2, 1078 Budapest, Hungary determine species’ range margins. Theory and empirical studies suggest that parapatric range limits are likely to form along strong environmental gradients when there is interspecific Natrix tessellata (Laurenti, 1768) is a diurnal piscivorous Eurasian snake species with a large competition. The parapatric salamanders Salamandra salamandra and S. atra meet in small distribution area. Along the River Danube it is present in the floodplain and along the lower contact zones in the European Alps and the maintenance of parapatry poses interesting ques- stretch of tributaries. The distribution of N. tessellata among the different microhabitats along tions to biologists. To identify determinants of their sharp range limits, this research com- the River Danube at Göd and its relationship with environmental parameters, e. g. current bines species distribution and habitat modelling. Macroecological models are used to explore velocity, and fish presence/absence, species composition, size distribution and abundance interspecific niche differentiation within contact zones and to assess the role of climate for was studied at the eight selected sections. Samples were collected twice a month from March the sharp range limits. To study patterns of local co-occurrence and to find evidence for inter- to October in 2008–2009, N. tessellata was found between May and early October. Altogether specific resource competition, species-habitat relationships are analysed using site-occupancy 26 N. tessellata sightings were recorded together with 497 individuals of 21 fish species along models. The results reveal a strong climatic gradient across range limits and show that the the same section, while another 5,547 individuals belonging to 28 species was caught in snake- salamanders generally occur at different conditions. The species show a moderate niche over- free areas. Microhabitat characteristics were more decisive than the presence of fish in deter- lap and conditions at species’ records vary among regions. On the habitat scale, the slope of mining along which section N. tessellata stays. It favoured slow-flowing sections with aquatic a site is important for S. salamandra, while none of the predictors explains the occurrence of macrophytes and a sandy or muddy bottom. In spite of their suitability, artificial habitats S. atra. There is no effect of species presence on each others’ site-occupancy. The results sug- were not much used may be due to increased risk of disturbance and predation/persecution. gest that the climate represents a determining factor for species’ range limits within contact The habitat preference of N. tessellata changed less between the two studied years with dif- zones. Interspecific niche overlap and the observation that both species can occur in a much ferent water level fluctuations than those of fishes and it was not identical with any of the wider range of climatic conditions and that no habitat predictor explains the occurrence of fish species caught. However, 15–30 mm and longer fish may influence the distribution of N. S. atra propose that interspecific interaction may play a role. However, there is no direct tessellata. evidence for resource competition. To fully understand the mechanisms that determine the parapatric range limits and to unravel the role of biotic interactions it is crucial to study the salamanders’ life history traits.

190 191 SYSTEMATICS ECOLOGY

Taxonomical developments in Triturus newts, driven by genetic data Anuran diversity of floating meadows in Amazonia

Ben Wielstra, Jan W. Arntzen Silas Wolf 1, Philipp Böning 1, Pablo J. Venegas 2, Stefan Lötters 1

Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands; [email protected] 1 University of Trier, Universitätsring 15, 54296 Trier, Germany; [email protected] 2 Centro de Ornitología y Biodiversidad (CORBIDI), Santa Rita 117, Huertos de San Antonio, Surco, Lima, Peru Genetic studies have revealed that the crested newt species ‘Triturus karelinii’ comprises more than one species. The first clue came from mitochondrial DNA: deep genetic divergence as The rivers of the Amazon basin harbor stationary formations of hydrophytes, known as already hinted at over twenty years ago, in one of Europe’s first phylogeographical surveys. ‘floating meadows’. These ecosystems are heterogeneously structured within and among Recently, a detailed mitochondrial DNA phylogeography revealed that T. karelinii comprises regions and underlay remarkable dynamics related from water level change. Biological diver- three mitochondrial DNA clades. A phylogeny based on full mitochondrial genome sequences sity is enormous, but little studied. Anuran amphibians exploit floating meadows for both showed these three clades to be as distinct as recognized Triturus species. Also ecologically foraging and reproduction. From the many questions arising from this, in this study we pro- speaking these mitochondrial DNA clades show differences comparable to those among real vide survey data (species richness) from four regions in the upper Amazon basin of Peru and species. To assess the validity of these results, an analysis of multiple nuclear DNA markers compare their species-turnover (beta diversity). Floating meadows were opportunistically sur- was required. A study employing allozymes confirmed genetic differentiation within T. karelinii veyed; perhaps not all anurans found actively chose floating meadows and rather ‘accidentally’ sensu lato, but sampling was too limited to allow firm conclusions. However, subsequently a occurred there (e. g. a few juvenile Rhinella marina). Species were identified on the basis of phylogeography of three nuclear DNA markers confirmed that T. karelinii sensu lato indeed morphology and advertisement calls (>25 taxa, with >85% of all species account for those in is composed of three species, as the mitochondrial DNA suggested. These three species are: the family Hylidae). Species richness was estimated using different incidence- and abundance- (1) an eastern species, covering the southern Caspian Sea shore, the Caucasus and the Crimea, based richness estimators (ACE, ICE, Chao1, Chao2, First-order jackknife). The taxon- (2) a central species, covering northern Asiatic Turkey, and (3) a western species, covering turnover was calculated using the Sørensen index, which was corrected by Jaccard similarity western Asiatic Turkey and the south-eastern Balkan Peninsula. The name T. karelinii should coefficient. Our results suggest that anuran species communities in floating meadows can be be restricted to the allopatric eastern species. The central and western species occur in para- remarkably different (likely related to structure and size) and that the similarity between two patry. We explain why the taxonomy of these two Triturus species is complicated and present floating meadows correlated with distance. an interim solution. Currently, only genetic data can be used to identify the three crested newt species comprising T. karelinii sensu lato. However, we hope our results stimulate what has been termed ‘reverse taxonomy’, i. e., the discovery of diagnostic morphological features for species initially identified based on genetic data only.

192 193 POPULATION GENETICS ECOLOGY

Distribution and genetic relationships among populations of Differential antipredator responses in two sympatric lacertid lizards the foam-nest treefrog Chiromantis hansenae (Cochran, 1927) in Thailand Anamarija Zagar 1,2, Katja Bitenc 2, Miguel A. Carretero 1, Al Vrezec 3

1 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Siriporn Yodthong, Anchalee Aowphol Vairão, 4485-661 Vairão, Portugal; [email protected] 2 Department of Biology, Biotechnical Faculty, University of Ljubljana, Vecna pot 111, 1000 Ljubljana, Slovenia Department of Zoology, Faculty of Science, Kasetsart University, 10900 Bangkok, Thailand; [email protected] 3 National Institute of Biology, Vecna pot 111, 1000 Ljubljana, Slovenia

The foam-nest treefrog, Chiromantis hansenae was previously reported for the geographic Syntopic populations of overall similar lacertid species are likely to share predators. However, distribution in northern, East-Central, and southeastern Thailand. Taxonomic status of predation pressures might be different. Lacertids are preyed by different predators with dif- Chiromantis hansenae has been unclear due to the high morphological similarity with con- ferent foraging strategies, namely avian predators attack from above, while terrestrial preda- generic species, C. vittatus. We investigated the distribution, and examined the genetic rela- tors attack by approaching on ground (in the open) or by searching for prey in their hiding tionship among C. hansenae populations in Thailand using partial sequences of mitochondrial places. Therefore lacertids are expected to employ different antipredatory behaviours, either 16S rRNA gene. Analyses of Maximum-likelihood (ML), and Bayesian methods revealed two crypsis or early predator detection (visually or chemically) followed by escape to shelter. distinct evolutionary lineages of C. hansenae in Thailand. Our results showed that C. hansenae We conducted an integrative evaluation of both the predator pressure and the antipredator Group I is distributed across northeastern, eastern, and southern Thailand whereas C. hansenae mechanisms for two morphologically and ecologically similar species: Iberolacerta horvathi Group II is distributed in northern and western Thailand. The mean pairwise distance and Podarcis muralis, occurring in syntopy in the Northern Dinaric region. We used three between Groups I, and II of C. hansenae was 4.7%. Our study suggested that the genetic dif- different approaches: (i) identifying and evaluating the predation pressure of avian predators ference between two groups of C. hanseane could be a result of geographic barrier. Further with plasticine models and automatic video cameras, (ii) experimental test of chemical recog- investigation at population level should be conducted by increasing genetic data and other nition of potential snake predators, and (iii) testing the refuge-escape tactics in the field. We evidence. have found very low predation pressure by avian predators. The recognition of snake chem- icals differed between species, I. horvathi, but not P. muralis, recognizing Coronella austriaca as a potential predator. Interestingly, none of the species responded to the scent of Vipera ammodytes. Adults of both species intrinsically differed in their escape behaviour when approached by the researcher simulating a ground predator attack. Specifically, I. horvathi tended to escape at greater distances and spent more time in the shelter before emerging (longer recovery time) regardless the air, surface or refuge temperatures and habitat type. Implications of these differential antipredator strategies for the coexistence of both species are discussed.

194 195 PRACTICAL CONSERVATION PHYLOGEOGRAPHY AND PHYLOGENY

Threats to the Moldavian meadow viper (Vipera ursinii moldavica) The puzzling phylogeny of the Vipera kaznakovi complex in eastern Romania Oleksandr Zinenko 1, Nikolaus Stümpel 2, Lyudmila Mazanaeva 3, Konstantin Shiryaev 4, 5 6 7 6 8 Stefan R. Zamfirescu 1, Alexandru Strugariu 1, Iulian Gherghel 2, Tiberiu C. Sahlean 3, Göran Nilson , Nikolai Orlov , Boris Tuniyev , Natalia Ananjeva , Robert Murphy , 9 Oana Zamfirescu 1 Ulrich Joger

1 1 Faculty of Biology, Alexandru Ioan Cuza University of Ias¸i, Bd. Carol I, 20A, 700505 Ias¸i, Romania; The Museum of Nature at V.N. Karazin Kharkiv National University, Trinkler St. 8, 61058 Kharkiv, Ukraine; [email protected] [email protected] 2 2 Department of Zoology, Oklahoma State University, 501 Life Sciences West, Stillwater, OK 74078, USA Staatliches Naturhistorisches Museum Braunschweig, Pockelstraße 10, 38106 Braunschweig, Germany 3 3 Faculty of Biology, University of Bucharest, Splaiul Independentei 91–95, 050095 Bucuresti, Romania Dagestan State University, Faculty of Biology, Department of Zoology, Apt. 13, M. Gadzhiyeva St. 37a, 367025 Makhachkala, Dagestan, Russia 4 Tula Regional Exotarium, Oktyabr’skaya St. 26, 300002 Tula, Russia One-fifth of the European reptile species are threatened because of fragmentation, degrada- 5 Göteborg Natural History Museum, Box 7283, 40235 Göteborg, Sweden tion, and loss of habitats. Vipera ursinii is probably the most threatened snake in Europe. In 6 Zoological Institute, Russian Academy of Science, Universitetskaya nab. 1, 199034 St. Petersburg, Russia eastern Romania V. u. moldavica inhabits steppe and deltaic habitats in Moldavia and the 7 Federal State Institution Sochi National Park, Moskovskaya St., 354000 Sochi, Russia Danube Delta, respectively, which are not free from anthropogenic alteration, despite legal 8 Centre for Biodiversity and Conservation Biology, Department of Natural History, Royal Ontario Museum, 100 Queen’s Park, Toronto, ON M5S 2C6, Canada protection. Human impact was documented tangentially in some studies and consequently, 9 Staatliches Naturhistorisches Museum Braunschweig, Gausstraße 22, 38106 Braunschweig, Germany our main objective is to inventory the anthropogenic threats and their potential harm to the Moldavian and deltaic viper populations, objective which is particularly important because We studied the phylogeny of vipers of the kaznakovi-complex using sequences of two mito- it would support future conservation efforts. Thus, we processed observations spanning chondrial genes, Cytb and COI, as well as the nuclear genes Bach1 and Rag1. Sampling cov- almost a decade and carried out in three Moldavian and three deltaic populations. Our pre- ered most of taxonomic diversity and distribution range of the complex in the Caucasian diction regarding the bioclimatic niche in the context of climate change scenarios indicates region (Vipera kaznakovi, V. dinniki, V. darevskii, V. orlovi, V. magnifica, V. pontica), including an extension of the species’ current range in eastern Romania. However, many potential habi- the type specimens of V. magnifica and V. pontica. Resulting trees reconstructed in MrBayes tats, although bioclimatically suited, might be improper due to profound anthropogenic and PhyML and TCS statistical parsimony networks show division of the complex into two alterations. In addition, the Moldavian viper populations are currently under anthropic pres- clades. One (Northern) is restricted to the territory of Krasnodar region in Russia and the sure which may lead to extinction regardless of the favourable climate or climate change. The range of the second (Southern) covers most of the territory to the south of the Main Caucasian human activities identified in the study areas are: grazing, mowing, fire, agriculture, roads, Ridge with some populations in the alpine zone of Northern Ossetia and Dagestan. Phylogeny constructions, leisure activities, afforestation, cynegetic species management, persecution, does not correspond to present taxonomy: the complex is not monophyletic and the Northern illegal collection, and littering. Human impact is stronger in one population of the Danube clade clusters with V. ursinii and V. renardi, while the Southern clade forms the sister lineage Delta than in Moldavia, but the deltaic population is larger and therefore probably more to the former clades. Closely related haplotypes of the Northern clade were found in all samples resistant to extinction. One steppe habitat (Moldavia) is threatened by urban sprawl in the from V. orlovi, V. magnifica and most of V. kaznakovi and V. dinniki from Russia. The Southern near future. The anthropogenic threats of the V. u. moldavica habitats and populations must clade further splits into lineages from Dagestan (alpine population of V. cf. dinniki or V. cf. be continuously monitored for identifying in time changes that may lead to population lotievi), the high altitude border region between Russia and Abkhazia (V. cf. dinniki), a high decrease and extinction. altitude population from Northern Ossetia (V. cf. dinniki or V. cf. lotievi), and the rest of high altitude and lowland populations of the V. kaznakovi complex from Georgia, Turkey and Armenia, including populations morphologically assigned to V. dinniki, V. kaznakovi, V. darevskii and V. pontica. Possible gene introgression has been identified by presence of hetero- zygous specimens in contact zones between Northern and Southern clades as well as between the Northern clade and the V. renardi complex. Vipera pontica being identical in mtDNA to V. kaznakovi from Turkey has nuclear alleles of V. kaznakovi and V. ammodytes, thus it rep- resents a hybrid between these two species.

196 197 ECOLOGY

Rounded or elongated body shape? Emys orbicularis hatchlings— juveniles display phenotypic plasticity when under different ecological conditions

Marco A. L. Zuffi, Giovanni D. Masucci

Museo di Storia Naturale e del Territorio, Università di Pisa, Via Roma 79, 56011 Calci, Italy; [email protected]

To test if habitat could shape carapace features starting from birth, in Emys orbicularis, we tested randomly chosen hatchlings from four clutches of a same locality, under two treat- ments, firm vs running water. After initial inertia and feeding stop during winter, youngster grew rapidly in March–May, then much more in June–September. On average, size and mass increased much more in river than in pond turtles. Biometric variables were all normally distributed (Komogorov-Smirnov test, all at P>0.05) (mother, treatment, season). The ratio carapace width/carapace length, was found as close to 1 (0.91-0.94; rounded) in pond Emys and was found lower (0.89–0.92; elongated) in river Emys. ANOVA for repeated measures (size=carapace width/carapace length, plastron width/plastron length and mass/

carapace length), multifactorial (mother, treatment, season) Fmother=35.996, P<0.0001, Ftreatment= 7.179, P=0.008, Fseason=5.157, P<=0.002; Finteraction mother*treatment=22.294, P<0.0001) gave sig- Poster Presentations nificant results. Newborns-youngsters Emys orbicularis reach a carapace shape on average rounded in pond treatment, and on average elongated in river treatment. The Shape-Function relationship starts at birth, and it seems adaptive, likely with a relatively low genetic inertia (this statement should be demonstrated).

198 PHYLOGEOGRAPHY PHYLOGEOGRAPHY

Inferring the effects of past climate fluctuations on the distribution Multiple dispersal out of Anatolia: biogeography and evolution of pattern of Iranolacerta (Reptilia, Lacertidae): evidence from Oriental green lizards mitochondrial DNA and species distribution models Faraham Ahmadzadeh 1,2, Morris Flecks 2, Dennis Rödder 2, Wolfgang Böhme 2, Faraham Ahmadzadeh 1,2, Miguel Angel Carretero 3, Dennis Rödder 2, Çetin Ilgaz 3, D. James Harris 4, Jan O. Engler 2,5, Nazan Üzüm 6, Miguel A. Carretero 4 David James Harris 3, Susana Nunes Freitas 3, Ana Perera 3, Wolfgang Böhme 2 1 Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti University, G. C., Evin, Tehran, Iran 1 Department of Biodiversity and Ecosystem Management, Environmental Sciences Research Institute, Shahid Beheshti 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; [email protected] University, G.C., Evin, Tehran, Iran; [email protected] 3 Dokuz Eylül University, Fauna and Flora Research and Application Center, 35150 Buca-Izmir, Turkey 2 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 4 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de 3 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Vairão, 4485-661 Vairão, Portugal Vairão PT-4485-661 Vairão, Portugal 5 Biogeography Department, Trier University, 54296 Trier, Germany 6 Adnan Menderes University, Faculty of Science and Arts, Department of Biology, 09010 Aydin, Turkey A variety of different methods are available for investigating the effects of past climate fluc- tuation on biota. Among them, molecular phylogeography and species distribution modeling The Oriental green lizards of the Lacerta trilineata group are widely distributed in Greece, approaches have been shown to be especially useful tools to trace past climate induced modi- Anatolia, the eastern Mediterranean, the southern Caucasus and the Zagros mountains in fications of species’ geographic distributions. The results of both analytical approaches are Iran. We studied their phylogeography using three mitochondrial markers with comprehensive here combined to better understand the influences of past climate changes on the fragmented sampling from most representatives of the group. Their phylogeny and divergence times distribution pattern of the lizard genus Iranolacerta that currently occurs in the northwest (implementing fossil-based molecular clock calibrations) were inferred using Bayesian methods, and central Zagros Mountains of Iran. While I. zagrosica is restricted to few localities and and haplotype networks were reconstructed to assess how genetic diversity and current dis- cannot be modeled, the potential distribution of I. brandti suggested a large region of unfa- tributional patterns were shaped. According to our phylogenetic analyses, the group consti- vorable habitat between northwest and the central Zagros Mountains populations. From the tutes a well-supported monophylum containing several distinct evolutionary lineages with phylogenetic perspective, both species display deep genetic separation likely predating the high haplotype diversity. Vicariance might explain the divergences within most lineages that Pleistocene. However, limited genetic diversity across this divided range of I. brandtii implies have accumulated by range restriction and expansion of populations due to Quarternary cli- that the current separation is recent. Deeper genetic divergences were, nevertheless, un- mate oscillations and glacial refugia. However, niche divergence seems to be a major force covered within the northwest population. Since no recent geological events explain these promoting speciation and large scale distributional patterns between lineages were shaped results, the most likely explanation for such a distribution pattern is due to past climate fluc- earlier by multiple, independent dispersals out of Anatolia during the Pliocene and early tuations. Both sources of evidence suggest that during the early Holocene the northwest and Pleistocene. Our results also suggest that the group is in need of a taxonomical revision, Zagros populations of I. brandtii were connected, and the current pattern was shaped with as identified lineages and genetic diversity are not congruent with the currently recognised their disconnection in the early Holocene. Further studies may identify potential glacial re- subspecies. fugia for other species in this biodiverse region.

200 201 ECOLOGY SYSTEMATICS

Symmetry of the cranium in island populations of Podarcis lizards Amietia angolensis and A. fuscigula (Anura: Pyxicephalidae) in analyzed with a high resolution 3D geometric morphometric approach southern Africa: hidden clades revealed

Raluca Ioana Ba˘ncila˘ 1,2,4, Ana Ivanovic 3, Jan W. Arntzen 4 Ninda Baptista 1, Rui Rebelo 1, Alan Channing 2

1 Institute of Speleology “Emil Racovit¸a˘”, 13 Septembrie Road, No 13, 050711 Bucharest, Romania 1 Centro de Biologia Ambiental, Departamento de Biologia Animal, Faculdade de Ciências da Universidade de 2 Faculty of Natural and Agricultural Sciences, University Ovidius Constant¸a, Aleea Universita˘¸iit nr. 1, corp B, Lisboa, Bloco C2, Campo Grande, 1749-016 Lisboa, Portugal; [email protected] 900527 Constant¸a, Romania 2 Biodiversity and Conservation Biology Department, University of the Western Cape, Private Bag X17, 7535 Bellville, 3 Institute of Zoology, Faculty of Biology, University of Belgrade, Studentski trg 16, 11000 Belgrade, Serbia South Africa 4 Netherlands Centre for Biodiversity Naturalis, Darwinweg 2, 2333CR Leiden, The Netherlands; [email protected] The taxonomy of the African amphibians is not totally understood. Since amphibians are considered an indicator of habitat quality and may play an important role in the flow of mat- We explored the influence of population isolation on shape and symmetry of the cranium ter and energy in the ecosystems, it is extremely important to know their systematics, in order in Podarcis lizards with CT-micro-scanning and 3D geometric morphometric techniques. to successfully address their effective conservation. A genetic-based study revealed that the 13 P. bocagei and 17 P. hispanica populations were studied from 13 island and 15 mainland two previously described species Amietia angolensis and Amietia fuscigula from southern Africa localities of the Ria de Arosa archipelago in northwestern Spain. The geological history were in fact composed by four different clades. In this work, we used advertisement calls and generated a natural laboratory with more or less isolated populations and thus allowed the morphology to evaluate this hypothesis. following predictions to be tested: small, random departures from symmetry (fluctuating Ninety-three advertisement calls, which are biphasic for this genus, were analyzed. Specimens asymmetry – FA) of the skull is higher (1) in island populations than in mainland popula- from museum collections and sampled in Angola, Lesotho, Namibia, South Africa and Zim- tions, (2) on remote islands than on islands close to the mainland, (3) on small islands babwe were analyzed for body measurements (44 adults) and coloration (61 adults and juve- than on large islands, and (4) in the resident species, P. hispanica than in the recent colo- niles). Discriminant Function Analyses were then performed on body ratios. nizer P. bocagei. Hypotheses 1–3 were partially confirmed, indicating that FA of the lizard Our data supports the hypothesis of four different clades. The integration of our data and cranium (as a measure of developmental stability and stress) is driven by population isolation. those from the genetics-based study allowed to propose a new division for these species, as We suggest that FA can be used in conservation biology as an early-warning system for con- follows: Amietia angolensis, known only from Angola, Amietia fuscigula, from the south-western ditions of stress. Cape in South Africa, Amietia quecketti, from South Africa, Lesotho and Zimbabwe, and Amietia poyntoni sp. nov. Channing & Baptista, 2013, from Namibia and South Africa. The advertisement calls were diagnostic for the four species, which could be distinguished by the duration of the first clicking phase: A. angolensis and A. quecketti have a mean dura- tion less than a second, sometimes reaching 1.2 s, A. fuscigula has a duration range between under 5 seconds and 14.3 seconds, and A. poyntoni sp. nov. has a duration range of 15.3–22.6 seconds. Amietia angolensis and A. quecketti can be distinguished by the duration and number of pulses in the second phase: A. angolensis has a mean duration under 0.2 seconds, and A. quecketti under 0.4 seconds. The DFA showed significant differences between A. fuscigula and A. poyntoni and also between A. angolensis and A. quecketti. Coloration pattern, namely the throat colour pattern showed differences between the clades, but there was some overlap between taxa.

202 203 PRACTICAL CONSERVATION ECOLOGY

Rearing conditions of tadpoles from the North African Call differences among brown frog species green water frog Pelophylax saharicus Carlo M. Biancardi 1, Anna Rita Di Cerbo 1, Edoardo Razzetti 2 Meher Bellakhal, Mouna Fartouna Bellakhal 1 Centro Studi Faunistica dei Vertebrati, Italian Society of Natural Sciences, C.so Venezia 55, 20121 Milano, Italy; [email protected] Exploitation of Aquatic Environments, High Institute of Fisheries and Aquaculture, Errimel, B. P. 15, 7080 Bizerta, 2 Museo di Storia Naturale, Università di Pavia, P.zza Botta, 9/10, 27100 Pavia, Italy Tunisia; [email protected] In anurans, different studies have shown that species sharing morphological characteristics Humans are the principal responsible of the decrease and the disappearance of some frog also show similar call characteristics. The following main features are used by females for species, not only by destruction of their natural habitats but also by their direct consumption. mate choice: call duration (CD), dominant frequency (DF) and pulses repetition rate (PRR). The development of frog legs market in many countries has stimulated research on frog Underwater calls of two close related species, Rana dalmatina and R. latastei, were recorded farming. Rearing conditions of the tadpoles of the North African green water frog Pelophylax in the same pond during one field session in Lombardy (Northern Italy), by means of a pro- saharicus are studied in order to assess the possibilities of intensive farming of this species. fessional hydrophone (Sea-Phone) and a digital recorder (M-Audio MicroTrack II), Audio The results showed that the temperature of 26 °C is optimal for a maximum weight gain files were analysed with the software Audacity and Sound Ruler. with a maximum rate of metamorphosis between 24 and 26 °C. The highest yields were The CD of R. dalmatina was variable, with a mean of 4.0 s (± 3 s SD). Each call was formed obtained at low densities from 1 to 10 tadpoles. The best survival and expedited metamor- by single impulses of 85.9 ±8ms, separated by an interval of 190 ±12ms; DF: 657 ±11Hz; phosis is obtained with a 35% protein food. The impact of protein level on food intake and PRR: 125 Hz. Rana latastei showed two different kind of call: one long “brum”, formed by weight increase is feeble. The maximum weight of tadpoles at metamorphosis is obtained several impulses, CD: 2.1 ± 0.2 s, interval 300 ± 40 ms; DF: 200 ± 27 Hz; PRR: 74 Hz. The with a rate of 40% protein. “meow” is a frequency modulated single call, CD: 551 ± 6 ms; DF: 783 ± 25 Hz; PRR: 64 Hz. Moreover we analysed two different calls of a Ligurian population of R. italica: one low inten- sity “grongron” (PRR: 58 Hz), and a higher “squack” (PRR: 330 Hz). The bioacoustics parameters of the calls of other 5 species of European brown frogs (R. temporaria, R. arvalis, R. graeca, R. iberica and R. pyrenaica) have been analysed, from refe- rence materials. The multiple comparisons (Pearson correlation index) of such parameters showed a substantial similarity among all the calls, with the exception of the “brum” call of R. latastei, which revealed negative correlation with almost all the other calls. Other species, such as R. iberica, R. italica and R. pyrenaica, emitted two different calls, but only one kind of brum-like call of R. iberica was similar to that of R. latastei (Pearson R = 0.552). These results can help to understand the role and the evolution of different calls in sym- patric close-related species.

204 205 ECOLOGY MORPHOLOGY, ANATOMY AND DEVELOPMENT

A preliminary study on predation pressure using plasticine models and Comparison of skull form in the snake genus Natrix automatic video camera traps in allotopic and syntopic populations of (Squamata: Serpentes: Natricinae) two lizards Bartosz Borczyk, Martyna Gorynia Katja Bitenc 1, Anamarija Zagar 1,2, Miguel A. Carretero 2, Al Vrezec 3 Department of Evolutionary Biology and Conservation of Vertebrates, University of Wroclaw, ul. Sienkiewicza 21, 50-335 Wroclaw, Poland; [email protected] 1 Department of Biology, Biotechnical Faculty, University of Ljubljana, Vecˇna pot 111, 1000 Ljubljana, Slovenia; [email protected] 2 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de Although snake skull structure attracts serious interest of researchers, quantitative studies on Vairão, 4485-661 Vairão, Portugal closely related species are relatively rare. Presented here is an analysis of 20 measurements 3 National Institute of Biology, Vecˇna pot 111, 1000 Ljubljana, Slovenia of skull dimensions of members of four Natrix species: Natrix natrix (N=19), N. tessellata (N=23), N. maura (N=6) and N. megalocephala (N=2).Because the latter is represented only Podarcis muralis and Iberolacerta horvathi are small, ecologically and morphologically similar by two specimens, all analyses were repeated including and excluding this species. Besides lacertid lizards who can be found living in sympatry in southern parts of Slovenia. Regionally, comparison of skull shape another test has been conducted to find if aquatic specialization the only substantial difference found thus far between the two species was the exhibited dif- and piscivory of tessellata and maura and terrestrial habitats on anurophagous natrix and ference in altitude of their preferred habitats. In order to try to recognise the mechanisms megalocephala are reflected in the mechanical advantage (MA) of snake jaw (ratio of the in- behind this kind of segregation we chose to research the effect of predators on these two lever to out-lever). Principal component analysis and discriminant function analysis enable species. In some systems, predation pressure can promote coexistence of prey species, while clear distinction of natrix, tessellata and maura specimens; megalocephala when considered is low predation pressure may be the driving force behind stronger expression of competitive placed within natrix morphospace. The characters being most important for discrimination interactions. The species could both be preyed on by other reptiles, birds and also mammals. between these species are skull height and jaw length, which may reflect adaptation to different Our choice for determining the strength or presence of avian and mammalian predator pres- habitat (aquatic vs. terrestrial). The results show that tessellata and maura have slightly lower sure in syntopic and allotropic locations were plasticine models of lizards, which we set up in MA compared to natrix, which is congruent with the general trend observed in aquatic, fish- groups of 100 in nine localities for one or two days, considering the microhabitat use of both eating snakes. species. Additional measures for detection of predators were provided in the forms of IR night vision automatic video camera (U-Way), focused on five plasticine models on exposed surfaces for the duration of the entire week. At the same time we also recorded presence of avian predators on all of the nine localities and estimated lizard relative abundance, along with collecting mammal scats to investigate the presence of lacertids in their diet. Preliminary results of these experiments will be presented with first analyses and photographs of predation events taken during video recording. Plasticine models had marks of small mammals on them and we recorded a beech marten (Martes foina) appearing to be eating one. Avian predators which we most frequently encountered on localities were raven (Corvus corax) and common buzzard (Buteo buteo).

206 207 ECOLOGY BEHAVIOUR

Reproduction and cold hardiness of the eggs of Siberian wood frog, Different effect of thermal and humidity conditions on duration of Rana amurensis in the North-East habitat chemical signals of two sympatric and sister lizard species

N. A. Bulakhova 1,2, O. N. Meshcheryakova 1, D. I. Berman 1 Carlos Cabido 1, José Martín 2

1 Institute of Biological Problems of the North, Far East Branch, Russian Academy of Sciences, Portovay St. 18, 1 Aranzadi Society of Sciencies, Alto de Zorroaga 11, 20014 Donostia, Spain; [email protected] 685000 Magadan, Russia; [email protected] 2 National Museum of Natural Sciences, Spanish Council for Scientific Research (CSIC), José Gutiérrez Abascal 2, 2 Research Institute of Biology and Biophysics, Tomsk State University, Lenina St. 36, 634050 Tomsk, Russia 28006 Madrid, Spain

Time periods and places of spawning, fecundity and cold hardiness of the eggs of Rana The sensory drive hypothesis states that there is natural selection for signals to effectively amurensis Boulenger, 1886, which is the northernmost in Asia Anuran species in two pop- stimulate the receiver sensory system and that the nature of the most effective signal design ulations of the Magadan region, located in the valley of the Tauy and El’gen rivers (59°45'N, depends on environment characteristics. Male lizards of many species possess holocrine epi- 148°32'E and 62°50'N, 152°22'E), were studied. The frogs spawned in shoal of standing and dermal glands that, during the breeding season, deliver secretions used in intraspecific com- weakly running oxbow and thermokarst ponds, often at a distance of hundreds of meters munication. Effective chemical communication may be also influenced by the environment away from places of wintering. Egg layings (up to 20 in one site) are disposed singly without and different habitats may promote chemical signal divergence even in closed related species. forming communal surface rafts. At the end of May, 2012, in the valley of the El’gen River The aim of the present study is to experimentally explore whether temperature and humidity embryogenesis came to the end, while near the coast of the Sea of Okhotsk spawning only may determine the duration of male chemical signals of two sympatric and closely related began. Number of eggs in one laying was 1131 ± 93.8 (n=21; 601–2152); three from 105 analysed lizard species occupying different microhabitats: Podarcis bocagei (ground-dwelling) and layings were unfertilized. Podarcis hispanica (rock-dwelling). We expected that selection would favour chemical signal Surviving, supercooling points (SCP) and dehydration in process of freezing of the whole adaptation to each environment (physical and/or social). Tongue-flick behaviour (TF) reflects laying, and SCP of single eggs, their jelly capsules and naked embryos, were analyzed. Stay chemical investigation of a stimulus and is often used as a quantitative bioassay of detection of the whole laying is icebound results in freezing and death of the periphery eggs; those, of chemical cues in experiments addressing chemosensory discrimination in lizards. We located in the center stay unfrozen and finish their development under temperature increase. examined signal detectability and persistence (time that signals remain detectable for the Laying eggs outside water (when the pool is dried up) freeze without supercooling between receivers) using TF rates of males to other male chemical signals that were exposed during 1, –0.2 °C and 0.5 °C, their mass halves due to dehydration. The eggs freeze with supercooling. 24 or 48 hours to normal temperature (26 °C; average temperature during reproduction sea- There is a slight difference between minimum SCP of unfertilized (–5.6 °C) and fertilized son in the capture area) or to 5 °C of increased temperature; and to an humidity of 60 or eggs (–6.2 °C). Mean SCP of the eggs increase from 15–18 stages to the end of embryogenesis 85%. We found that both, temperature and humidity, affect differentially to signalling per- from –4.5 ±0.1 °C to –3.2 ±0.2 °C. Jelly capsules have higher SCP (up to –5.3 °C), in com- formance of each species. That is, higher temperature after 24 h and higher humidity after 1 h parison with embryos of any state of development (up to –8.5 °C) or early larvae (up to –7.9 °C). of exposure reduces detectability of P. hispanica male signals. Between species differences in Different layings of one state of development have mean SCP of the eggs differed by 1.3 °C, mating system may explain why P. bocagei chemical signals seems to be more stable than P. of their jelly capsules or naked embryos by 0.2 °C. hispanica ones. Obtained characteristics of the eggs’ cold hardiness and its variation tendencies are similar to those, of syntopic Salamandrella keyserlingii and North American Rana sylvatica. The eggs of these species have ability to supercooling, that is not inherent to the whole laying as evo- lutionary secondary structure.

208 209 MONITORING PRACTICAL CONSERVATION

A preliminary checklist of lizards of Qatar Status of amphibian breeding sites in Derdap National Park, Serbia

Aurora M. Castilla 1,2, Aitor Valdeón 3,4, Dan Coga˘lniceanu 5, Alberto Gosá 3, Jelka Crnobrnja-Isailovic´ 1,2, Dragana Milojkovic´ 3, Marija Ilic´ 2, Biljana Macura 4,5 Paloma Mas 6, Ali Alkuwary 7, Essam O. H. Saifelnasr 8,9, Noora Al-Jaidah 1, Renee Richer 10 1 Faculty of Sciences and Mathematics, University of Nisˇ, Visˇegradsˇka 33, 18000 Nisˇ, Serbia; [email protected] 2 Institute for Biological Research “S. Stankovic´”, University of Belgrade, Despota Stefana 142, 11000 Belgrade, Serbia 1 Department of Biodiversity, Qatar Environment and Energy Research Institute (QEERI), Qatar Foundation, 3 Public Enterprise National Park “Derdap”, Kralja Petra I 14a, 19220 Donji Milanovac, Serbia Education City, P.O. Box 5825, Doha, Qatar 4 Department of Land, Environment, Agriculture and Forestry, University of Padova, 35020 Agripolis-Legnaro (PD), 2 Forest Sciences Centre of Catalonia (CTFC), Spanish National Research Council (CSIC), Spain Italy 3 Department of Herpetology, Aranzadi Society of Sciences, Zorroagagaina 11, San Sebastián, Spain; 5 Centre for Evidence-based Conservation, School of Environment, Natural Resources and Geography, [email protected] Bangor University, Gwynned LL57 2UW, United Kingdom 4 Department of Geography and Regional Planning. University of Zaragoza. Pedro Cerbuna 12, Zaragoza, Spain 5 University Ovidius Constant¸a, Faculty of Natural Sciences and Agricultural Sciences, Aleea Universita˘¸iit 1, corp B, 900740 Constant¸a, Romania Derdap National Park is an internationally recognized protected area because of its diverse 6 Department of Biodiversity and Evolutionary Biology, National Museum of Natural Sciences, habitats, geological structures, archaeological sites and beautiful landscapes. Twelve amphibian Spanish National Research Council (CSIC), José Gutiérrez Abascal 2, 28006 Madrid, Spain species have been recorded there and additional four species were listed as potentially occur- 7 Wildlife Research Section, Ministry of Environment, P.O. Box 7635, Doha, Qatar ring (16 in total), which represents 76% of the overall amphibian species diversity in Serbia. 8 Genetic Resources Department, Biotechnology Centre, Ministry of Environment, P.O. Box 200022, Doha, Qatar 9 Agricultural Research Center (ARC), Ministry of Agriculture, Egypt In a sample of 30 breeding sites, the commonest amphibian species in the area seems to be 10 Weill Cornell Medical College, Qatar Foundation, Education City, P.O. Box 5825, Doha, Qatar Rana dalmatina (found in 50% of registered reproductive sites). Pelophylax esculentus complex (47%), Bufo bufo (37%) and Salamandra salamandra (30%) are also common Salamandra Qatar is a peninsula projecting from the Arabian mainland 186 km in the Arabian Gulf and salamandra prefers water bodies far from human settlements (p<0.01) in shady places (p<0.05) sharing 85 km terrestrial borders with the kingdom of Saudi Arabia. Qatar terrestrial territory with low amount of submersed vegetation (p<0.05). Pelophylax esculentus complex choose covers 11,180 km2 and is represented by a relatively flat desert with ca. 100 m a. s. l. Its climate more open landscape (p<0.05), and ponds rich in submersed vegetation (p<0.05). Both species is hot and arid with an average annual mean temperature of 27 ºC and 75 mm rainfall per are avoiding heavily exploited water bodies (p<0.05). Rana dalmatina prefers temporary stag- year. Diversity of habitats is little and few natural resources are available other than abundant nant waters (p<0.01) with less or no flotant vegetation (p<0.05). gas and oil reserves. Overall the harsh desert conditions, the relatively uniform habitat, The highest number of amphibian species occurred in proximity of the entrance of smaller the increasing industrialization and growing human impact have contributed to sparsely and rivers into the Danube. Main local factors of threat are loss of traditional agriculture and locally disseminated lizard populations usually with low individual densities. Very few her- alteration of breeding sites into fish ponds. Establishment of amphibians-friendly artificial petological surveys have been conducted till now and very little is known about the distribu- ponds should be a priority in sustainable management. tion and the ecology of the lizard species present in Qatar. A recent survey and inventory study of lizards has confirmed the presence of 21 lizard species. The only neighbouring coun- try is Saudi Arabia that hosts over 30 lizard species, and some of which might also exist in Qatar. The most abundant family found in Qatar is Gekkonidae with nine species (Bunopus tuberculatus, Cyrtopodion scabrum, Hemidactylus robustus, H. flaviviridis, H. persicus, Steno- dactylus arabicus, S. slevini, S. doriae, Pseudoceramodactylus khobarensis), followed by Agamidae with three species (Trapelus flavimaculatus, Uromastyx aegyptia, Phrynocephalus arabicus), Lacertidae with four species (Acanthodactylus schmidti, A. opheodurus, Mesalina brevirostris, M. adramitana), Scincidae with two species (Scincus mitranus, Trachylepis septemtaeniata), and Varanidae (Varanus griseus), Sphaerodactylidae (Pristurus rupestris) and Trogonophidae (Diplometopon zarudnyi) with only one species each. Our results will help to understand the terrestrial biodiversity in Qatar, and will lead to establishing better management and conser- vation activities.

210 211 ECOLOGY PHYSIOLOGY

Population dynamics and habitat use of two sympatric newt species, Cellular bases of body size variance in the fence lizard Triturus dobrogicus and Lissotriton vulgaris at the breeding site Sceloporus undulatus (Phrynosomatidae) from different latitudes

Gábor Deák 1, Péter Sály 2, István Kiss 1 Natalia Derus, Marcin Czarnoleski

1 Szent István University, Department of Zoology and Animal Ecology, Páter K. u. 1, 2100 Gödöllo˝, Hungary; Jagiellonian University, Institute of Environmental Sciences, Life History Evolution Group, Gronostajowa 7, [email protected] 30-387 Kraków, Poland; [email protected] 2 HAS Centre for Ecological Research Balaton Limnological Institute, Klebelsberg Kunó u. 3, 8237 Tihany, Hungary No one knows why many species of ectotherms and endotherms attain larger body sizes at Newt populations were investigated in two small water bodies (a pond and a drainage ditch) higher latitudes. Mammals, birds and turtles usually follow this so-called Bergmann’s rule, in 2010. Newts were caught by funnel traps on nine sampling occassions during the breeding but not lizards and snakes. In contrast to general tendencies of lizards, in the North American season, and were identified individually by their belly patterns. The relation of abundances lizard Sceloporus undulatus individuals from northern populations are typically larger than and habitat characters was analyzed by a Generalized Linear Models (GLM). In total, 149 individuals from phylogenetically related populations from the south. Our study evaluates smooth newts (L. vulgaris) and 94 Danube crested newts (T. dobrogicus) were caught in the how differences in body size between four geographically distant populations of S. undulatus pond, while 42 smooth newt and 74 crested newts were collected in the drainage ditch. The (from Nebraska, Florida, Indiana and Texas) link to changes in the size of cells in two tissue number of L. vulgaris females was higher than that of T. dobrogicus at both breeding sites. types—red blood cells and skeletal muscles. We discuss results in the context of emerging Individuals arrived earlier, usually spent longer time at the breeding site. First and second theory on changes in the adaptive value of cell size across thermal environments. Under- recapture rates (RR) were quite similar for the two species in the pond habitat (L. vulgaris standing how and why cell size changes with latitude in S. undulatus can help future work first RR 20%, second RR 4.1%; T. dobrogicus first RR 19.7%, second RR 5.6%). On the con- that aims at finding ultimate explanations of Bergmann’s clines in ectotherms. trary, RR were substantially smaller and differed remarkably between species in the drainage The work is supported by the National Science Centre (grant N N304 373238). ditch (L. vulgaris first RR 13.5%, second RR 0%; T. dobrogicus first RR 4.6%, second RR 3.0%). The average distance of moving among the traps was quite similar for both species: 9.2 m (L. vulgaris) and 10.8 m (T. dobrogicus) in the pond, and 12.2 m (L. vulgaris) and 10.0 m (T. dobrogicus) in the ditch. The abundance of smooth newt showed a positive association with duckweed and reed-mace covering alike. This finding suggests that the meso scale habitat heterogenity is favourable for this species. The abundance of Danube crested newt showed only a weak positive association with the open water surface. Our results demonstrate that population dynamics of newt species can substantially vary depending on the type of the breeding habitat still within a single breeding season. Considering the sharp contrast between climates of subsequent years, the small, temporary aquatic habitats should receive greater importance in the conservation of newts, especially in a fragmented landscape.

212 213 MORPHOLOGY, ANATOMY AND DEVELOPMENT PHYLOGEOGRAPHY

How accurate do 2D geometric morphometrics quantify shell shape Late Pleistocene herpetofauna from the Baranica Cave (Serbia) in chelonians? D. Djuric´ 1, K. Bogic´evic´ 2 Marko Djurakic 1, Anthony Herrel 2, Anne-Claire Fabre 2, Raphael Cornette 2 1 Natural History Museum, Njegosˇeva 51, 11000 Belgrade, Serbia; [email protected] 2 Department of Paleontology, University of Belgrade, Faculty of Mining and Geology, Kamenicka 6, P.O.Box 227, 1 Department of Biology and Ecology, Faculty of Sciences, University of Novi Sad, Trg Dositeja Obradovica 2, 11000 Belgrade, Serbia 21000 Novi Sad, Serbia; [email protected] 2 UMR 7179, CNRS/MNHN, 55 rue Buffon, 75005 Paris, France Baranica cave is located in the southeastern part of Serbia, about four kilometers south of Among the 330 recognized extant species of turtles and tortoises, near 50% are listed as Knjazevac. Archaeological research of this cave was carried out in period 1994–1997 (Faculty “Threatened” which makes them among the most endangered of the major groups of verte- of Philosophy from Belgrade and the National Museum of Knjazevac). A large amount of brates. A key challenge for conservation plans is to suppress threats that may reduce the fit- fossil vertebrate remains was collected during the archeological excavation. The fossil mate- ness of a population. For that reason it is crucial to identify straightforward and reliable rial includes remains of large and small mammals prevailing over the other vertebrates (fish, methods that can recognize important units for conservation. Up to date, many chelonian amphibians, reptiles and birds). The revealed profile has four visible layers while layers 2 and studies have analyzed shell shape variation from the conservation point of view and its asso- 4 showed a significant wealth of fossil remains. The age of these layers was determined as ciation with other variables (physiological, historical, and environmental) as well. The shell Upper Pleistocene by absolute dating of bones of large mammals (23,520 ± 110 B. P. for Layer shape is usually quantified by traditional morphometrics and, less commonly, using geo- 2 and 35,780 ± 320 B. P. for Layer 4). This paper presents the herpetofauna collected in exca- metric morphometrics (GM). Despite the recognized advantage of both 2D and 3D GM over vations at Baranica I in 1995. The collected material is partially damaged (broken) and traditional morphometrics in characterizing shapes, their utilization is limited to museum includes isolated bones of representative amphibians and reptiles, while the most numerous collections or dead specimens. However, long-term comparative field studies on free-living items are snake vertebrae. Following taxa were identified in analysis of remains: Natrix natrix, specimens are a key for conservation research and the use of GM would be beneficial. Here Vipera berus, Elaphe longissima, Vipera ammodytes, Lacerta sp., Rana cf. ridibunda. we compared how precise and accurate 2D GM quantifies shell shape and proposed its uti- lization for field studies. We analyzed carapaces (vertebral and costal plates) and plastrons of 21 specimens belonging to five chelonian species using both 2D and 3D GM. 2D GM quanti- fied shell shape with notably high precision for the symmetric component of shape variation. In contrast, the asymmetric component of shape variation exhibited less precision probably because of confounding variation introduced by the position of the camera. Geometric prop- erties of a shell shape matched closely between 3D and 2D GM suggesting appropriate accu- racy of a 2D GM approach for the symmetric component of the shape variation. In summary, 2D GM can be used in studies of free-living chelonians giving opportunities for a more rigorous estimation of shell shape and its association with fitness in the field.

214 215 ECOLOGY CLIMATE CHANGE

Habitat preference of fire salamanders (Salamandra salamandra) in Adaptive potential and phenotypic plasticity of two anuran species, Stiavnické Vrchy Mountains Rana temporaria and Bombina variegata, from northern Steigerwald (Bavaria, Germany) in a response to thermal heterogeneity Gergely Dobay 1, István Kiss 2, Dániel Kaczkó 3 Sanja Drakulic 1, Heike Feldhaar 2, Theresa Spatz 3, Michael Seiler 2, 1 Institute of Botany and Ecophysiology, Szent István University, Páter Károly u. 1, 2100 Gödöllo˝, Hungary; Carolin Dittrich 1, Mark-Oliver Rödel 1 [email protected] 2 Department of Zoology and Animal Ecology, Szent István University, Páter Károly u. 1, 2100 Gödöllo˝, Hungary 1 3 Department of Aquaculture, Szent István University, Páter Károly u. 1, 2100 Gödöllo˝, Hungary Museum für Naturkunde, Invalidenstraße 43, 10115 Berlin, Germany; [email protected] 2 Department of Animal Ecology I, University of Bayreuth, 95440 Bayreuth, Germany 3 Department of Animal Ecology and Tropical Biology, University of Würzburg, 97070 Würzburg, Germany In case of amphibians like fire salamander (Salamandra salamandra) the habitat loss and the major changes causes decline or the extinction of the local populations. That is why habitat In order to survive, organisms have to cope with rapid, natural or human-induced changes preference surveys are becoming more important. In our survey we observed a fire salaman- in habitat structure. Dramatic declines in amphibian species populations worldwide are a der population in Slovakia. We captured the metamorphosed salamanders between 2007 and result of the complex interaction among multiple stressors, with climate change implicated 2011 along a 745 m long line transect on the western side of Makoviste mountain in the to be one of the principal factors. Our study focuses on the adaptive potential of two anuran Stiavnické Vrchy Protected Landscape Area (CHKO Stiavnické Vrchy). 9% of the 177 cap- species, the European common frog, Rana temporaria, and the yellow-bellied toad, Bombina tured and identified individuals were recaptured at least one time. We managed to characterize variegata, in the region of northern Steigerwald (Bavaria, Germany), and their ability and the habitat in 5–5 quadrats in the 4 different parts of the forest using 12 environmental x potential to answer to the rapid changes in environmental, namely climatological, condi- parameters. We have complemented these 20 quadrats with 2 5 quadrats, where we had tions. We examined the functional responses of those species to potential environmental information about the presence and absence of the species. We made a Principal Component changes, as well as the scale on which the responses are manifested: at the individual, pop- Analysis (PCA) with the gained datas. According to our results the habitats differ mainly in ulation or species level. In order to assess the adaptive potential of focal species to variations the ratio of uncovered and gravel surface coverage, and in the average of the tree trunk cir- in temperature conditions, we tested the differences in preferred temperatures among larvae cumferences. We found the highest density of fire salamanders in two habitats, where forest of populations originating from thermally differing microhabitats (exposed vs. closed micro- stands has completely different age and species composition. These habitats has nearly the habitats; i. e. open areas against forest). Furthermore, we experimentally examined the larvae’s same surface coverage ratios. Analysis of the data gained from the quadrats situated according developmental/growth rate adaptations to local thermal conditions, tracking the differences to presence and absence showed, that while the species composition of the forest vegetation in developmental progress under different environmental temperatures. appears to be irrelevant, the ratio of uncovered and gravel surface coverage were those factors which influenced the presence of the species. According to the results of our survey we ascer- tained that in the observed area fire salamanders prefer habitats where the ratio of uncovered surface is relatively high and evade those, where the extent of surface covered with rocks and gravel is more than 50%.

216 217 ECOLOGY BEHAVIOUR

What is the ‘ideal’ stream-adapted anuran larva? Bet-hedging in tadpole deposition in the Neotropical frog Allobates femoralis Anna Eckerlein, Stefan Lötters, Michael Veith, Bruno Viertel Magdalena Erich, Max Ringler, Walter Hödl, Eva Ringler Department of Biogeography, Trier University, Universitätsring 15, 54296 Trier, Germany; [email protected] Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; Stream adaptation has convergently evolved in tadpoles of about 20 anuran families. The evo- [email protected] lutionary change enabling adaptation to lotic waters resulted in different ‘morphotypes’. Based on this, gastromyzophorous, oroadhesive and pleuroadhesive larvae can be distin- Within anurans terrestrial oviposition has independently evolved several times, presumably guished from a ‘standard type’ adapted to lotic habitats. But the degree of stream adaptation as an adaptation to high predation pressure on eggs and the risk of competing stray sperm can also be defined stressing the habitat component. We ask if there is a common pattern of from rivals in aquatic environments. Terrestrial egg deposition is observed almost exclusively which ecomorphological characters were modified as an adaptation to the lotic environment, in tropical regions where it is enabled by the high humidity which prevents eggs from desic- and which combination of adaptations may characterise an ideal stream adapted anuran cating. The dilemma between terrestrial eggs and the aquatic anuran larvae is resolved in larva. Based on a large and world-wide body of literature, we compiled morphological char- most dendrobatid species by obligatory parental care in the form of tadpole transport. In the acters that have been identified as important in conjunction with stream adaptation. We sta- Neotropical frog Allobates femoralis (Dendrobatidae), clutches are laid in the leaf litter and tistically analysed how often these characters were in fact found in stream dwelling larvae. tadpole transport to water bodies is mainly performed by males. Facing the risks of aquatic This allowed us to eliminate those parameters that most obviously are not of primary adap- predators in, or desiccation of ephemeral pools, the partitioning of larvae over different pools tive value for a life in lotic habitats. Degree and combination of different ecomorphological would be a suitable bet-hedging strategy to overcome the risk of total clutch loss. In the present adaptations to lotic habitats vary among species, depending on the habitat itself (fast versus study we assessed the distribution of transported offspring across water bodies in a wild pop- slow flowing water). Some characters were more often found than others, such as numerous ulation of A. femoralis in French Guiana in spring 2010. We used seven highly polymorphic labial tooth rows and oral sucker (e. g. Heleophryne), large abdominal sucker (e. g. Atelopus) microsatellite loci and DNA samples of adults and of larvae from 30 artificial pools which or funnel-shaped mouth (e. g. Leptodactylodon). Our analysis suggests that no tadpole com- had been installed in 2009. Single pools contained on average 43.25 larvae (SD= 57.40). The bined all features identified as important adaptation to life in running water. The variety of reconstructed pedigree revealed that of the A. femoralis males that were identified as fathers combinations realised in larvae of different anuran families reflects (i) the different types of of more than one tadpole, 65.1% had partitioned successive clutches and 25.40% had spread lotic habitats and (ii) hints at numerous phylogenetic constraints on the morphological evo- single clutches across multiple pools. The partitioning of single clutches was significantly lution of stream adaptation. more often observed in males that were in their second or third breeding season, suggesting older males had probably more experience regarding the location of the artificial pools. Our results clearly demonstrate that A. femoralis males use bet-hedging in tadpole deposition by distributing single and successive clutches across several water bodies.

218 219 ECOLOGY BEHAVIOUR

Home range and movement patterns of the Neotropical freshwater Release and function of alarm pheromones in agile frog turtle Hydromedusa maximiliani (Testudines: Chelidae) in southeastern (Rana dalmatina) tadpoles Brazil Gábor Fera, Katalin Pásztor, Attila Hettyey Shirley Famelli 1,3, Leonardo Ramos Adriano 1, Fabrízio P.A. Pereira 1, Franco Leandro Souza 2, Bernd Gruber 3, Arthur Georges 3, Jaime Bertoluci 1 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected]

1 Departamento de Ciências Biológicas, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, In response to a predator attack, many prey species release chemical alarm signals that alert Av. Pádua Dias 11, 13418-900 Piracicaba, São Paulo, Brazil; [email protected] 2 Universidade Federal de Mato Grosso do Sul, Centro de Ciências Biológicas e da Saúde, 79070-900 Campo Grande, other potential prey to the threat. These chemicals, called Schreckstoff, are present in many Mato Grosso do Sul, Brazil aquatic vertebrates. While the chemicals and their release mechanisms have been identified 3 Institute for Applied Ecology, University of Canberra, ACT 2601, Australia in a few species, these seem to vary among species and we know little about how these alarm substances are released in larvae of anuran amphibians. Also, it has been suggested, that A key factor in conservation biology is a suitable design of protected areas. However, long- Schreckstoff may not only function to alert other individuals, but may also enhance the term studies are required when determining the size, structure, and shape of these areas for releaser’s survival probability by attracting further predators that interfere with the primary freshwater turtles, thereby integrating essential information to guarantee the reproductive attacking predator. Using tadpoles of the agile frog (Rana dalmatina), we experimentally success, recruitment of new individuals into the population and persistence of the population investigated if disturbed individuals release Schreckstoff actively, and if common predators of over the years. Nineteen adult animals (10 males and nine females) were fitted with radio- tadpoles are attracted to chemical alarm signals. As predators, we used larvae of the southern transmitters to analyze the home range, movement patterns, habitat use and selection of hawker (Aeshna cyanea) and of the great diving beetle (Dytiscus marginalis), young individuals Hydromedusa maximiliani, an endemic and vulnerable freshwater turtle from Atlantic Rain- of the western pike (Esox lucius), and adult males of the smooth newt (Lissotriton vulgaris). forest in Brazil, one of the most threatened ecosystems of the world. Another 18 adult animals We exposed predators and groups of tadpoles to (1) homogenized tadpoles (all types of cues (11 females and seven males) were monitored with thread-bobbins. Minimum Convex Polygon, released by prey), (2) the smell of tadpoles disturbed with a glass stirrer (only actively secreted Kernel, and Cluster were applied to measure home range size, which varied between 0.2 and chemical cues), or to (3) the smell of undisturbed tadpoles (no alarm substances) and recorded 1.5 ha over a year of monitoring. Mean size was 0.4 ha for both the wet and the dry season. their behaviour. Our results suggest that R. dalmatina tadpoles can not only passively, but Home range measured using different methods did not differ despite the time difference in also actively release alarm pheromones. Further, predators moved towards the source of alarm their application (72 hrs for thread-bobbins and 1 yr for telemetry). This result reflected the pheromones and, thus, seemed to be attracted by the chemicals released by disturbed tad- small scale of movements of H. maximiliani (3 to 138 m/day). Frequent use of burrows was poles. It will be interesting to see in future studies to what degree the predator attractant associated with sedentary behaviour and strong fidelity to site by the turtles within the river. property of alarm pheromones enhances the survival probability of attacked individuals under The importance of terrestrial habitat for these turtles is established, and suggests that long- realistic conditions. term monitoring is necessary to study this aspect of their biology. Such knowledge will con- tribute to future conservation strategies together with identification of nesting sites, and habitat use and selection of young turtles.

220 221 CLIMATE CHANGE ECOLOGY

The invasion potential of pet herps under climate change in Europe The effect of a common herbicide on survival of agile frog (Rana dalmatina) tadpoles in the presence of lethal predators Katharina J. Filz, Stefan Lötters Zoltán Gál, János Ujszegi, Zsanett Mikó, Attila Hettyey Trier University, Biogeography Department, Universitätsring 15, 54296 Trier, Germany; [email protected] “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, Biological invasions are driven by human-mediated dispersal of individuals beyond their Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] native ranges and worldwide cause severe ecological and economic disruptions. The magni- tude of these damages continues to grow in times of growing mobility and economic develop- Roundup® is one of the world’s most widely used herbicide. It consists of two components: ment with almost all ecosystems being heavily impacted by humans. Pet trade for aesthetic glyphosate, which inhibits the synthesis of essential amino acids, and polyethoxylated tallow motivations or research needs accounts for the majority of herpetological introductions. amine (POEA), which acts as a surfactant. This surfactant can be highly toxic to aquatic ani- Today, most amphibian and reptile species intentionally released or unintentionally escaped mals. It causes morphological changes in tadpoles and decreases their survival. In our study we are not capable of building invasive populations. The main reason might be a climatic mis- observed how the acute and chronical presence of the herbicide affects the survival of agile match between their original habitat climate and the climatic conditions at the receiving frog (Rana dalmatina) tadpoles. We raised tadpoles in outdoor mesocosms at two Roundup® location. We asked whether anthropogenic climate change, as predicted for the future, does concentrations (0 or 1.5 mg a. e/l glyphosate) and three predator treatments [no predator, increase the invasion potential of common pet herp species in Europe by employing species caged smooth newt (Lissotriton vulgaris), caged dragonfly larva (Aeshna cyanea)] in a full- distribution modeling approaches. Potential ranges of 25 amphibians and reptiles originating factorial design, using randomized spatial blocks. Twenty days after start we removed tadpoles from subtropical climates were computed under their current climatic niche (using occur- from rearing boxes, marked them individually, and presented them to free-ranging predators rence records from both native and, if applicable, invasive ranges) and projected onto climatic in clear water or in water containing Roundup®. Bioassays lasted for 24 hours. At termination, simulations for the years 2050 and 2080. The majority of taxa is more likely to occur under we counted survivors and identified from which rearing environment they had come from. climate change in Europe than under current climate, raising concerns that the rate of estab- As we expected, predator-induced responses resulted in significantly elevated survival proba- lishment and hence the magnitude of ecological and economic impacts might increase. bilities of tadpoles. Surprisingly, however, tadpoles reared in the presence of the herbicide also enjoyed higher survival rates in the presence of free-ranging predators, compared to those reared in the control treatment. The acute presence of the herbicide during the bioassays also lowered predation rates. Consequently, while spill-over of environmental contaminants into pristine habitats can be very harmful to some species, it may also have unexpected positive effects on others. This result supports the notion that effects of contaminants need to be tested on a row of species and investigations relying on one single or just a few indicator species can be misleading.

222 223 BEHAVIOUR PHYLOGENY

Diversity of lateralization patterns in Tarentola geckos Phylogenetic signal of the advertisement call in tropical hylids (Smilisca and Triprion) Enrique García-Muñoz 1,2,3, Catarina Rato 2, Bárbara Mesquita2, Miguel A. Carretero 2 Roberto García-Roa 1, Diego Llusia 2, David Buckley 2, Rafael Márquez 2 1 CESAM, Centro de Estudios de Ambiente o do Mar, Universidade de Aveiro, Campus Universitário de Santiago, 3810-193 Aveiro, Portugal; [email protected] 1 Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), José Gutiérrez Abascal 2, 2 CIBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos, Universidade do Porto, Campus Agrário de 28006 Madrid, Spain; [email protected] Vairão, 4485-661 Vairão, Portugal 2 Departamento de Biodiversidad y Biología Evolutiva, Museo Nacional de Ciencias Naturales (CSIC), 3 Departamento de Biología Animal, Biología Vegetal y Ecología, Universidad de Jaén, Campus de las Lagunillas s/n, José Gutiérrez Abascal 2, 28006 Madrid, Spain 23071 Jaén, Spain

The existence of laterally-biased behaviours in vertebrates is traditionally interpreted in terms Advertisement calls are important in species recognition and reproduction in many animal of saving costly duplication of neural circuitry and decreasing contradictory orders from the taxa and, hence, sexual selection drives sound signals to be species-specific traits. However, two hemispheres. However, when applied to antipredator behaviour, this is only valid from the evolution of these sexual traits also likely reflects the evolutionary history of taxa. Here, point of view of individuals. At higher levels, being prey behaviour lateralized could carry we study the phylogenetic signal underlying advertisement calls in tropical hylids (Smilisca negative implications, especially if predators may make prediction after multiple encounters and Triprion), which may provide insights into the evolution of sound signals and the debated with prey. These conflicting pressures, namely between the advantages for individuals, and phylogenetic relationships of species within these genera. We generate a phylogenetic hypo- the disadvantages for populations could be concealed if higher-level lateralization would arise thesis based on datasets available from GenBank and the Animal Sound Library of MNCN from the combination of lateralized behaviours of individuals which are mutually dependent. (Fonozoo). We compare our results with previously published studies, discussing the utility Here, we investigated the lateralization patterns in escape behaviour by six species of the gecko of advertisement calls in phylogenetic reconstruction. Moreover, we reconstruct, on the pro- genus Tarentola undergoing a predatory attack simulation in a “T” maze experiment. Contrary posed phylogeny, the ancestral character states of advertisement calls (i. e., temporal and to early claims for invariant lateralisation patterns in vertebrates, results showed that gecko spectral features), and we perform phylogenetic generalized least squares (PGLS) analyses populations and species displayed variable degrees of lateralization. While showing a general to examine the evolutionary changes in the characters and their correspondence with the trend for escaping to the right refuge, some species showed left- and no refuge preference. phylogenetic history of the group. We discuss the results of the phylogenetic and compara- tive study in reference to the historical scenarios for the diversification within the group and the current distribution of species to understand the different evolutionary processes shaping the evolution of sound signals in these tropical hylids.

224 225 PRACTICAL CONSERVATION ECOLOGY

Sublethal concentrations of glyphosate affect the life history and Differences in metabolic needs may relax competition and enable antipredatory behaviour of Iberian amphibians, which could reduce coexistence of several species of tadpoles within a Mediterranean their probability of survival anuran community

Ion Garin-Barrio 1, Carmen Martínez-Saura 2, Carlos Cabido 1 Núria Garriga, Eudald Pujol-Buxó, Gustavo A. Llorente

1 Aranzadi Society of Sciencies, Alto de Zorroaga 11, 20014 Donostia, Spain; [email protected] Departament de Biologia Animal, Facultat de Biologia, Universitat de Barcelona, Av. Diagonal 643, 2 Southeastern Association of Naturalists (ANSE), Plaza Pintor José María Párraga 11 (Bajo), 30002 Murcia, Spain 08028 Barcelona, Spain; [email protected]

Sublethal doses of glyphosate may have hidden effects which could reduce the probability of Volterra’s models suggested that n species could not coexist indefinitely on fewer than n survival and cause delayed mortality in amphibians. For instance, the time to complete meta- resources, but numerous examples of coexistence apparently violate this competitive exclu- morphosis is a significant ecological pressure, especially for species occupying temporary sion principle. The anuran community of our study area (Girona Province, NE Spain) com- ponds. If sublethal doses of glyphosate affect growth and delay metamorphosis, this may sup- prises a total of 9 anuran species that, although showing different preferences, are usually pose that larvae will not conclude the metamorphosis quickly enough or that they will with forced to coexist in their larval stages. Thus, with a high niche overlap, this system is highly a smaller size, compromising their long-term survival. On other hand, amphibian larvae suitable to evaluate the mechanisms developed for their coexistence. The aim of the study was reduce their activity level when they detect a predator to prevent to be localized. However, to evaluate the metabolic mechanisms that may enable coexistence by assessing food exploita- this also involves costs (lower feeding and development rate) that may affect long-term sur- tion abilities of all tadpole species present in the study area. This was done using a standard- vival. Sublethal doses of glyphosate could also alter these delicate trade-offs causing indirect ized experimental protocol, analyzing metabolic and morphologic parameters of all species in mortality (due to predation, for instance). Thus, first we determined by laboratory tests the the same conditions. We first determined which variables better explained the growth rate lethal concentrations (LC50-96h) for seven species of Iberian amphibians. The most resistant of tadpoles, so then we could explore the specific differences in the tactics used to exploit species were Alytes obstetricans and Pelophylax perezi, whereas the most vulnerable were Pelodytes resources among species. Therefore, we were able to classify species into different general punctatus, Bufo calamita and Rana temporaria. Second, we calculated the glyphosate concent- strategies and relate these strategies with ecological and evolutionary implications. Growth ration that is expected in small ponds following the recommended herbicide use in Spain Rate was the response variable and unexpectedly the only one to show a significant phylo- (expected environmental concentrations, EEC). Then, we examined growth, time to meta- genetic signal. However, Consumption rate and Production efficiency were the most important morphosis and size at metamorphosis of P. perezi and P. punctatus tadpoles, after being exposed variables defining individual Growth rate, surpassing species in importance and leaving to EEC or clean water. We also examined growth and antipredatory behaviour (activity level Assimilation efficiency and morphologic aspects like Gut length a marginal importance. of larvae before and after perceiving the odour of a predator) of A. obstetricans tadpoles Consumption rate and production efficiency defined a gradient of “consumer vs. efficient” exposed to 1/3 of their LC50 or clean water. Larvae exposed to EEC presented lower rate of strategies in which our species are spread according to several ecological patterns. Our data growth, and metamorphosed earlier and at a smaller size. In addition, A. obstetricans larvae on the exploitation abilities of each species may lead to a better comprehension of the guild exposed to sublethal glyphosate concentration presented overall lower rate of activity and a structure of European tadpole communities in general as a good example of a competing further reduction of their activity in face of a predator. grazers’ community.

226 227 DISEASES AND PARASITES BEHAVIOUR

Batrachochytrium dendrobatidis detection in amphibian samples Reactions of green lizards (Lacerta viridis) to the defensive secretion of using real-time PCR with SYBR GREEN chemistry Graphosoma lineatum (Heteroptera: Pentatomidae)

Elena Grasselli, Giulia Vecchione, Simone Di Piazza, Mirca Zotti, Martina Gregorovicˇová Sebastiano Salvidio Department of Zoology, Faculty of Science, Charles University, Vinicˇná 7, 12844 Praha 2, Czech Republic; Dipartimento di Scienze della Terra, dell'Ambiente e della Vita (DISTAV), Corso Europa 26, 16132 Genova, Italy; [email protected] [email protected] Chemical defence of true bugs (Heteroptera) is based on repellent secretions (synthesized or Chytridiomycosis is an emerging infectious disease of amphibians caused by the fungus sequestered), which may signal the unpalatability or be directly toxic for the predator. We Batrachochytrium dendrobatidis (Bd). Boyle et al. (2004) developed a method for detecting have tested the reaction of wild-caught green lizards (Lacerta viridis) as potential predators Bd using real-time Taqman PCR assay. Starting from this method, we have set up and vali- of true bugs in order to find out (1) whether green lizards avoid the chemically defended true dated a real-time PCR assay based on the cheaper SYBR GREEN chemistry. Primers used bugs, and (2) which of the components of true bug defensive secretion are aversive for lizards. for real-time PCR were designed on the basis of the Boyle’s ITS-1/5.8S rDNA sequence with The striated shieldbug (Graphosoma lineatum) was used as a model prey species. The following some modification for forward primer. Efficiency has been determined by a ten-fold serial chemicals, which form major components of shieldbug defensive secretion, were tested sep- dilution ranging from 100 to 0.1 Bd GE. By using fast real-time PCR assay we analyzed the arately (applied on palatable larvae of Tenebrio molitor [mealworms]): mixture of three alde- extracted total nucleic acids from a toeclip of the invasive bullfrog Lithobates catesbeianus, hydes (2-hexenal, 2-octenal, 2-decenal), the same aldehyde mixture enriched with tridecane, caught near Turin (Piedmont, NW Italy) from an infected population. Histological prepara- oxoaldehyde (4-oxohex-2-enal) and direct extract from metathoracic scent glands of adult tions from the same individual were also performed for comparison. Primers amplified a 146- Graphosoma lineatum. Hexane and mealworm without any chemicals added were used as bp product in ITS-1/5.8S rDNA region with a 100% efficiency. We were able to detect 1 Bd controls. Aversive reactions were evaluated on the basis of latencies to the beginning of GE in total nucleic acids extracted from the bullfrog in 1:10 diluted sample (final concentra- attacking/eating the prey with particular chemical components. The longer the response time, tion detected: 10 Bd GE). This assay constitutes a further powerful tool to help the detection the stronger the intensity of aversive reaction. The strongest rejection of prey was elicited by of Bd in the wild, thus contributing to the monitoring and the conservation of amphibian direct secretion of G. lineatum. The results show that the mixture of aldehydes is aversive populations. for green lizards. On the contrary, the mixture of aldehydes and tridecane has smaller effect. Oxoaldehyde caused stronger aversive response than mixture of three aldehydes and tridecan in latencies of eating the prey. The effectiveness of oxoaldehyde was enhanced by predator’s experience with living shieldbugs. Unlike other chemicals tested it seems that the oxoalde- hyde does not work as an olfactory signal but it becomes aversive in association with direct experience after eating the prey with oxoaldehyde compound. The project was supported by grant GACR P505/11/1459.

228 229 PHYSIOLOGY POPULATION GENETICS

Ecotoxicological risk of cadmium, lead, and mercury to early life Genetic characterisation of isolated populations of Aesculapian snakes stages and the reproductive function of reptiles (Zamenis longissimus)

Britta Grillitsch 1, Luis C. Schiesari 2, Heinz Grillitsch 3 Ina Groß

1 University of Veterinary Medicine of Vienna, Veterinaerplatz 1, 1210 Wien, Austria; Institute of Pharmacy and Molecular Biotechnology, Im Neuenheimer Feld 364, 69120 Heidelberg, Germany; [email protected] [email protected] 2 University of São Paulo, Av. Arlindo Betio, 1000 São Paulo, SP 03828-000, Brazil 3 Natural History Museum of Vienna, Burgring 7, 1010 Wien, Austria Zamenis longissimmus occurs over a wide range of Europe and in one separate population in northern Iran. Apart from the contiguous distribution several isolated populations can be Cadmium, lead, and mercury are harmful environmental contaminants of global concern, found. Though Aesculapian snakes occur over a broad range they seem to decline in num- with a high potential to cause developmental and reproductive disorders in humans and ani- bers in many regions. While isolated populations are most severely affected, numerous pop- mals. For reptiles, high accumulation levels of these metals were reported, but related risk is ulations within the main range are also facing problems. The most serious threats to Zamenis unknown. Considering that such effects may directly affect population viability, the present are mainly human-associated like habitat loss and fragmentation. When large connected study aimed at rating the risk of Cd, Pb, and Hg to reptiles by comparing internal exposure areas of suitable habitat are getting altered (e. g. by intensive agriculture, human dwelling, levels of reproductive and early life stage (RELS) compartments reported in the literature as road-building) and only small patches of the former habitat remain. Depending on the the best available indicators of toxicological availability in target tissues with threshold levels amount of the remaining habitat patches and the distance between them, these factors are causing reproductive and developmental effects in reptiles and birds as the established surro- leading to a reduced population size, as well as to the fragmentation of populations. A seri- gate taxon. Overall 54 original publications (Pubs) reported concentrations of these metals in ous problem for small and isolated populations is that they are more prone to stochastic fac- 18 different RELS tissues (including male and female gonadal system as well as egg, embryo tors like genetic drift, inbreeding and loss of genetic diversity, which can be another cause and hatchling compartments) for 24 reptile species. Despite this patchy data set, maternal of extinction. transfer was confirmed (10 Pubs), internal exposure of RELS was almost ubiquitous, the Within the framework of a dissertation these genetic factors are investigated more closely observed concentration levels matched those reported for birds and exceeded the thresholds to evaluate the risk of extinction of Zamenis in isolated populations. Therefore several molec- causing relevant sublethal effects in birds by factors of 10–100. Regarding the effects of Cd, ular methods have been applied already to compare isolated with non-isolated populations of Pb, and Hg to RELS of reptiles, the database turned out to be particularly small as it included Aesculapian snakes. Until now genetic samples from two completely isolated populations only 11 Pubs (9: Cd, 1: Pb; 1: Hg) out of which there was evidence for interactions with the from Germany (Schlangenbad and Hirschhorn) and from one area within the main distribu- gonadal axis of the endocrine system (for Cd and Pb), developmental disorders (Cd) and tion area in Hungary (around Budapest) have been analysed. Besides sequence analyses of changed gene expression in ovary and embryos (Cd) at exposure concentrations at factors mitochondrial genes (CytB, ND1) different genetic markers for the investigation of genomic 100–1000 below the observed RELS tissue concentrations. In conclusion, this review indi- DNA [Inter-Simple-Sequence-Repeats (ISSR), microsatellites, multi-locus fingerprinting] cates that for reptiles, both toxicological availability and sensitivity of RELS to Cd, Hg, and have been applied. These results indicate a significant decreased genetic diversity within iso- Pb may be widespread even at the micro-pollution scale, and research appropriate to under- lated populations. stand source to adverse outcome pathways is urgently needed.

230 231 ECOLOGY TROPICAL HERP BIODIVERSITY

Age structure and body size in two populations of the Phylogeography of Leptopelis frogs across the sky islands of the northern banded newt Ommatotriton ophryticus (Berthold, 1846) Eastern Arc Mountains in East Africa from Turkey Václav Gvozˇdík 1, Jean Mariaux 2, Michele Menegon 3, Simon Loader 1 Çiçek Gümüs¸, Nazan Üzüm 1 Department of Environmental Sciences, Biogeography, University of Basel, Klingelbergstraße 27, 4056 Basel, Switzerland; [email protected] Adnan Menderes University, Faculty of Arts and Sciences, Department of Biology, Aydin, Turkey; 2 Muséum d’Histoire Naturelle de Genève, Dpt. des Invertébrés, CP 6434, 1211 Geneva, Switzerland [email protected] 3 Tropical Biodiversity Section, Trento Science Museum, Via Calepina 14, 38100 Trento, Italy The aim of the current study was to determine of age structure and body size of two different Five Leptopelis species are distributed in the “sky island” system of the Eastern Arc Mountains populations of Ommatotriton ophryticus from northern Turkey. of Kenya and Tanzania. Four of these species are confined to forest habitats scattered along We studied 50 samples (20 males and 30 females) from Bahçesultan/Bilecik (1015 m a. s.l.) different mountain blocks, typically at high elevations. The sky island system is an ideal area and 60 samples (30 males and 30 females) from Tosya/Kastamonu (1239 m a.s.l.). Adult for a comparative phylogeographical study to investigate common patterns in shaping genetic males and females caught during the breeding season all possessed external secondary sexual variation in different species. We studied mitochondrial and nuclear DNA sequences of the characters and were considered mature. We measured the snouth-vent length (SVL) of indi- four montane species, L. barbouri, L. parkeri, L. uluguruensis, L. vermiculatus, and one wide- viduals to compare the body size of the populations. The age was determined by skeleto- spread species common in lower and mid-elevations, L. flavomaculatus. A common pattern chronology performed counting lines of arrested growth (LAGs) of the phalanges. showed the presence of numerous highly divergent lineages between most montane popula- Results showed that the maximum lifespan was 9 years in Tosya and 10 years in Bahçesultan. tions concordant with geography, while the widespread L. flavomaculatus is genetically rela- Age at sexual maturity of both males and females was 4–5 years in the Tosya while 6 years for tively uniform. This pattern appears to be the result of the complex geological history of the both sexes in the Bahçesultan population. No significant differences were found between the region and Quaternary climatic oscillations. Such change affected the extent of forest habi- age distributions of the sexes in Bahçesultan (t=–0.05, p=0.961) while there was a significant tats, and thus enabled or disabled gene flow between different mountain blocks. difference in Tosya (t=–2.337, p=0.023). The age structure of males (t=–2.987, p=0.004) varied significantly between populations whereas there was no significant difference in females (t=–1.482, p=0.144). The SVL (snout-vent length) ranged from 64.09 to 78.87 mm for males and from 57.64 to 70.72 mm for females in Bahçesultan. It ranged from 58.41 to 71.55 mm for males and from 50.33 to 61.83 mm for females in Tosya. Significant differences were found between sexes and also between the populations in terms of mean SVL. A signif- icant correlation between age and snout-vent length (SVL) was found for females while no significant relationship was determined for males in both populations.

232 233 TROPICAL HERP BIODIVERSITY ECOLOGY

Herpetofauna of northern Cambodia: biodiversity, ecology and Habitat use and daily activity of Lacerta viridis and Lacerta agilis zoogeography in a cemetary

Timo Hartmann 1, Peter Geissler 1, Markus Handschuh 2, Flora Ihlow 1, Botond Heltai 1, Péter Sály 2, István Kiss 1 Dennis Rödder 1, Rattanak Seng 3, Wolfgang Böhme 1 1 Department of Zoology and Animal Ecology, Szent István University, Páter Károly u. 1, 2100 Gödöllo˝, Hungary; [email protected] 1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; 2 HAS Centre for Ecological Research, Balaton Limnological Institute, Klebelsberg Kunó u. 3, 8237 Tihany, Hungary [email protected] 2 Angkor Centre for Conservation of Biodiversity (ACCB), Kbal Spean, Phnom Kulen National Park, Siem Reap, Cambodia Cemeteries tend to contain diversified habitat patches and can play significant role in tem- 3 Department of National Parks, Ministry of Environment (MoE), 48 Samdech Preah Sihanouk, Tonle Bassac, poral or permanent establishment of reptiles in urban areas due to their moderate anthro- Khan Chamkarmorn, Phnom Penh, Cambodia pogenic disturbance. The aim of our study was to examine the population size of the lizard species occurring in an urban cemetery, and investigate their daily activity pattern. The sur- After years of civil unrest hampering any research, the recent herpetofaunal investigations in vey was carried out in eight parcels with different age and vegetation structure of the ceme- Cambodia focused on the mountainous areas in the Southwest (Cardamom Mountains) and tery of Dunaújváros town, Hungary, during 1–7 July, 2012. Parcels were characterized by the East (the Greater Annamites Ecoregion) of the country. Between 2008 and 2011 we conducted area of the grassy, bushy and arboraceous patches, the sunny and the shaded ground patches. extensive herpetological baseline surveys within the Phnom Kulen National Park and the Lizard observations were made in the mornings, at middays and in late afternoons. Occurrence Kulen Promtep Wildlife Sanctuary located in northern Cambodia, which revealed a surpris- data were analysed by a generalized linear mixed model (GLMM) using binomial distribu- ingly high herpetofaunal diversity taken the relative low-land character (only up to 487 m tion in which the days of observations and the cemetery parcels were used as random factors. a. s. l.) of the protected areas into account. A total of 84 species (25 amphibian species, 59 rep- During the survey, two lizard species (Lacerta agilis and L. viridis) were detected. In the eight tile species) was found, of which four (Kaloula mediolineata, Leiolepis rubritaeniata, Spheno- parcels, the total number of the L. agilis observations was 178 and that of the L. viridis obser- morphus lineopunctulatus, Tropidophorus cocincinensis) were recorded for the whole of Cambodia vations was 79. In general, lizards were more abundant in the more patchy, some decades old for the first time. To get insight into the mostly understudied feeding ecology of the recorded parcels than in the more homogenous, recently formed ones. However, L. viridis tended to species, stomach content analyses of 623 specimens were undertaken. Finally, based on a occur in the more bushy parcels than L. agilis. Both species showed similar daily activity pat- zoogeographical comparison of species communities with better-studied neighbouring geo- tern. The probability of occurrence was associated to the temperature and observational time. graphical areas, we were able to integrate the revealed herpetofaunal community of northern According to the GLMM model, lizards showed the highest probability of occurrence between Cambodia into a more general view of broader-scale distribution patterns in Southeast Asia. 12 a. m. (ca. 31 °C) and 14 p. m. (ca. 33 °C). The variance of occurrence probability associated to the cemetery parcels was 4.28 times greater than the variance originating from the differ- ences of the observational days. Consequently, it seems that direct local environmental factors influence more the dial activity of lacertid lizards than the differences among the observational days do within a short (ca. one week) time period.

234 235 DISEASES AND PARASITES ECOLOGY

Amphibian parasites in the Hortobágy National Park: The importance of direct chemical cues for predator detection preliminary findings in tadpoles

Dávid Herczeg 1, Judit Vörös 2, Zoltán Barta 3, Zsolt Végvári 4, Daniel R. Brooks 5 A. Hettyey 1,2,3, Z. Tóth 1,3, K. E. Thonhauser 1, D. J. Penn 1, J. Van Buskirk 2

1 University of Debrecen, Department of Evolutionary Zoology and Human Biology, Egyetem tér 1, 4032 Debrecen, 1 Konrad Lorenz Institute of Ethology, Department of Integrative Biology and Evolution, University of Veterinary Hungary; [email protected] Medicine Vienna, Savoyenstraße 1a, 1160 Wien, Austria; [email protected] 2 Hungarian Natural History Museum, Baross u. 13, 1088 Budapest, Hungary 2 Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Winterthurerstraße 190, 3 MTA-DE, “Lendület” Behavioural Ecology Research Group, Egyetem tér 1, 4032 Debrecen, Hungary 8057 Zürich, Switzerland 4 Hortobágy National Park Directorate, Sumen u. 2, 4024 Debrecen, Hungary 3 MTA-ATK, “Lendület” Evolutionary Ecology Research Group, Herman Ottó út 15, 1022 Budapest, Hungary 5 University of Toronto, Department of Ecology and Evolutionary Biology, 25 Willcocks Street, Toronto, M5S 3B2, Canada The ability of prey to respond adaptively to the presence of predators depends on the avail- ability of reliable cues. Chemical cues are generally considered the most important modality Parasites are becoming recognized as significant participants in the evolutionary game and are for predator detection in the aquatic environment, especially so in turbid waters. Chemical seen as perfect model systems for general evolutionary studies. Our present knowledge of the cues on predation threat include general prey metabolites that are excreted upon stress, spe- adult platyhelminth parasites of vertebrates in Hungary especially in the Hortobágy National cial disturbance cues that are costly to produce and are released actively upon attack, cues that Park is defective. Our aim of examination was to reveal the helminth fauna of the Pelophylax are passively released from injured prey tissue, cues originating directly from the predator, esculentus complex and compare our findings with the results of previous studies and com- and constituents of prey tissue released through digestion by predators. While digestion pare the faunistical changes now with a baseline from more than 40 years ago. Fifty-nine released cues may reveal significant information about the predator species present and the individuals of Pelophylax spp. were collected between May and September of 2012 (35 P. ridi- types of prey it has been feeding on, we know rather little about to what degree prey take bundus and 24 P. kl. esculentus) and we found the following species of parasites. Digeneans: advantage of the information coded in the excretions of their predators. Here we investigated Haematoloechus variegatus, Opisthioglyphe ranae, Diplodiscus subclavatus, Pleurogenoides medians; whether tadpoles of the common frog (Rana temporaria) use the information present in the nematodes: Oswaldocruzia filiformis, Rhabdias esculentarum; acanthocephalans: Acantho- excretions of predators by investigating whether tadpoles adjust their induced defences de- cephalus ranae. Rhabdias esculantarum is a new species for the Hungarian fauna and P. ridi- pending on the diet of predators even when cues released by prey upon attack are excluded. bundus represents a new host record for Rh. esculentarum in Europe, while D. subclavatus is We reared tadpoles in the presence of caged predators (Aeshna cyanea). We fed predators dif- also a new species for the fauna of the Hortobágy. The prevalence was 84.7% [95% CI: ferent types of prey outside the mesocosms. Two hours after food provisioning, we removed 73,05–92,05] in the examined water frog population. The mean intensity was 2.44 (± 1.49) remaining prey, washed the predators and replaced their water. Another two hours later, we helminth species per host. No host contained more than 6 species: 15 (25.4%) of 59 Pelophylax placed predators back into the mesocosms. To assess variation in the induced defences of spp. harbored 1 species, 13 (22%) harbored 2 species, 12 (20%) harbored 3 species, 6 (10%) tadpoles, we monitored behaviour and assessed morphology. Our results showed that the harbored 4 species, 3 (5%) harbored 5 species, and 1 (1.6%) harbored 6 species. among-treatment pattern in the magnitude of behavioural changes was very similar to that observed in the morphological changes of tadpoles. Also, the strength of the response de- pended on the feeding history of predators. These results support the hypothesis that tad- poles extract information from the excretions of their predators.

236 237 ECOLOGY DISEASES AND PARASITES

Reduced impact logging and climate extremes: mutual determinants Occurrence of blood parasites in Spanish terrapins of homogenisation in tropical amphibian communities? (Mauremys leprosa) from the southwestern Iberian Peninsula

Monique Hölting 1,2, Raffael Ernst 1 Alex Ibáñez 1, José Martín 1, Pilar López 1, Alfonso Marzal 2

1 Museum of Zoology, Senckenberg Natural History Collections Dresden, Königsbrücker Landstraße 159, 01109 Dresden, 1 Dpto. de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, C.S. I. C., José Gutiérrez Abascal 2, Germany; [email protected] 28006 Madrid, Spain; [email protected] 2 Department of Biodiversity Dynamics, Institute of Ecology, TU Berlin, Rothenburgstraße 12, 12165 Berlin, Germany 2 Dpto. de Biología Animal, Universidad de Extremadura, 06071 Badajoz, Spain

The analysis of variation in species composition can provide fundamental insight into the Studies of blood parasites of turtles have been scarce and were based on visual examination processes leading to either homogenisation or diversification of tropical ecosystems that are of blood smears to detect parasites. Here, a total of 54 adult Spanish terrapins (Mauremys lep- exposed to human disturbances and/or climatic extremes. In our study we investigate beta- rosa) from two localities on Southwestern population of Iberian Peninsula, were screened for diversity pattern changes of amphibian communities in the lowland forest of Guyana that are apicomplexan parasites of the three genera Haemoproteus, Plasmodium and Hepatozoon using exposed to (a) selective logging and (b) extreme weather anomalies. We specifically analysed a recently developed polymerase chain reaction method. Overall prevalence was 57%. compositional change at different spatial scales and different species rarity-dominance weight Prevalence of Hepatozoon spp. (54%) and Plasmodium spp. (9%) was detected. No individual ratios. Data was collected in three consecutive years (2009–2011) within a controlled poly- was infected with Haemoproteus spp. Of the infected turtles, 6% carried mixed infections. We cyclic timber harvesting scheme implemented by our project partner Iwokrama International also found four genetically distinct parasite lineages infecting terrapins (2 Plasmodium, 2 Centre for Rainforest Conservation and Development. In addition to the logging impacts, Hepatozoon). Turtles from different localities significantly differed in their overall prevalence. the study area was exposed to an extreme weather anomaly during the first sampling year, However, overall prevalence did not differ between sexes. These new methods might have resulting in a prolonged drought period. A pressing question that needs to be addressed is strong impact on several aspects of research on turtle blood parasites and could be applied for thus whether such extreme events aggravate the effects of logging disturbance and conse- management and conservation of wild populations. quently shape diversity patterns through an additive amplifying process? Our results show that it is important to consider these synergistic effects when interpreting diversity pattern changes. Both extreme climatic events and selective logging had a rapid but also complex effect on amphibian composition. Beta-diversity patterns differed not only depending on time since logging, but also with regard to the emphasis given to rare or abundant species in the analysis. As particular species disappeared, others colonised or increased in abundance. And while impacted communities remained stable under the climate extreme event, pristine communities were very sensitive to drought. The next step in our analysis will be to investigate how changes in species composition will affect trophic networks in our study sites and how this in turn will impact ecosystem processes.

238 239 PRACTICAL CONSERVATION ECOLOGY

Are alien tree plantations ecological traps for amphibians? Home range and habitat selection of the endangered Effects on immune response and secondary sex characteristics of Euphrates softshell turtle Rafetus euphraticus (Daudin, 1812) in palmate newts a fragmented habitat in southwestern Iran

Maider Iglesias, Carlos Cabido Flora Ihlow 1, Hanyeh Ghaffari 2, Michael V. Plummer 3, Barbod Safaei Mahroo 2, Dennis Rödder 1 Aranzadi Society of Sciencies, Alto de Zorroaga 11, 20014 Donostia, Spain; [email protected] 1 Herpetological Department, Zoologisches Forschungsmuseum Alexander Koenig, Adenaueralle 160, 53113 Bonn, In the last century, the natural environment has been conversed to anthropized environment Germany; [email protected] on an unprecedented scale. The new simplified habitats have different selective pressures 2 Department of Environmental Science, Graduate School of the Environment and Energy, Science and Research Branch, Islamic Azad University, Tehran, Iran comparing with natural ones, so they are exploited by species adapted to them that can tol- 3 Department of Biology, Harding University, Searcy, AR 72149, USA erate dramatic changes. However, although some species identify these habitats as right ones, when they are not able to adapt to the new environmental conditions, anthropized habitats We present the first information on home range size, space use and habitat selection for the may act as ecological traps. The palmate newt (Lissotriton helveticus) has a wide distribution, endangered Euphrates softshell turtle Rafetus euphraticus (Daudin, 1802) from Karkheh often appearing in eucalyptus and pine plantations. However, since tree plantations may be Regulating Dam Lake in southwestern Iran. Fourteen turtles were trapped, including two acting as ecological traps, the health condition of individuals could be worse than those living juveniles and 12 adults, and subsequently fitted with radio tracking transmitters. Each turtle in natural habitats. To examine this hypothesis, we studied the health status in breeding was relocated between 21 and 51 times between May 2011 and July 2012 resulting in a total of males captured in six eucalyptus and six pine plantations, and six oak forests. We estimate the 254 fixes. The mean home range length (LHR) was 2.54 ± 0.83 km. The LHR was variable capacity of response of the immune system with phytohematoglutinin tests. Moreover, males (CV = 33%) among individuals but there was no statistically significant relationship between show secondary sex characteristics as hind feet webs, caudal crest and caudal filament. These linear home range size and straight carapace length (SCL) and linear home range size and seem to be honest signals that depend on the quality of the individual and the reproductive body mass. The mean river channel area was 55.35 ± 17.98 ha. The mean minimum convex success of each individual will depend on them. Therefore, they can also be use as quality polygon home range size (MCP) was 47.49 ± 23.36 ha and the mean 95% kernel home range indicators of the individuals living in different habitats. Hence, we measured the caudal fila- size was 21.75 ± 9.44 ha. Home range overlap was high. On average, individual MCP home ment’s length and the crest’s height of the captured individuals. We found that both health ranges overlapped with MCP home ranges of 7.5 other turtles, individual KDE home ranges status and sexual signals were different between populations: those living in eucalyptus plan- overlapped with those of 7.3 other turtles, and core areas overlapped with those of 5.5 other tations had worse immune response and less development of sex characteristics. Our results turtles. indicate that eucalyptus plantations have a negative effect on this species, with direct conse- quences, as worse immune capacity, and evolutionary ones, as the alteration of sexual selec- tion mechanisms.

240 241 CLIMATE CHANGE PHYLOGEOGRAPHY

What limits range overlap of common and Syrian spadefoot toads: Mitochondrial phylogeography of three species of slow worms climate or competition? (Anguis) in the Balkans

Ruben Iosif 1, Ciprian Samoila 1, Monica Papefl 2, Dan Coga˘lniceanu 1 Daniel Jablonski 1, Georg Dzˇukic´ 2, David Jandzˇik1,3, Dusˇan Jelic´ 4, Katarina Ljubisˇavljevic´ 2, Peter Mikulícˇek1, Jirˇí Moravec 5, Nikolay Tzankov 6, 1 Ovidius University Constant¸a, Faculty of Natural and Agricultural Sciences, Aleea Universita˘¸iit 1, corp B, Václav Gvozˇdík 5,7 900470 Constant¸a, Romania; [email protected] 2 Oklahoma State University, Department of Zoology, 501 Life Sciences West, Stillwater, Oklahoma 74078, USA 1 Department of Zoology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynska dolina B-1, 84215 Bratislava, Slovakia; [email protected] Four species of the spadefoot toad genus Pelobates, mostly allopatric, are distributed over 2 Department of Evolutionary Biology, Institute for Biological Research “S. Stankovic´”, 11060 Belgrade, Serbia parts of Europe, North Africa, Caucasus, and the Middle East. Ranges of P. fuscus (Pf) and P. 3 Department of Ecology and Evolutionary Biology (EBIO), University of Colorado, Ramaley N122, Campus Box 334, syriacus (Ps) overlap over a narrow strip in Dagestan and along Danube, in the Balkans. Our Boulder, CO 80309, USA 4 study focused on investigating (i) factors limiting the range overlap of the two species (cli- Croatian Institute for Biodiversity, Croatian Herpetological Society Hyla, I. Breznicka 5a, 10000 Zagreb, Croatia 5 Department of Zoology, National Museum, Cirkusová 1740, 19300 Prague, Czech Republic mate or competition), and (ii) possible changes in their geographic ranges under future cli- 6 Department of Recent and Fossil Amphibians and Reptiles, National Museum of Natural History, mate change scenarios. We computed climatic suitability models using MaxEnt and projected Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria their distribution onto future (i. e., A1B, A2, B1 scenarios) and past conditions (i. e., Last 7 Laboratory of Molecular Ecology, Institute of Animal Physiology and Genetics, Academy of Sciences of the Interglacial and Last Glacial Maximum). Additionally, we used fossil occurrences to test the Czech Republic, 27721 Libechov, Czech Republic predictive accuracy of past projections. Finally we assessed the degree of range overlap The Balkan Peninsula has played a role of a speciation and radiation centre in evolutionary between the studied species using metrics implemented in ENMTools. The present climatic history of slow worms (Anguis spp.). Four from five known species are present in the region. niche was slightly extended outside the known geographic range, in the Caucasus and the However, knowledge of the detailed distribution ranges of the species and their contact zones Balkans, south for Pf and north for Ps. The last interglacial distribution of Pf included British in the Balkans remains insufficient. In the first step of this project we genotyped 193 individuals Isles, and broad areas in western, central and northern Europe, and extended farther north from 152 localities of this region based on mitochondrial DNA (ND2) to map distributions in the Balkans for Ps. Validation with fossil records revealed an excellent predictive perform- and contact zones of mitochondrial lineages/species and to screen their mtDNA diversity. ance (Omission error = 4.1% for Pf and 16.6% for Ps). Last Glacial Maximum maintained According to our results, A. fragilis is relatively widespread in the north-western and central only isolated refugia in southern Europe, Pannonian Basin and Caucasus for Pf, and Israel, Balkans, south-westward from the Danube River. Relative genetic uniformity of this species Southern Balkans, and Caucasus for Ps. Present potential distributions revealed a narrow suggests a recent dispersal within the Balkans. Anguis colchica has been identified in the range overlap (Range Overlap = 0.44), while under Last Glacial Maximum the two species north-eastern and eastern Balkans in Romania, Serbia and Bulgaria. This species forms several had nearly disjunct allopatric distributions (Range Overlap = 0.17). The range overlap is divergent evolutionary lineages within the Balkans, probably related to different Pleistocene projected to increase towards the end of this century. The species’ absence from climatically refugia. Anguis graeca was, beside Greece, Albania and southernmost Montenegro, newly suitable areas supports the hypothesis that dispersal barriers and competition limit their geo- detected in Republic of Macedonia. In comparison to the other species, this Balkan endemic graphic range. shows much higher and complex genetic variation. This finding corresponds to the hypo- thesis of multiple Pleistocene microrefugia in the southern Balkans, the pattern similar to other endemic species. Further investigations will evaluate variation in nuclear genes allowing detection of historical or recent hybridization events.

242 243 PHYLOGEOGRAPHY PRACTICAL CONSERVATION

Phylogeography of Stellagama stellio: resolving the relationships Creating open water surfaces substantially increases amphibian within its subspecies biomass in eutrophic wetlands

Emmanouela Karameta 1, Nikos Poulakakis 2, Spyros Sfenthourakis 3, Patrik Katona 1,2,3, Balázs Vági 1,2,4, Tibor Kovács 1,2 Petros Lymberakis 2, Heinz Grillitsch 4, Çetin Ilgaz 5, Panayiotis Pafilis 1 1 Hungarian Biodiversity Research Society, Hunyadvár utca 43/a, 1165 Budapest, Hungary; [email protected] 2 1 Section of Zoology and Marine Biology, Department of Biology, University of Athens, Panepistimioupolis, Ilissia, Birdlife Hungary Amphibian and Reptile Conservation Group, Költo˝ utca 21, 1121 Budapest, Hungary 157-84 Athens, Greece; [email protected] 3 Department of Evolutionary Zoology and Human Biology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, 2 Department of Biology and Natural History Museum of Crete, University of Crete, Knossos Av., P.O. Box 2208, Hungary 71409 Irakleio, Greece 4 MTA-ATK, “Lendület” Evolutionary Ecology Research Group, Herman Ottó út 15, 1022 Budapest, Hungary 3 Department of Biological Sciences, University of Cyprus, University Campus, P.O. Box 20537, 1678 Nicosia, Cyprus 4 Herpetological Collection, Natural History Museum Vienna, Burgring 7, 1010 Wien, Austria Amphibians and reptiles could have vast biomass and usually play a crucial role in trophic 5 Department of Biology, Faculty of Science, Dokuz Eylül University, Tinaztepe Yerleskesi, 35160 Buca-Izmir, Turkey networks in wetland areas, therefore suitable habitat management for herpetofauna is vital in wetland ecosystems. In 2012 open channels had been created by sweeping in an exceedingly The roughtail rock agama Stellagama stellio (Linnaeus, 1758), until recently one out of 20 eutrophic area of the Ócsai Öregturján, a Natura 2000 Habitat Directive site, which had species of the genus Laudakia, comprises the commonly recognized subspecies: stellio (Greece, improved habitat complexity. In this research we made herpetofaunistic assessments in fixed Turkey, Syria, Lebanon, Israel, Jordan), daani (Greece and Turkey), cypriaca (Cyprus), vulgaris transects along the newly swept and older channels to follow changes in species composition (Egypt), brachydactyla (Jordan, Saudi Arabia, Sinai, Israel), picea (Jordan, Syria, Saudi Arabia) and biomass during vegetation periods and between years. We investigated if species abun- and salehi (Sinai, Israel). Despite extensive research focusing on the morphology-based tax- dance and colonization in the channels depended on the distance to undisturbed habitats, onomy of S. stellio, the phylogenetic relationships among the aforementioned subspecies aquatic and riparian vegetation cover, channel size parameters and regulation of water level. remain unclear. This is the first study aiming to resolve the phylogeography of the stellio com- In the area members of the water frog group (Pelophylax spp.) were the most abundant. plex along the species’ distribution and illustrate the resulting taxonomic consequences. For Density of water frogs had been already increased in the first year after sweeping. The greatest this purpose, we used more than 80 Stellagama stellio specimens belonging to all above sub- number of metamorphs appeared in channels with dense vegetation and transects in the species. Two partial sequences of a mitochondrial (ND4) and a nuclear (Nktr) gene were vicinity of undisturbed habitats. Colonization of new habitats had already started in the first targeted and several phylogenetic (Maximum Likelihood and Bayesian Inference) and phylo- year, nevertheless unbalanced population density remained between channels with different geographic (BEAST) approaches were performed. Taxonomic implications regarding the status vegetation cover, size parameters and habitat surroundings. Further monitoring may reveal if of the various subspecies and the distribution of the produced phylogenetic lineages in view aggregated distribution of anurans is the consequence of habitat alteration, and hence to be of the known palaeogeography of the region are discussed. compensated in the following years, or caused by other ecological background variables and will persist for a long time period. Our research supports that amphibian populations can be corroborated rapidly by sweeping in eutrophic habitats, and undisturbed habitats should be also preserved as they act as primer sources of colonization.

244 245 PRACTICAL CONSERVATION PRACTICAL CONSERVATION

On the status of water frogs Pelophylax sp. (Fitzinger, 1843) in Herpetological education projects in Hungary: effective tools for the Austrian state of Salzburg: morphological and preliminary protection and a meaningful support for research projects molecular data András Kéri 1,5, Mihály Tóth 2,4, Béla Mester 2,4, Tímea Mechura 3,4, Miklós Puky 4,5 Peter Kaufmann 1, Andreas Maletzky 1, Peter Mikulícˇek2 1 Eötvös Loránd University, Department of Physical Geography, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary 2 University of Debrecen, Faculty of Science and Technology, Department of Hidrobiology, Egyetem tér 1, 1 University of Salzburg, Department of Organismic Biology, Hellbrunnerstraße 34, 5020 Salzburg, Austria; 4032 Debrecen, Hungary [email protected] 3 Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 2 Comenius University Bratislava, Department of Zoology, Mlynska dolina B-1, 84215 Bratislava, Slovak Republic 4 Varangy Akciócsoport Egyesület, IX. utca 40, 1172 Budapest, Hungary 5 MTA Centre for Ecological Research, Danube Research Institute, Jávorka S. utca 14, 2131 Göd, Hungary Three water frog taxa occur in the Austrian state of Salzburg: Pelophylax ridibundus, P. lessonae and their hybrid P. esculentus. Due to the complexity of exact species identification, earlier Regular herpetological education started in the 1980s in Hungary with toad rescues over records and studies most often did not differentiate between those three taxa. Pelophylax roads and ad hoc translocation projects including highly conflicting areas between humans lessonae is strictly protected by means of the EU Habitats Directive (Annex IV) and presum- and herps such as swimming pools in villages and snake nesting areas under motorways. By ably declining in many parts of Austria, while P. ridibundus showed a recent spreading there, today, many people are also active in less demanding herp-related citizen science activities most likely due to human induced habitat changes. such as mapping species distribution by sending photos with the indication where they were To gain insight into the current distribution and taxa composition, we sampled 268 water shot to Varangy Akciócsoport Egyesület, a 28 year old NGO working in amphibian and rep- frogs in Salzburg and Upper Austria in 2011 and 2012. Morphometrical parameters of frogs tile conservation, research and education. Besides traditional talks and nature walks unusual were measured and species identification was carried out accordingly. Additionally tissue was activities, such as “Frog concerts”, also generated much interest. Similarly, launching the sampled via buccal swabs for subsequent genetic analyses. Amphibian of the Year, followed by the Reptile of the Year, was also an international award- Morphometrical results indicate that P. lessonae is present only in few isolated and natural winning event. In some cases, where there is a strong educational element, it may be advis- wetlands, while P. ridibundus is more widely distributed than assumed previously. The distri- able for scientists to give the lead in that sub-project for educators, as it turned out to be often bution as well as the recent spreading of P. ridibundus leads to the hypothesis that it could be the best way while making Carpathian Basin Children Competitions in Hungary, Romania, an introduced and potentially invasive species for Salzburg. Slovakia and Ukraine. It is especially advisable with young children, e.g. in programmes made Preliminary genetic analysis of mtDNA and microsatellite markers test for this hypothesis for kindergartens. In such events, the co-operation of professionals and educators is a key and also aim to affirm morphometrical species determination, obtain data on phylogeo- element of success. In any case when scientists work with volunteers, however, easily under- graphic relationships and evaluate genetic diversity of sampled populations. standable protocol and data validation is needed, and if the organizers fail to make these steps, more difficult changes had to make later, as another organization experienced it facing a high number of obviously incorrect herpetofauna species determination by its volunteers. Consequently, participants have to be informed and trained about what to do and they need to be informed about the results, too. With no doubt volunteers can successfully be involved in data collection, however, species determination and data analysis are still predominantly the field for professionals.

246 247 ECOLOGY POPULATION GENETICS

Ecological survey of a rediscovered urban fire salamander Isolation by time in the yellow-bellied toad, Bombina variegata? (Salamandra salamandra) population in Budapest Konstantin Knorr, Heike Pröhl István Kiss 1, Judit Vörös 2, Sára Borbála Balogh 1, Erika Jáger 3 Institute of Zoology,University of Veterinary Medicine, Bünteweg 17, 30559 Hannover, Germany; [email protected] 1 Szent István University, Department of Zoology and Animal Ecology, Páter K. u. 1, 2100 Gödöllo˝, Hungary; [email protected] 2 Department of Zoology, Hungarian Natural History Museum, Baross u. 13, 1088 Budapest, Hungary In species with prolonged breeding season groups of individuals might reproduce at different 3 Environmental Consultants Ltd., Kresz G. u. 18, 1132 Budapest, Hungary time periods. Gene flow could be restricted between early and late reproducers even within a population in the same habitat when the reproductive time period is heritable. This restric- The fire salamander, Salamandra salamandra was thought to be extinct for the last 100 years tion of gene flow caused by reproduction in different time periods is called “isolation by from the Buda Hills. In 2008 a small population was rediscovered in Budapest in an isolated time”. The reproductive period of the critically endangered yellow-bellied toad (Bombina habitat along a stream surrounded by family houses. The goal of our study was to measure variegata) in Germany starts in April and runs until September. The long reproductive peri- the sex ratio, age classes, body size pattern, seasonal activity, migratory movement, and larval od allows that groups of yellow bellied toads might be behaviorally and genetically isolated development characteristics of the population. from each other due to isolation of time, as it was found in other amphibian and vertebrate The study was performed between 2008–2012. The juvenile and adult salamanders were species. For this study we collected buccal swabs at four different time points in two consec- surveyed after rain or the day after, from March to November. We identified the sex, meas- utive breeding seasons from one population of B. variegata in southern Lower Saxony. Using ured the body size, body weight and saved the GPS coordinates of the exact location of every ten highly variable molecular markers (microsatellites) we measured the genetic differences specimen. Individuals were identified by the colour pattern that is characteristic to every within and among the different groups of toads. Here we report our first results of this study specimen. Larvae were surveyed every 10 days between April and September, on a 500 m long and discuss the importance of our findings for the conservation of B. variegata. section of the stream, during the night, using flashlight. From 2008 we caught 1860 specimens (juvenile or adult). In the last 3 years adult sex ratio was male-biased and varied between 1:1.1 and 1:1.3. The average body size did not differ between males and females, but body weight was higher in females. We defined three age classes: recently metamorphosed, juveniles (1–3 years), adults (over 3 years). Half of the pop- ulation consisted of adult specimens. Every year we observed one activity peak in spring and one in autumn. The sampling success and number of captured individuals was concordant with the quantity of rainfall. According to the capture-recapture locations we tracked the movement pattern of individuals. The movement of the most commonly recaptured individuals suggested that salamanders in this population use a 869–1125 m2 area. The females started to deposit their larvae at the end of March and finished within a week. Due to heavy rainfall in 2011 the number of larvae reduced with 44%. The larvae were inhabited mostly those sec- tions of the stream where small pools were formed.

248 249 BEHAVIOUR MONITORING

Escape strategy of Schreiber’s green lizards (Lacerta schreiberi) Photographic method for the European snake-eyed skink is determined by simple environmental factors but not ecological (Ablepharus kitaibelii fitzingeri) individual identification context or sex Dániel Kovács, István Kiss Renáta Kopena 1, Gábor Herczeg 2, José Martín 1 Department of Zoology and Animal Ecology, Szent István University, Páter K. u. 1, 2100 Gödöllo˝, Hungary; [email protected] 1 Departamento de Ecología Evolutiva, Museo Nacional de Ciencias Naturales, CSIC, José Gutiérrez Abascal 2, 28006 Madrid, Spain; [email protected] 2 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, The snake-eyed skink occurs only in the Carpathian Basin. Its main distribution area is located Pázmány Péter sétány 1/c, 1117 Budapest, Hungary in Hungary. Due to its secretive life and under-researched status, our knowledge about its life, history and population biology is quite deficient. It is essential to develop a harmless, Antipredator behaviour is a key behavioural component of animals’ fitness. Antipredator escape safe identification method for surveys. Until now, there had not been reliable methods for behaviour shows great variation not only between, but also within species or even within the individual identification of this subspecies. populations and individuals. One well-established source of variation is the physical envi- Our object was to test the I3S (Interactive Individual Identification System) software utility ronment, including factors such as the distance to the nearest. Another important source of in the identification of the European snake-eyed skink. The photographic capture-recapture variation is the ecological context such as being in reproductive vs. non-reproductive state. method has previously been successfully used in several surveys. The identification procedure However, the relative roles of the above sources are rarely assessed within the same study. assumes that the shape and size of the reptile scale patterns are various with unique disting- Here, we studied the escape strategy of Schreiber’s green lizards, Lacerta schreiberi. We meas- uishing marks and the probability of finding two identical ones is low. ured the distance that lizards allowed a potential simulated predator to approach before flee- Field sampling was carried out during the summer and autumn of 2012. We took photo- ing in the field (approach distance) to test whether the physical environment, the ecological graphs of the supralabial scales on each individual. The use of the software is based on pointing context, or their interaction are the most important determinants. The environment had out a minimum of twelve connection points among scales. In total, 25 Ablepharus kitaibelii strong direct effects on the lizards’ escape strategy; approach distance showed strong positive fitzingeri individuals were caught. We collected 84 images to a database and we were able to correlation with distance to refuges on grass substrates, whereas no such correlation was recapture 13 individuals. observed on rocky substrates. Further, refuge distance and the escape angle had a complex, We concluded that the unique size and shape of the supralabial scales are suitable for indi- but substrate-independent, effect on escape strategy: either short refuge distances or high vidual identification. The requirements for application are good quality photos, and the pre- escape angles resulted in short approach distances. In contrast, neither season (reproductive cise digitalization of the pictures. The accuracy of the identification was 92%. We could elimi- vs. non-reproductive), nor sex had any effect on the escape strategy. Our results suggest that nate the program errors by visual comparison of the unique shape and size of the supralabial the escape strategy of Schreiber’s green lizards is determined by the environmental settings, scales’ pigmentations. We found that the supralabial pigmentations of the recaptured indi- irrespective of the ecological context or sexual roles. viduals did not change in shape after shedding, or we observed very slight differences. A combi- nation of the software and pigmentation identification increased the accuracy of the iden- tification of individuals to 100%.

250 251 MONITORING PRACTICAL CONSERVATION

Mapping Budapest herpetofauna using GIS techniques Czech action plan activities for Zamenis longissimus

Tibor Kovács1,2, Ciprian Samoila 3 Antonin Krasa

1 Amphibian-Reptile Conservation Group, MME-BirdLife Hungary, Költo˝ u. 21, 1121 Budapest, Hungary; Nature Conservation Agency of the Czech Republic, Kaplanova 1, 14800 Praha, Czech Republic; [email protected] [email protected] 2 Hungarian Biodiversity Research Society, Hunyadvár u. 43/a, 1165 Budapest, Hungary 3 Ovidius University Constant¸a, Faculty of Natural and Agricultural Sciences, Aleea Universita˘¸iit 1, corp B, Zamenis longissimus is the rarest snake in the Czech Republic. There are three distinct popu- 900470 Constant¸a, Romania lations which differ in the abundance, area and isolation from other populations. Best known Budapest harbours a remarkable herpetofauna including 18 amphibian and reptile species. is the smallest and most isolated one living in Poohrˇí in West Bohemia. The biggest one lives The territory of the Hungarian capital contains extended semi-natural or natural and pro- in Podyjí National Park in South Moravia. Third one lives in Bílé Karpaty in East Moravia tected areas. Cultivated fields, grasslands, wetlands and forests cover more than 25% of the on the border with Slovakia. All of them are strongly related to human activities such as small city and the proportion of gardens and parks is also significant. This relatively airy archi- scale agriculture, non intensive forestry and keeping of animals. tecture allows the persistence of amphibian and reptile species occupying different habitat Action plan (AP) for this species was approved by Ministry of the Environment of the CR types and so raises both their numbers and species richness compared to other big cities. The in 2008. Maintenance of special biotopes, migration corridors and breeding sites and protec- Amphibian and Reptile Conservation Group of MME-BirdLife Hungary initiated a survey tion of individuals on and near roads are the most important activities of AP. Of course, program on Budapest herpetofauna in 2010 involving six guided and couple of dozens regular monitoring, searching new potential localities and diverse research are not missing. unguided volunteers who uploaded their observations on our herp-mapping portal www. Most diverse spectrum of activities is used in Poohrˇí where 400–600 individuals are living herpterkep.mme.hu. in area of only 970 ha. This population faced two main problems which have been solved. A total of 1598 occurrence records were exported from the online portal and imported in a There is a main road where a lot of snakes were dying, particularly juveniles. The most dan- file geodatabase using ArcGIS Desktop 10.1 for mapping and spatial analysis. Species richness gerous part has been modified and special device has been installed there, so that snakes are or number of records were calculated in a 1 x 1 km grid covering the Budapest city boundaries. not able to move to the road and they have to go through holes going under the road. The Highest values of species richness (7–10 species) occurred in 8 grid cells, all located in second problem was deficiency in natural breeding facilities, which is why 26 artificial ones forests or wetlands. The most common reptile species was the common wall lizard (Podarcis have been built to enhance reproduction success. 2 muralis) occupying densely built up areas too, even in the downtown or bare blockhouse There are 3000 to 4000 individuals living in Podyjí (77 km ), where the most important estates. Among the amphibians, the green toad (Bufo viridis) showed a similarly high level of activity is clearing of special biotopes (old watermills, agricultural terraces and wineyards). urbanization, although its observed number was considerably less. The green lizard (Lacerta Area of this national park was formerly strongly influenced by man, but after WWII forest viridis) was also a dynamic conqueror intruding suburban gardens. Even the Natura 2000 has regrown it what causes problems today. In Bílé Karpaty population size and inhabited species European pond turtle (Emys orbicularis) formed stable populations within the bound- area are not clear yet. Thus we concentrate on gaining more information there. aries of Budapest. Further spatial analysis and predictive modelling is planned to estimate the potential dis- tribution of herpetofauna species in Budapest.

252 253 DISEASES AND PARASITES POPULATION GENETICS

Amphibian egg mortality in the Bialowiez˙a Forest, Poland Species composition of Triturus populations in a three way contact in the Waldviertel (Lower Austria) Renata Krzys´ciak-Kosin´ska 1, Miroslawa Orlowska 2 Patricia Lagler 1, Pim Arntzen 2, Günter Gollmann 3, Ben Wielstra 2 1 Bialowiez˙a National Park, Research Unit, Park Palacowy 5, 17-230 Bialowiez˙a, Poland; [email protected] 2 Department of Biology, Medical University of Bialystok, Kilin´skiego 1, 15-089 Bialystok Poland 1 University of Natural Resources and Life Sciences, Gregor Mendel Straße 33, 1180 Wien, Austria; [email protected] The Bialowiez˙a Forest is the last remnant of lowland deciduous forest of primeval character 2 Naturalis Biodiversity Center, P.O. Box 9517, 2300RA Leiden, The Netherlands in Europe. Central part of the forest is strictly protected for over 80 years and human activity 3 University of Vienna, Department of Theoretical Biology, Althanstraße 14, 1090 Wien, Austria is very limited. Selected water bodies were monitored in the spring during the breeding sea- son of two most common in the Bialowiez˙a Forest amphibian species: Rana temporaria and The crested newt Triturus cristatus superspecies consists of seven members and is distributed Rana arvalis. Sites were visited every second day to assess the survival of embryos. At each site in western Eurasia. The mostly parapatric species distributions are delimited by sharp contact there were ten randomly selected egg clumps which were studied carefully for visible changes. zones. In such zones, hybridisation and mitochondrial DNA introgression occur. Lower Number of developing eggs and those which failed to develop in each of selected clumps were Austria is characterized by a transition of climate and biota. Here, three Triturus species meet: counted. The study shown that up to 86% of eggs failed to develop. the great crested newt (T. cristatus), the Danube newt (T. dobrogicus) and the Italian newt (T. Samples of eggs were taken to the laboratory. As the substratum for growing fungus-like carnifex). organisms, cannabis seeds and eggs of R. temporaria and R. arvalis were used. After 72 hours In the Waldviertel, the north-western part of Lower Austria, Triturus species distribution is of incubation in the temperature of 15 °C, the first microscope analysis were carried out. still unclear. In general, morphology is quite reliable for species identification but in contact Then, the samples of mycelia were transported onto Sabouraud [bioMerieux] substratum zones mixed phenotypes appear, due to hybridisation, which makes this method less useful. where the culture continued to grow. The samples were examined every seven days. The The aim of our study is to clarify the situation of species composition and hybridisation. experiments were carried out for three weeks. Identification of fungus-like organisms was Therefore tissue samples of 93 individuals from 6 populations (covering the Waldviertel from based on their morphology and biometric data of antheridia and oogonia. There were seven the East to the West) were collected. Next to morphological examinations, genetic analyses Saprolegnia species isolated from the collected material. These were Saprolegnia anisospora, were done, using 52 nuclear markers and one mitochondrial DNA marker. S. diclina, S. ferax, S. hypogyna, S. monoika, S. parasitica, and S. uliginosa. The most often Based on morphology, the Triturus newts mostly resemble T. carnifex. However, only T. isolated species was Saprolegnia ferax which was present in all analysed samples. dobrogicus (eastern) or T. cristatus (western situated populations) mitochondrial DNA haplo- It is presumed that fungi-like organisms contributed to the high mortality of amphibian types are present. These results are a clear indication that hybridisation must have occurred eggs as there are several species of Saprolegniaceae reported so far as causing mortality of in the recent history of these populations. Analysis of the nuclear markers provides a more amphibian eggs but two species are most frequently blamed: Saprolegnia parasitica and S. detailed insight into the gene flow among the three Triturus species and allows us to formu- ferax. late a historical biogeographical scenario.

254 255 PHYLOGENY ECOLOGY

Systematics of the ‘flap-headed frogs’ of the dicroglossid genus Phenotypic variability in the Pelophylax saharicus complex along an Limnonectes Fitzinger, 1843 altitudinal gradient in the region of Marrakech and its surroundings (Morocco). Discriminant analysis and implications in terms of Markus Lambertz 1, Timo Hartmann 2, Shannon Walsh 3, Peter Geissler 2, conservation David S. McLeod 3 Aziza Lansari 1, Tahar Slimani 1, El Hassan El Mouden 1, Alain Pagano 2 1 Institut für Zoologie, Rheinische Friedrich-Wilhelms-Universität Bonn, Poppelsdorfer Schloss, 53115 Bonn, Germany; [email protected] 2 Sektion Herpetologie, Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany 1 Cadi Ayyad University, Faculty of Sciences, Department of Biology, Laboratory Biodiversity and Ecosystem 3 University of Kansas, Biodiversity Institute, 1345 Jayhawk Blvd., Lawrence, Kansas 66045, USA Dynamics, P.O. Box 2390, 40000 Marrakech, Morocco; [email protected] 2 University of Angers, Campus Belle Beille, 49045 Angers, France Most of the 55 currently recognized species of Limnonectes (Ranoidea: Dicroglossidae) are rather unremarkable, medium-sized brown frogs. The males of four species (L. dabanus, L. The influence of climate and space global change has been studied by comparing the size and gyldenstolpei, L. macrognathus, and L. plicatellus), however, possess a conspicuous dorsal orna- color morphs of the North African green frog (Pelophylax saharicus Boulenger, 1913) in mentation of their heads. To date, no study has examined this structure in detail. The goals Morocco. We examined the morphological variation of four populations subject to different of this study were to address the systematics of the flap-headed species and assess the func- environmental variables in the region of Marrakech and its surroundings. The processing of tional and evolutionary significance of this feature. Morphological analyses of these struc- morphometric data using the univariate (ANOVA) and multivariate (ACP and AD) statis- tures (via gross anatomical, histological, and radiographic means) in all four species revealed tical analysis, revealed the existence of at least two morphotypes of this species which are great differences in external characteristics such as shape and extent. Histological architec- clearly separated by the ACP. The use of discriminant analysis determined that the most ture, however, was nearly identical in all species, suggesting that the appendages indeed could discriminating variables are the first toe of the left foreleg and the length of the femur of the have a common evolutionary origin. Additionally, we found that all four species have a con- posterior leg of the same side. cave lateral curvature of the anterior ramus of the pterygoid bone, which is convex in all other Overall, frogs from medium and high elevation sites (respectively Oukaimeden and Tigha- Limnonectes. Results of molecular phylogenetic analyses of the 12S–16S mtDNA gene region douine) are larger than those from the Atlantic coast (Had Adra). Many various coloration provided well-supported evidence for the monophyly of this group. Based on these two inde- patterns were noted in the different surveyed sites. These variations in size and color may cor- pendent lines of evidence, we conclude that these four species represent a monophylum and respond to a phenotypic plasticity in response to the instability of the habitat and therefore that the head appendage is a reliable diagnostic feature of this group. a substantial taxonomic differentiation like what is observed in Europe, and thus could be of considerable conservation interest from which this interest in arid areas where this type of study, in our knowledge, done before.

256 257 PHYSIOLOGY MONITORING

Habitat effect on the digestive efficiency of a Mediterranean lizard Spatial and temporal patterns of the ladder snake (Rhinechis scalaris) in a human modified landscape: preliminary results from Victoria Litsi-Mizan 1, Emmanouela Karameta 1, Konstantinos Sagonas 2, a radio-telemetry monitoring program in eastern coastal Spain Efstratios D. Valakos 2, Spyros Sfenthourakis 3, Panayiotis Pafilis 1 Marcial Lorenzo 1, Fernando Martínez-Freiría 2, Miguel Lizana 1 1 Section of Zoology and Marine Biology, Department of Biology, University of Athens, Panepistimioupolis, Ilissia, 57-84 Athens, Greece 2 Section of Animal and Human Physiology, Department of Biology, University of Athens, Panepistimioupolis, Ilissia, 1 Departamento de Biología Animal, Universidad de Salamanca, Campus Miguel de Unamuno, 37007 Salamanca, 57-84 Athens, Greece Spain; [email protected] 3 Department of Biological Sciences, University of Cyprus, University Campus, P.O. Box 20537, 1678 Nicosia, Cyprus; 2 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto, [email protected] Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal

The particular ecological and environmental features of a given habitat have a direct impact The ladder snake (Rhinechis scalaris) is a Mediterranean snake, widely distributed in the on numerous aspects of the overall animal biology. Among them thermal quality and pre- Iberian Peninsula. Its secretive behaviour has hampered a good knowledge of populations’ dation pressure play a pivotal role in shaping the performance of physiological traits such as ecological patterns which were inferred using radio- telemetry monitoring but for limited digestion. The later represents a measure of successful survival and, especially in ectotherms, periods over the annual cycle of activity (e. g. in spring). Here we present the preliminary has been proved to be strongly temperature dependent. Digestive efficiency is an important results of spatial and temporal patterns for four specimens (two males and two females) link between the gut physiology and the ecological aspects of energy budgeting. Apparent inhabiting a human modified landscape (mostly consisting of citric cultures, area = 132 ha) in digestive efficiency (ADE) is defined as the ability to absorb energy through food and eastern coastal Iberia (Castellón, Spain) throughout an annual cycle of activity (2012–2013). depends on gastrointestinal motility, enzymatic activity and body temperature. Snakes (avg. SVL=986.5 mm; avg. weight=558.75 g) were captured in spring 2012 and In this study we aimed to clarify whether habitat quality (in terms of temperature and pre- radio-transmitters (SOPI 2380; weight=10 g) were implanted by surgery (assisted by a veteri- dation regime) affects digestive efficiency in the roughtail rock agama (Laudakia stellio). We narian). After 5 days of post-surgery, snakes were released and radio-telemetry monitoring presumed that environmental temperatures (as they are sketched by operative temperatures) was performed 20 days per month till the end of March 2013. A total of 817 records were and predation would induce shifts in ADE. ADEs of the three major nutrient components obtained, including GPS coordinates and data on habitat and micro-habitat composition (proteins, lipids and sugars) were examined in 20 individuals from two eastern Mediterranean and activity parameters of snakes. Geographical Information Systems were used for inferring islands differing in climate conditions and predation pressure, Cyprus and Naxos (Cyclades, spatial patterns whereas linear regression analyses were performed for habitat descriptors. Aegean Sea). Higher temperatures have been repeatedly reported to favor effective digestion. Snakes were mostly found on citric cultures (48.95% of records) along the year, without On the other hand predation restricts the time dedicated to foraging and thus ADE should differences among individuals and sexes. Selected micro-habitats for the four individuals were be high to ensure that animals would take the most of every meal. mostly composed by herbaceous vegetation in the surroundings of stone walls, but each indi-

ADElipids and ADEsugars did not differ between the two populations. ADE for proteins was vidual used different micro-habitats according to the period of year (log-lineal; p =0.98). marginally higher in the case of Cyprus and the difference disappears if excluding one indi- Home range sizes were different for males (kernel estimator =1.48 ha) and females (kernel vidual from analyses. Both populations are well adapted in their habitats and achieved high estimator = 0.53 ha). Movement rates revealed a winter dormancy with low movement digestive performance. Future experiments that will focus on gut passage time will reveal the (December–January: 1 m/day) and several peaks of activity February–March (69.63 m/day), full repertoire of digestive procedure. June–July (95.9 m/day) and October–November (56.8 m/day). Considering that these results are a preliminary approached, new data is shown about the ecology of this species, especially those related with spatial and temporal patterns along whole one year.

258 259 BEHAVIOUR ECOLOGY

An experimental approach for testing the role of olfactory and visual Activity patterns and habitat preference of European pond turtles stimuli in snakes climbing towers behaviour (Emys orbicularis L.) in Lake Naplás

Marcial Lorenzo 1, Pablo Garcia-Diaz 1,2, Fernando Martínez-Freiría 3, Miguel Lizana 1 Zsófia Lovász 1, Tibor Kovács 2 , Péter Sály 3, István Kiss 1

1 Departamento de Biología Animal, Universidad de Salamanca, Campus Miguel de Unamuno, 37007 Salamanca, 1 Szent István University, Department of Zoology and Animal Ecology, Páter K. u. 1, 2100 Gödöllo˝, Hungary; Spain; [email protected] [email protected] 2 School of Earth and Environmental Sciences, The University of Adelaide, Adelaide, South Australia 5005, Australia 2 Department of Animal Taxonomy and Animal Ecology, Eötvös Loránd University, Pázmány P. s. 1/c, 1117 Budapest, 3 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidadedo Porto, Hungary Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão, Portugal 3 HAS Centre for Ecological Research Balaton Limnological Institute, Klebelsberg Kunó u. 3, 8237 Tihany, Hungary

Ladder snakes (Rhinechis scalaris) are suffering an unusual non-natural mortality due to electro- Lake Naplás of Budapest is a remarkable habitat of the European pond turtle in spite of the cutions in the largest electricity network of Spain (Iberdrola S. A.).Research has shown spatio- fact that its vicinity is highly disturbed by human activities. The goal of our investigation temporal patterns in electrocutions consistent with the hypothesis that the snakes climb to was to count and estimate the number of specimens living in the lake and the surrounding medium tension power towers (MPVT) attracted by prey’s stimuli. However, experimental swamp, to study sex ratio, seasonal activity, fidelity to locations where trapping took place works are required to confirm this hypothesis. Here, we used a captivity experiment for and recapture rate. We intended to reveal the relationship between these variables and the studying ladder snakes’ preferences (28 individuals) for two 3 m replicas of MPVT (one con- morphometric data of the captured specimens. trol and the other treatment) with different attractants: prey odours (olfactory detection), In 2008, we trapped turtles at 21 occasions using 4 cage-traps at each trial while in 2009, preys (visual detection), heat emitters (heat detection, simulating wires from MPVT) and dif- we used traps with changing numbers at 18 occasions. In 2008, we collected 146 and in 2009, ferent combinations. Snakes’ body size and activity rhythms were also analysed. 73 turtles. Altogether 113 specimens were identified, out of which 69 were females and 36 Ladder snakes climbed significantly more often to the MVPT with prey odour in light con- males. The recapture rates were low, the number of turtles recaptured once was 7 in 2008 and ditions (hurdle GLM model: p<0.03), but there were no significant differences regarding 6 in 2009 while multiple recaptures were rarer. Mainly males were recaptured. other attractants and conditions. Moreover, activity of the snakes had significant effects (hurdle The estimated size of the population was 129 specimens (95% confidence interval ranged GLM model: p<0.01 in both cases). Additionally, the proportion of snakes that climbed to between 72–289). The hypothesis that smaller and therefore younger individuals showed the treatment MPVT was significantly higher for prey odour treatment in either light or dark higher recapture rate could not be proven. The average carapax length of females was 157.2 conditions (Binomial test: p<0.05 in both cases). Finally, there were no statistical significant (± 42.4 SD) mm while that of the males was shorter 138 (±18.6 SD) mm. Also the average patterns in the time since the beginning of the experiment to the first climbing event to the bodyweight was bigger in females (mean ±SD: 643.6 ±330.2 g) than that of the males (mean± treatment tower for any of the conditions and treatments (Kruskal-Wallis test: p=0.36). SD: 416.5± 142.4 mm). These results support the hypothesis of an olfactory attraction to MPVT. However, how Trapping seemed to prove that most of the turtles occurred in the willow swamp habitat snakes can detect odours from birds located at 16 m height is still unknown. patch and in the sunny edge of the reed bed. These habitat patches were used mainly by the smaller individuals. The majority of the turtles appeared in another habitat only once during the study year and this change was observed in both sexes. The highest activity of the turtles.

260 261 MONITORING ECOLOGY

Three years of population monitoring of newts (Lissamphibia, Habitat features and syntopic species of crested newts ponds in Salamandridae) in isolated karstic ponds South Moravia, Czech Republic

Martina Luzˇnik 1, Boris Krysˇtufek 2 Zdeneˇk Macˇát 1, Lenka Jerˇábková 2, Antonín Reiter 3, Martin Rulík 1

1 Faculty of Mathematics, Natural Sciences and Information Technologies; University of Primorska; Glagoljasˇka 8, 1 Department of Ecology and Environmental Sciences, Palacky´ University, Slechtitelu˚ 11, 78371 Olomouc, Czech 6000 Koper, Slovenia; [email protected] Republic; [email protected] 2 Science and Research Centre, University of Primorska, Garibaldijeva 1, 6000 Koper, Slovenia 2 Nature Conservation Agency of the Czech Republic, Kaplanova 1931/1, 148 00 Praha 11, Czech Republic 3 South Moravian Museum Znojmo, Prˇemyslovcu˚ 6, 66945 Znojmo, Czech Republic We conducted a population study of two syntopic salamandrids, the smooth newt Lissotriton vulgaris and the Italian crested newt Triturus carnifex, at six ponds in Karstic Plateau, SW Crested newt use different habitats throughout their lives; knowledge about habitat features Slovenia. Both newts, which are classified as “vulnerable” in the national Red list, are partic- are useful and important for their continued protection. Previous investigations proved coex- ularly under threat in the study area due to rapid habitat loss in karstic landscape which lacks istence and hybridization among three crested newt species: Triturus cristatus, T. carnifex and the surface water outlet. A detailed CMR survey was conducted in a pond in the village T. dobrogicus. We studied ponds features and species richness of amphibian populations in 31 Kastelec during breeding seasons 2006–2008 and population abundance was estimated using breeding ponds in the Czech region of Znojmo. Data for analysis was collected during sea- model POPAN available in MARK. Abundance in further five ponds with less intense field sons 2010–2012, during which time we recorded eight habitat features: pond area (<100, sampling was evaluated using the Petersen index. Populations of both species, smooth and 100–1000, 1000–5000, >5000 [m2]), maximum depth (<30, 30–100, >100 [cm]), fish presence Italian crested newts, were most abundant at Kastelec pond. Over three years of survey, the or absence, origin (natural or man made) and presence or absence of human use. Also we abundance of smooth newt was quite stable at about 300 males and 300 females which sug- assessed density of submerged vegetation and shade (both in 25% class). Each pond was cate- gest this population to be above the minimal viable population (MVP) threshold. The pop- gorized on type (natural pond, urban pond, quarry, stagnant ditch). The effect of this eight ulation of Italian crested newts at Kastelec was estimated at about 150 individuals in 2007. pond features on crested newt pond occupancy and frequency of syntopic species was calcu- The number of captured individuals in the remaining years was too low for statistically lated. According to our results, pond depth between 30–100 cm, shade between 26–50%, robust population assessments. Populations of the Italian crested newt in karstic ponds type of pond and area between 100–1000 m2 was evaluated as the essential parameters increasing allegedly do not meet the MVP threshold which may pose risk on their long-term persist- attractivity of the ponds for the occupancy of adult crested newts. The smooth newt (Lisso- ence. Populations of both species are further threatened by introductions of fish in some of triton vulgaris) and agile frog (Rana dalmatina) were recorded at 22 and 20 ponds, respectively, the ponds. An ongoing population genetic study based on microsatellite loci is expected to together with crested newt. Our conclusions about optimal habitat features of pond from the shed light on the metapopulation structure of newts in a system of karstic ponds. The final point of view of crested newts correspond in general with results of similar studies. Unlike to goal is to better define conservation status of the two newt species in an area, which is hos- other studies, fish presence and absence of submerged vegetation were found to be the major tile to amphibians with limited dispersal potential. causes for population density of crested newts. It seems that the population structure of syn- topic species is unique for different areas within distribution of crested newt. Further data collection and analyses are required to support the outcomes of this study.

262 263 POPULATION GENETICS ECOLOGY

The genetic mating system of Hyalinobatrachium valerioi Spatial distribution and population density of Pelophylax perezi (Ranidae: Anura) in Tablas de Daimiel National Park (Central Spain) Alexandra Mangold, Katharina Trenkwalder, Walter Hödl, Max Ringler, Eva Ringler Ignacio Martín, Oscar Rodríguez de Rivera, Isabel Angulo, Yaiza Vélez Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; [email protected] Zoology Department, School of Forestry and Natural Environment Engineering, Polytechnic University of Madrid, Avda. Ramiro de Maeztu s/n, 28040 Madrid, Spain; [email protected] The reticulated glass frog Hyalinobatrachium valerioi (Centrolenidae) is a small leaf-breeding frog which occurs in the tropical rainforests of Costa Rica, Panama, Columbia and Ecuador. The present work establishes the population density and spatial distribution of Pelophylax There are a few studies focusing on the reproductive behavior of this frog species, but to date perezi in Tablas de Daimiel National Park, Ciudad Real (Central Spain). The project was pro- the genetic mating system has not been resolved. The H. valerioi population under study moted by the Management of the National Park, and it is related with I+D+I initiatives of occurs along the Quebrada Negra, a small lowland stream close to the tropical research sta- the Department National Park belonging to the Spanish Ministry of Environment. The field tion La Gamba, at the edge of the Piedras Blancas National Park, Costa Rica. Field data and data were obtained through several “listening stations” carried out during 2011 and 2012. The samples were collected from August to November 2012, covering almost one entire breeding samplings were done in March, April, May and June. May was divided into another two peri- season. Along a 400-m river transect, we monitored 144 individual frogs (96 males, 48 ods “May 1” and “May 2”, due to the high amount of activity, doing samplings every 15 days. females) during the whole study period and took tissue samples of all adult frogs and clutches For this specie the sex-ratio is established by 1.9:1 in favor of males. Similar results have been for the subsequent molecular analyses. The integration of observational field data as well as obtained in different points of the country. For the data processing is used the following 2 molecular parentage analyses allowed us to gain precise insights in individual mating pat- expression D=Ln(n/n1)*n/m(r ), with D being the number of individuals/ha, n the number

terns, the adult sex ratio, potential reproductive rates, the effective number of breeders and of individuals found in the sampling plot, n1 the number of individuals found out of the samp- also data on site fidelity of a H. valerioi population. ling plot, m the number of plots in each station and r the radius of each of it. Using the cor- relation coefficients we could observe the amount of relationship between the different variables and the density of P. perezi. However, this only indicates a slight influence of any of these variables. For a deeper understanding, the use of statistical models, particularly regres- sion models, could help us to make predictions of the presence and density of that species. Furthermore, these models are useful to analyze which variables alone or related have influence over the presence and abundance of P. perezi. In case of not having enough record inference it could be the best option to find a reliable data distribution, augmenting to simulate and obtain good predictions and strong understanding of the environmental variables.

264 265 ECOLOGY ENPARTS

Population density and spatial distribution of Hyla arborea Amphibian road mortality: comparing spatial statistical methods (Hylidae: Anura) in Tablas de Daimiel National Park (Central Spain) to detect hotspots

Ignacio Martín, Oscar Rodríguez de Rivera, Isabel Angulo, Yaiza Vélez Cátia Matos, Neftalí Sillero, Elena Argaña

Zoology Department, School of Forestry and Natural Environment Engineering, Polytechnic University of Madrid, Centro de Investigação em Ciências Geo-Espaciais (CICGE), Universidade do Porto, Faculdade de Ciências, Avda. Ramiro de Maeztu s/n, 28040 Madrid, Spain; [email protected] Observatório Astronómico Prof. Manuel de Barros Alameda do Monte da Virgem, 4430-146 Porto, Portugal; [email protected] The present work establishes the population density and spatial distribution of H. arborea in Tablas de Daimiel National Park, Ciudad Real (Central Spain). The project was promoted Due to their habitat requirements, amphibians became one of the most commonly road- by the Management of the National Park, and it is related with I+D+I initiatives of the killed group and this may affect their population viability. Implementation of mitigation Department National Park belonging to the Spanish Ministry of Environment. The field measures (e. g. crossing structures) may overcome the problem, still several factors influence data were obtained through several “listening stations” carried out during 2011 and 2012. measures effectiveness. Crossing structures location is the most influential factor forecasting The samplings were done in March, April, May and June. May was divided into another two effectiveness. However, road network density and dimension requires a cost-effective appli- periods “May 1” and “May 2”, due to the high amount of activity, doing samplings every 15 cation and a better understanding in where they should be implemented. As amphibians days. For this specie the sex-ratio is established by 1.66:1 in favor of males. For the data pro- present a clustered road-kills distribution, their mortality hotspots can be identified using 2 GIS (Geographic Information Systems). Hotspots or critical sites can be predicted using spa- cessing is used the following expression D=Ln(n/n1)*n/m(r ), with D being the number of tial statistical methods that are available and have been tested in several road-kill studies. individuals/ha, n the number of individuals found in the sampling plot, n1 the number of individuals found out of the sampling plot, m the number of plots in each station and r the Nonetheless it is lacking a comparison analysis of them in order to understand their pros and radius of each of it. Using the correlation coefficients we could observe the amount of rela- contras. The aim of this study was to analyse possible differences between global, multi-scale tionship between the different variables and the density of H. arborea. However, this only and local spatial analysis methods in defining hotspots using amphibian road fatality data indicates a slight influence of any of these variables. For a deeper understanding, the use of collected in northern Portugal country roads. Nearest neighbor index, Moran’s I were calcu- statistical models, particularly regression models, could help us to make predictions of the lated in order to compare the global clustering of points in seven sampled roads, and three presence and density of that species. Furthermore, these models are useful to analyze which were identified as clustered. We used Ripley K-function the best scale for Malo’s equation, variables alone or related have influence over the presence and abundance of H. arborea. In Kernel density analysis in detecting hotspots and we compared their detection performance case of not having enough record inference it could be the best option to find a reliable data with Local Indicators of Association (LISA) (i. e. Local Moran’s I and Getis-ord Gi*). Three distribution, augmenting to simulate and obtain good predictions and strong understanding different GIS software applications were used: ArcGis, Quantum GIS with R (opensource) of the environmental variables. and GeoDa (opensource). Results showed the importance of using multi-distance spatial cluster analysis to define the best scale for hotspot detection with Malo’s equation and Kernel density analysis. Here we also suggest the advantages of Local Indicators of Association (LISA) for detecting clusters with the contribution of each individual observation (Local Morans I and Getis-ord Gi*).

266 267 PRACTICAL CONSERVATION ECOLOGY

Educational work and protection of salamanders in three European Ant predation by the wall lizard (Podarcis muralis) countries within a Sparkling Science project Claude Mermod Magdalena Meikl 1, Enrico Romanazzi 1, Juan Ramón Fernández Cardenete 2, Ana Gimeno Represas 1, Javier Benavides Sánchez de Molina 2, Stefano Doglio 1, Institute of Biology, University of Neuchâtel, 2000 Neuchâtel, Switzerland; [email protected] Alexandra Pitt 1, Ulrike-Gabriele Berninger 1 Podarcis muralis, the wall lizard, is widespread in western and southern Switzerland. It is an opportunist predator, mainly on arthropods, but also on earthworms and small molluscs. 1 Department of Organismic Biology, University of Salzburg, Hellbrunnerstraße 34, 5020 Salzburg, Austria; [email protected] Analyses of alimentary diet show that ants are an important part of the lizard diet. During 2 Asociación Herpetológica Granadina, c/Granada n° 30-b, 18213 Jun-Granada, Spain an eigth year monitoring of a small population (about 200 days of observation per year) cap- ture of winged adults was observed in ten occasions. However, we never ascertained predation The Alpine salamander (Salamandra atra atra), an endemic amphibian species to the Alpine on workers ants. Although several ant species inhabit the area, this behaviour was directed regions across Europe, and the fire salamander (Salamandra salamandra salamandra), the best only against one species, Diplorhopthrum (Solenopsis) fugax. Workers are very small (1.5–3 mm), known salamander species in Europe because of its eye-catching coloration, are among pale yellow, but winged females and males measure 4 to 6 mm and are black. For most of the threatened species in Austria because their habitats are prone to destruction due to human year, these ants live under stones or pavement. They steal brood of other ant species entering invasions. The golden Alpine or Aurora salamander (Salamandra atra aurorae), the Pasubio their nest and inducing workers to flee by emitting a repulsive substance. Swarming takes salamander (Salamandra atra pasubiensis) and the Lanza’s salamander (Salamandra lanzai) live place between July and September. On these occasions, a great number of workers accompany at the south-eastern and western border of the Alps. Their distribution areas are very small winged males and females above ground, and are very active during about 30 to 90 minutes. and they are threatened by forestry, tourism and illegal collection. The Aurora salamander is Lizards are attracted to the spot of swarming and rush to capture winged individuals. From a priority animal in Europe, listed in Annex II and IV of the European Habitats Directive. three to seven lizards, males, females and youngs participate in this rush, without any show Salamandra lanzai is also listed in Annex IV. The south of the Iberian peninsula houses two of agressive behaviour among them. They often come from more than 20 to 40 meters away, subspecies of the fire salamander, the Penibaetic salamander (S. s. longirostris) and S. s. since most of the observed individuals were not residents of our study site. We suppose that morenica. Salamandra s. longirostris is one of the most localized and scarce amphibians in they are attracted by chemicals (pheromones?) emitted by ants during swarming. All along Europe. It is considered as “Vulnerable” in its distribution area, with very small populations, this predation, lizards are totally indifferent to the presence of worker ants. After the end of some of them totally isolated, owing to the disappearance of its breeding areas. Within the the swarming episode, lizards forage further for some minutes on the spot, before scattering framework of a Sparkling Science project, Austrian, Italian and Spanish herpetologists work and disappearing. together and involve more than 25 schools to bring the knowledge of salamanders into the children’s mind. The children become aware of the importance of salamanders by means of workshops, excursions and exhibitions. With the children’s help, measures will be designed to protect salamanders and their environment. The networking platform for children and sci- entists within the project is www.alpensalamander.eu, where salamander observations all over Europe are collected on a database. We believe that the protection of amphibians and their habitats is only possible by actively involving the public, starting with the youngest ones.

268 269 PRACTICAL CONSERVATION DISEASES AND PARASITES

Effects of burning and grazing on anurans in the Fekete-rét Blood parasite load and individual quality in male European green alkali marsh (Hortobágy) lizards (Lacerta viridis)

Béla Mester 1, Mónika Szalai 2, Miklós Puky 3, Szabolcs Lengyel 1 Boglárka Mészáros 1, Katalin Bajer 2, János Török 1, Orsolya Rita Molnár 3, Gábor Herczeg 1 1 University of Debrecen, Faculty of Science and Technology, Department of Ecology, Conservation Ecology Research Group, Egyetem tér 1, 4032 Debrecen, Hungary; [email protected] 1 Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 2 Körös-Maros National Park Directorate, Department of Nature Conservation and Grants Coordination, Anna-liget 1, 2 Laboratório de Biogeografia e Macroecologia, Universidade Federal do Rio Grande do Norte Centro de Biociências, 5540 Szarvas, Hungary Departamento de Botânica, Ecologia e Zoologia, Campus Universitário, Lagoa Nova, Natal-RN, 59078-900, Brazil 3 MTA Centre for Ecological Research, Danube Research Institute, Jávorka S. utca 14, 2131 Göd, Hungary 3 Department of Biological Sciences, Dartmouth College, 78 College Street, Hanover, NH 03755, USA

Due to pollution, fragmentation, loss of habitats and diseases amphibians have been declined During mate choice, the selecting gender (usually females) use these traits to estimate male since the last three decades. Habitat restoration and conservation management of wetlands quality and choose the male which they can produce the highest quality offspring with. These can have an important role in the protection of amphibians, which may be useful indicators features are called sexual signals and can be connected to individual quality in various ways. to monitor post-restoration success. Our goal was to examine how the controlled burning of One possible mechanism is parasite-mediated sexual selection (Hamilton & Zuk hypothesis), reed and year-round cattle grazing affect the species richness and abundance of anurans in which presumes that parasite infection can influence the elaboration of sexual signals. Anti- Fekete-rét alkali marsh (Egyek-Pusztakócs marsh system) during the spring of 2010. We sur- parasite defence is less expensive for more resistant individuals, so these can invest more energy veyed anurans along five transects each in six differently managed areas: (i) long-burnt (fire into the expression of sexual signals; hence, more intensive signals indicate better individual in 2007) and grazed, (ii) long-burnt and not grazed, (iii) recently burnt (fire in 2009) and quality. Females should therefore prefer males with intensive signals, as this way their off- grazed, (iv) twice burnt and grazed, (v) unburned and grazed and (vi) control area with no spring would also inherit better genes. Previous research showed that male European green management. Twice-burnt and recently burnt areas had significantly higher numbers of species lizards’ (Lacerta viridis) ultraviolet-blue throat colour affects female mate choice (intersexual and individuals than other areas, mainly due to the eradication of reed by recent burning. By selection) and male-male competition (intrasexual selection) positively, as a condition dependent summer, reed grew back in most areas, except in grazed sites, in which density of anurans was complex multiple signalling system. The aim of this study was to test the predictions of the as high as 20,000 individuals per hectare. Late-summer burning of reed provides short-term Hamilton & Zuk hypothesis on male European green lizards. In order to do so, we took effects by creating suitable breeding sites the next spring. In contrast, cattle grazing can have blood samples from 30 adult male green lizards during the breeding season and, after fixation an important role after the re-colonization of reed by maintaining diverse habitats. and Giemsa staining identified the blood parasites under microscope. We found different morphotypes from the Haemogregarinidae family, with prevalence being 96.7%. The infec- tion intensity showed negative correlation with throat and belly colour intensity in line with the predictions of the Hamilton & Zuk hypothesis. Furthermore, we found positive cor- relation between the amount of parasites and body size and condition and negative correlation with the level of asymmetry and number of ticks. This study shows that throat patch colour of European green lizards possibly acts as an honest sexual signal of health state in accordance with the Hamilton & Zuk hypothesis.

270 271 ECOLOGY PHYSIOLOGY

Relationship of Scaevola taccada tree with the nest site selection of Evaluation of metal accumulation and its effect on shell deformations Chelonia mydas at Penang Island, Peninsular Malaysia in the European pond turtle (Emys orbicularis)

Sarahaizad Mohd Salleh 1, Shahrul Anuar Mohd Sah 1, Mansor Yobe 2 Tamás Molnár 1, János Gál 2, Ildikó Benedek 1, István Pálinkás 3, Marcell Molnár 1, Miklós Marosán 2 1 School of Biological Sciences, Universiti Sains Malaysia, Minden, 11800 Penang, Malaysia; [email protected] 1 Kaposvár University Faculty of Animal Science, Guba S. 40, 7400 Kaposvár, Hungary; [email protected] 2 Penang Department of Fisheries, Gelugor, 11700 Penang, Malaysia 2 Szent István University Faculty of Veterinary Science, István u. 2, 1078 Budapest, Hungary 3 Károly Róbert College, Insitute of Environmental Sciences, Mátrai u. 36, 3200 Gyöngyös, Hungary The study site is locate at Penang Island, the northern part of Peninsular Malaysia. The coor- dinates of Kerachut Beach are 5°27'4"N, 100°10'58"E. The objectives are to determine the Turtles have been reported as useful biomonitors of metal contamination in the environ- preferable nest site selection locate between vegetation zone, open beach or grass areas, and ment; especially, carnivorous species may accumulate greater concentrations of chemicals the preferable nest site selection locate between intervals 0–10 m, 10–40 m, and >40 m from through trophic transfer during their long life. The aim of the recent investigation was to esti- high tide line. Research was conducted from December 2009 to August 2010. During the mate the relation of heavy metal contamination and the occurrence of scute anomalies and season, all 43 nests laid were identified by intensive nocturnal survey. Results show that green ulcerative shell disease (USD) in two European pond turtle (Emys orbicularis) populations. turtles preferred to nest in the shade of Scaevola taccada, compared to open beach or grass Turtles were collected in Boronka and Csombárd Nature Protected Areas (N=205 and 62, areas (Kruskal-Wallis: p<0.01), and at interval of 10–40 m from tide line, compare to inter- respectively). The number of scute anomalies and the severity of USD were determined. vals 0–10 m and >40 m (Kruskal-Wallis: p<0.01). 61.9% of the nests were located in the Furthermore, liver and muscle samples from 19 perished turtle bodies were examined by shade of S. taccada, and 85.7% nests located between intervals 10–40 m. We conclude the inductively coupled plasma mass spectrometry (ICP-MS) to evaluate the heavy metals con- reasons are due to vegetation of S. taccada gives a shelter and dark environment, the sand taminations. temperature and relative humidity within the S. taccada are optimum and suitable for green The occurrence of scute anomalies and USD in the Boronka population (N=5.3% and 59%) turtle to nest, which are 29 °C and 0.9–1.0%. The reasons turtle fail to dig nest at open beach was higher than in Csombárd (N=1.6% and 41%). The Cd (2.12±0.5 and 1.49±0.1 mg/kg) are due to nest could easily rupture due to sand lack of binding from roots, disturbances by and Hg (2.27±0.9 and 1.29±0.2 mg/kg) and Zn (45.9±7.8 and 34.6±10.8 mg/kg) content in smooth-coated otter and feral dogs, and expose to human disturbances, while turtle fail to the liver samples was significantly higher in Boronka population while As (0.64±0.5 mg/kg) build nest at the grasses areas are due to the rough, dry, and low humidity of the sand, and was found only in the Boronka population. In case of Pb (3.86±4.9 and 1.43±1.7 mg/kg) and less shelter. Therefore, we suggest that the vegetation of S. taccada should be preserve, pro- Cu (25.4±9.3 and 18.29±2.9 mg/kg) the differences were not significant due to the high devi- tect and increase the plantation to up growing the distribution, and as well as to maintain ations. The sex of the turtles had no significant effect. Based on our preliminary data the the natural habitat. higher occurrence of shell deformations and USD in Boronka population could be the result of higher environmental contamination. Further examinations are proceeded including more neighbouring populations and the examination of the status of turtles’ immune system. The study was carried out in the frame of TÁMOP-4.2.2.A-11/1/KONV-2012-0038 project.

272 273 TROPICAL HERP BIODIVERSITY ECOLOGY

Notes on the herpetofauna of western Bas-Congo, Clutch size and egg characteristics in the Souss Valley tortoise, Democratic Republic of the Congo Testudo graeca soussensis Pieh, 2001 (Testudines: Testudinidae) from a coastal and an arid inland area in West-Central Morocco Zoltán T. Nagy 1, Chifundera Kusamba 2, Václav Gvozˇdík 3,4 Hichami Nawal, Mohammed Znari, Mohamed Naimi, Salwa Namous 1 Royal Belgian Institute of Natural Sciences, Joint Experimental Molecular Unit, Rue Vautier 29, 1000 Brussels, Belgium; [email protected] Laboratory “Biodiversity and Ecosystem Dynamics”, Department of Biology, Faculty of Science, Semlalia, Cadi Ayyad 2 Laboratoire d’Herpétologie, Département de Biologie, Centre de Recherche en Sciences Naturelles, CRSN, Lwiro, University, Bd Prince Moulay Abdellah, P.O. Box 2390, 40000 Marrakech, Morocco; and the Natural History Democratic Republic of the Congo Museum of Marrakech, Cadi Ayyad University, Avenue Allal El Fassi, Marrakech; [email protected] 3 National Museum, Department of Zoology, Cirkusova 1740, 19300 Prague, Czech Republic 4 Department of Environmental Sciences, Biogeography, University of Basel, Klingelbergstraße 27, 4056 Basel, Switzerland Reproduction in female Souss Valley tortoises (Testudo graeca soussensis), the family Testu- dinidae, in two populations from a coastal (North to Essaouira) and arid-inland (Central While we have fairly comprehensive knowledge on the herpetofauna of Europe, North America Jbilet mountains) areas of West-Central Morocco, was studied in April–May 2012 using oxy- and some other temperate zones of the Earth, data deficiency in tropical regions is still a major tocin to induce egg deposition in individuals (n = 5 and 4 females with eggs, respectively). problem in herpetofaunistic research. During the last few years we conducted several field Clutch size varied from 1 to 5 eggs with no significant difference between the two popula- expeditions to Central Africa, including the Congo Basin. In 2012 we visited four sites in the tions and averaged 3.44±1.33 eggs. Carapace length and mass of females, which were lower in westernmost part of the Democratic Republic of the Congo. These included the Mangroves the females from arid-land (172±4.4 vs. 185.1±7.7 mm, and 1157.1±91.7 vs. 1486.9±223.4 g), National Park in the region of the Congo River estuary as well as the Mayombe region char- were not significantly correlated neither with clutch size nor with any egg variable in both acterized by low mountains covered by rainforest, all north of the Congo River, and an addi- populations. Eggs averaged, respectively in the arid and coastal populations, 36.24±2.49 x tional site south of the Congo River with savannah habitats. Remarkably, no recent herpeto- 29.99±1.33 mm and 32.06±2.21 x 28.96±0.88 mm in dimensions, 17.05±1.87 and 14.13± logical research was done in this part of the country. Regular visual and acoustic encounter 1.81 cm3 and 19.34±2.14 and 17.28±1.09 g, respectively in volume and in mass. The eggs were surveys were carried out during day and night in order to collect amphibian and reptile spe- significantly longer in the arid-land tortoises. The relative egg masses and relative clutch cimens. We surveyed a wide spectrum of habitats including mangroves, savannah, forests, masses were significantly higher in tortoises from arid-lands (1.70±0.28 vs. 1.15±0.15 and village surroundings and agricultural sites. Approximately 21 species (or representatives of 4.06±1.32 vs. 3.09±1.52, respectively). The obtained data were compared to those on other species complexes) of amphibians and 23 species of reptiles have been recorded. Voucher populations of T. graeca across its distribution range and the differences between the studied specimens are deposited in the RBINS. Tissue samples are used in DNA barcoding and phy- populations are discussed in terms of adaptation to their corresponding environments. logeographic studies focusing the Congo Basin. For some widespread species (e. g. selected colubrid and viperid snakes), remarkable ‘intraspecific’ divergence was observed.

274 275 ECOLOGY ECOLOGY

Niche partitioning in seven syntopic anuran species in a pond in Edge effect on leaf-litter anuran of the Atlantic forest of southeastern Benin City, Nigeria Brazil

S. O. Ogoanah, E. Uchedike Carolina Ortiz 1, Bruna Pereira de Azevedo 1, Katia M. P. Micchi de Barros Ferraz 2, Jaime Bertoluci 1 Department of Animal and Environmental Biology, University of Benin, P.M. B. 1154 Benin City, Nigeria; [email protected] 1 Departamento de Ciências Biológicas, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, Av. Pádua Dias 11, 13418-900 Piracicaba, São Paulo, Brazil; [email protected] Very few diet studies on anurans exist in Nigeria and that only recently. To add to the existing 2 Departamento de Ciências Florestais, Escola Superior de Agricultura Luiz de Queiroz, Universidade de São Paulo, knowledge, the diet of seven syntopic anuran species belonging to four families—Amieto- Av. Pádua Dias 11, 13418-900 Piracicaba, São Paulo, Brazil phrynidae, Arthroleptidae, Dicroglossidae and Ptychadenidae—was determined using sto- mach flushing method. The most important prey category for Hoplobatrachus occipitalis were Anurans are extremely sensitive to abrupt transitions of microhabitats and microclimates. molluscs (V=4545.07mm3) while Ptychadena mascareniensis had arthropod parts (V= 2698.34 Fragmentation and habitat loss threat tropical forests. Much damage found in forest patches mm3). Ptychadena oxyrhynchus and P. bibroni both had coleoptera as most important prey can be attributed to edge effects, which changes the forest structure and faunal composition. (V=3899.96 mm3 and 616.64 mm3), respectively. Leptopelis occidentalis and L. hyloides both We intended to measure the edge effect on leaf-litter anurans of a continuous fragment of had molluscs as most important prey (V=3261.66 mm3 and 1650.99 mm3), respectively. Niche Atlantic Forest in southeastern Brazil. We sampled five transects positioned perpendicularly 2 breadth was highest in H. occipitalis (6.42), a generalist feeder and lowest in L. hyloides (1.6), to the forest edge. Each of them had seven 25 m plots, systematically distributed every 25 m. a tree frog. Amietophrynus maculatus which is a sit-and-wait forager had a niche breadth of Besides, we sampled the anurans along a transect located >1 km from the forest edge. To 2.56 and Hymenoptera as the most important prey (V=1830416.04 mm3). Though living syn- indentify differences in species diversity along the edge distance gradient, the Shannon and topically, niche partitioning is observed based on their food preferences and niche breadth. Pielou indices were used. We assessed the influence of temperature and air humidity on species richness through regression analysis. Preliminary records presented 26 species from seven families. Species richness varied from four (in the edge) to 12 species (75 m from the edge). Species diversity was lower in distance zero (H’= 0.92) and increased considerably only 75 m further in the forest (H’= 1.82). The highest level of diversity was recorded 150 m inside the forest (H’= 2.06), which was higher than that found >1 km far from the edge (H’= 1.86). Low diversity found in the edge is even lower than the patterns found in open areas of other tropical forests. Furthermore, the equability in the edge (J’= 1.52) was lower than 150 m inside the forest ( J’= 1.97), and temperature (p=0.50) and air humidity (p = 0.62) did not correlate to species richness. Our results suggest negative effects on species richness until 50 m from the edge, where just a few species are favored, leading to a simple structure of the anuran community. Thus, the edge effects could influence the spatial distribution of the anurans.

276 277 ECOLOGY PHYLOGEOGRAPHY

Testing new methods to extract skin toxins from common toad Disentangling the Pelomedusa subrufa complex (Bufo bufo) tadpoles Alice Petzold 1, William R. Branch2, Louis Du Preez 3, Leon Meyer 3, 4 5 1 Katalin Pásztor, Anikó Kurali, Zoltán Tóth Alfred Schleicher , Margaretha D. Hofmeyr , Uwe Fritz

1 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, Museum of Zoology (Museum für Tierkunde), Senckenberg Dresden, A. B. Meyer Building, 01109 Dresden, Germany; Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] [email protected] 2 Department of Herpetology, Port Elizabeth Museum, P.O. Box 13147, 6013 Humewood, South Africa 3 Faculty of Natural Sciences, Hoffman Street, 2520 Potchefstroom, South Africa Previous studies investigated the behavioural, morphological and physiological aspects of the 4 Namibia predator-induced phenotypic plasticity, but plasticity in chemical defences—which is thought 5 University of the Western Cape, Bellville, South Africa to be important in amphibians—has been rarely investigated. In order to study plastic responses in chemical defence both qualitatively and quantitatively, and without killing the In our project we investigated the differentiation of the widely distributed African helmeted study subjects, non-invasive methods are needed. With such methods, one may investigate turtle (Pelomedusa subrufa complex) by means of morphological and molecular genetic analyses. the extent of plastic responses accurately and design repeated measure experiments e. g. to Recent studies identified approximately 10 deeply divergent mitochondrial lineages, which examine their fitness costs. In this study, we used a hormone (norepinephrine), hormone-like could represent distinct species. The aims of our research are (1) to correlate possible morpho- substances (phenylephrine and oxymetazolin) and electro-stimulation in vivo to extract skin logical differences with genetic lineages and (2) to unravel the genetic identity of previously toxins from common toad (Bufo bufo) tadpoles and analyzed the quantity and quality of the described taxa to allocate the available names to genetic lineages by using DNA sequences of obtained toxins and investigated the effect of these treatments on individuals’ behaviour and historical type specimens. survival.We also used the standard invasive method, which requires to sacrifice the study sub- ject, as a control, to which we could compare the accuracy of the tested alternative methods. We found decreased survival and reduced activity after the electro-stimulus and oxymetazolin treatments, which implies that these methods are harmful for the tadpoles and therefore should not be used in further experiments. On the other hand, norepinefrine and phenil- ephrin treatments had no adverse effects thus can be considered as suitable methods for non- invasive skin toxin extraction in future studies. We expect to get the results of the qualitative and quantitative analysis of the extracted toxin later this year.

278 279 ECOLOGY PHYLOGENY

Population size, survival and viability of the fire salamander Phylogenetic signal in vocalizations of Neotropical harlequin frogs (Salamandra salamandra) in Central Croatia (Atelopus)?

Nina Popovic´ Julia Rautenberg, Stefan Lötters

Karlovac University of Applied Sciences, Trg J.J. Strossmayera 9, 47000 Karlovac, Croatia; [email protected] Department of Biogeography, Universitätsring 15, 54296 Trier, Germany; [email protected]

Population viability analysis (PVA) has become a commonly used tool to estimate extinction Acoustic signals are important in intraspecific communication of most anuran species. To risk and to model population dynamics of stage-structured populations, such as amphibians. optimize the cost-efficiency balance, natural and sexual selection often blur phylogenetic signals. There have been no studies of long-term population dynamics in Croatia. In this study I We study advertisement calls and their evolution in harlequin frogs (Atelopus). Vocalizations combined capture-mark-recapture data collected over a 3 year period with population viability in this bufonid group are suggested to be ‘conservative’, perhaps because Atelopus species are analysis. The aim of this study was to analyse data on population size and adult survival of largely allopatric and probably are only weakly affected by competition with non-congenerics, the fire salamander (Salamandra salamandra) and use it to confirm the predictions of popu- given that in general there are only few diurnal riparian anurans. Moreover, streams may be lation model. Population sizes, annual survival and viability of the fire salamander were esti- noisy and many taxa lack a middle ear. We here test the hypothesis that in these anurans mated in an area of 4.25 ha in the temperate forest in central Croatia, where 4 permanent acoustic traits reflect phylogenetic signal. By using simple Mantel tests, we compare Euclidean plots were installed. Drift fences and pitfall traps were used as sampling technique. Jolly- distances derived from quantitative call characters of so called ‘pulsed calls’ (i. e. advertisement Seber mark-recapture model was used for the estimation of population density and survival. calls) from 12 harlequin frog populations with genetic P-distances derived from mtDNA Data describing survival rates for distinct salamander population, complemented by a litera- markers and relate traits to a phylogeny. Although sampling is limited, our results show sig- ture review on additional demographic parameters, were used to define population matrix nificant correlations in two out of four traits studied. This leads us to tentatively confirm our and carry out simple viability analysis using a stage-structured model in the RAMAS GIS hypothesis. platform. Estimated population density ranged between 12.5 and 90 ind. ha–1. The mark- recapture analysis of adult salamanders showed low rates of annual survival of 0.393. These trends are consistent with simulations of the system, which predicted that studied population had high extinction risk. Estimated quasi-extinction risk within a 50 year period ranged from 88.2% to 100%. Simulation predicted median time to extinction of 17 years. The PVA model simulation estimated also a negative trend in abundance for the S. salamandra population. These data offers some insight into the likely viability of the study population and suggests a link between local adult survival and salamander population trend. Predictions of PVA can be used as a tool to better understand the demography of amphibian populations.

280 281 ECOLOGY ECOLOGY

Adaptive variation in development time in response to habitat type in A spatial approach for testing the cryptic role of dorsal pattern design Rana temporaria: a joint assessment of inter- and intrapopulation in Vipera seoanei variation and variation within clutches André Rocha 1, Luisa Viana 1, Xavier Santos 2, Fernando Martínez-Freiría 2 Jane Reniers, Luc Brendonck, Bram Vanschoenwinkel 1 Faculdade de Ciências da Universidade do Porto, Rua Campo Alegre, 4169-007 Porto, Portugal 2 CIBIO/InBIO, Centro de Investigação em Biodiversidade e Recursos Genéticos da Universidade do Porto, Laboratory of Aquatic Ecology, Evolution and Conservation, KU Leuven, Charles Deberiotstraat 32, 3000 Leuven, Instituto de Ciências Agrárias de Vairão, R. Padre Armando Quintas, 4485-661 Vairão Portugal; Belgium; [email protected] [email protected]

In anurans the timing of metamorphosis is a crucial life history trait with important fitness Crypsis and aposematism are oppossing evolutionary strategies derived from predator-prey consequences. For instance, postponing metamorphosis can result in fitter adults, but also interactions which have been ascribed to the zigzag pattern of European vipers. Previous increases the risk of larval mortality when the habitat dries out. To assess the relative impor- experiments using plasticine models have demonstrated its aposematic role, while no evidence tance of local adaptation, phenotypic plasticity and bet hedging contributing to the repro- has been found regarding a possible cryptic function. Moreover, the high frequency of mela- ductive success of pond breeding frogs, we set up a large scale laboratory common garden nistic specimens at high altitude or latitude was suggested to play an important role in speci- breeding experiment for 9 populations of the widespread generalist breeder Rana temporaria mens’ thermoregulation. Vipera seoanei is an endemic species to Iberia restricted to the northern (the common frog), from habitats that differ in permanence and habitat predictability. More region and with Euro-Siberian affinity. It is highly polymorphic showing four types of dorsal specifically, freshly laid eggs were collected from shallow wheel tracks with short hydroperiods, colouration patterns (bilineata, cantabrica, classical and uniform) and high frequency of from permanent fishless habitats and from semi-permanent ponds that typically dry up once melanistic specimens in mountains. In this work we aim to give insights into the cryptic role a year but may experience variable hydroperiods and both development time and develop- of V. seoanei dorsal pattern phenotypes by checking its relationship to environmental and mental plasticity in response to a water drop were quantified. Results revealed adaptive vari- habitat factors. A total of 490 georeferenced specimens from all of its Iberian range were double- ation in development time with frogs metamorphosing earlier in more short lived habitats. blind classified according to a dichotomic key dealing with diagnostic characters of dorsal Simulated pool drying reduced development time in all populations but there were no signi- colouration. Then, the geographical coherence of phenotypes distribution was tested using ficant differences among populations or habitat types. Intra-population variation in development Geographic Information Systems, and Ecological niche models using 19 low correlated vari- time was higher in populations from semi-permanent breeding ponds and resulted mainly ables (lithological composition, solar radiation, climate and habitat variables). Five clear pheno- from variation among, rather than from variation within clutches. Overall, these results suggest types were identified for 448 specimens. Intermediate phenotypes for classic with cantabrica that R. temporaria uses a combination of strategies to buffer against reproductive failure, and bilineata were also identified (n=42). Each phenotype had a strong geographic coherence including variation in life history traits within populations, local adaptation and a base-level with intermediate phenotypes mostly occurring in contact zones. Environmental (e. g. tem- of developmental plasticity. Bet-hedging via life history variation within clutches seems to be perature, precipitation) and habitat (e. g. abundance of pastures and grasslands) variables were relatively unimportant in this system. related to one or more phenotypes but showing distinct response curve profiles. It is discussed the role of these factors as drivers of crypsis and thermorregulative performance and also the relation of phenotypes distribution to the genetic structure of the species.

282 283 SYSTEMATICS ECOLOGY

Systematics, niche evolution and biogeography of the Middle Eastern Population structure dynamics of a common frog population at the newt (Neurergus) northern limit of its range

Dennis Rödder 1, Faraham Ahmazadeh 1, Timo Hartmann 1, Morris Flecks 1, Daniela Rosioru 1,4, Rodica Plaiasu 2, Raluca Ba˘ncila˘ 2, Claude Miaud 3, Flora Ihlow 1, Thomas Schöttler 2, Jürgen Fleck 3, Jonas Astrin 1, Wolfgang Böhme 1 Dan Coga˘lniceanu 4

1 Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; 1 National Institute for Marine Research and Development “Grigore Antipa”, Blvd. Mamaia nr. 300, [email protected] 900581 Constant¸a, Romania; [email protected] 2 Breslauer Straße 12, 65307 Bad Schwalbach, Germany 2 “Emil Racovit¸a˘” Institute of Speleology, 13 Septembrie Road nr. 13, 050711 Bucharest, Romania 3 Pfarrer-Hufnagel Straße 23, 63454 Hanau, Germany 3 UMR 5175, CEFE, Ecole Pratique des Hautes Etudes, Laboratoire Biogéographie et Ecologie des Vertébrés, 1919 route de Mende, 34 Montpellier, France It is well known, that evolutionary specialization of taxa to a particular climatic regime lim- 4 University Ovidius Constant¸a, Faculty of Natural Sciences, Aleea Universita˘¸iit 1, corp B, 900470 Constant¸a, Romania its their dispersal. Especially slowly and conservatively evolving organisms like ectotherm newts are likely to be strongly affected by environmental variables. Herein, we studied the Populations from high latitude, at the limit of their geographical range experience extreme correlation of environmental niche overlap and molecular similarity to reveal insights of the climate. These populations are the most vulnerable to climate changes and require special systematic relationship within the Middle Eastern newt, genus Neurergus. A molecular phy- conservation measures. We studied the temporal variation of age and body size in an isolated logenetic analysis based on two mitochondrial genes including all known taxa and recently population of the common frog (Rana temporaria) from Kilpisjarvi, Lapland, Finland. Body discovered geographically distinct colour morphs of the salamandrid newt genus Neurergus size and age (estimated by skeletochronology) were measured for 120 adult frogs and 51 sub- was computed to reveal the valid species of Neurergus. Species distribution models of the adults captured in 2003, 2009 and 2010. genetically validated species of Neurergus were calculated to gain well-defined potential Age ranged between 4–9 years for females, 3–10 years for males and 2–7 years for subadults. distributions for each species and provide a detailed zoogeographic picture. Based on this, Males were on average younger than females, but the median age for males was significantly niche occupancy profiles were created to evaluated niche conservatism between species of the higher in 2010 than in 2009. We found no significant difference between sexes in age at sexual genus Neurergus and show the correlation of environmental niche overlap and molecular sim- maturity, males and females reaching the sexual maturity at 3 to 5 years. Body size varied highly ilarity. Eight morphologically distinguishable and geographically separated taxa respective- within age classes and although a consistent positive relationship between body size and age ly colour morphs of Neurergus crocatus were included in the molecular analyses. Discovering was found, the relationship was not significant for either sexes or subaduls. discordance between mtDNA and morphology, our molecular analyses identified only four The results of our study indicated that males have a slight tendency to live longer. Body size well-defined evolutionary lineages which represent valid species of Neurergus. Niche overlaps was a poor predictor of individual age possibly because of different resource allocation during among these four species are low, suggesting only a minor degree of niche conservatism in growth periods. Neurergus. Still, a tendency towards slightly higher niche conservatism was discovered compar- ing the more closer related taxa.

284 285 PHYLOGEOGRAPHY PHYLOGENY

Frog biogeography and evolution in the Congo Basin: Phylogenetic relationships and phylogeography of the genus Lacerta some preliminary insight in Greece

Daniela Rössler 1, Jos Kielgast 1,2, Rayna Bell 3, Eli Greenbaum 4, Kostas Sagonas 1, Nikos Poulakakis 2,3, Petros Lymberakis 3, Aristeidis Parmakelis 4, Maximilian Dehling 5, Václav Gvozˇdík 6,7, Zoltán T. Nagy 8, Werner Conradie 9, Panayiotis Pafilis 5, Efstratios D. Valakos 1 Patrick J. McLaughlin 10, Susanne Müller 1, Marius Burger 11, Stefan Lötters 1 1 Department of Human and Animal Physiology, Faculty of Biology, University of Athens, Panepistimioupoli 1 Department of Biogeography, Universitätsring 15, 54296 Trier, Germany; [email protected] Zografou, 15784 Athens, Greece; [email protected] 2 2 Natural History Museum of Denmark, Universitetsparken 15, 2100 Copenhagen, Denmark Department of Biology, University of Crete, Vassilika Vouton, P.O.Box 2208, 71409 Irakleio, Crete, Greece 3 3 Department of Ecology & Evolutionary Biology, Cornell University, Ithaca, NY 14853 USA Natural History Museum of Crete, University of Crete, Knosos Ave, P.O.Box 2208, 71409 Irakleio, Crete, Greece 4 4 Department of Biological Sciences, University of Texas at El Paso, 500 West University Avenue, El Paso, TX 79968, Department of Ecology and Taxonomy, Faculty of Biology, University of Athens, Panepistimioupoli Zografou, USA 15784 Athens, Greece 5 5 Abteilung Biologie, Universität Koblenz, Universitätsstraße 1, 56070 Koblenz, Germany Department of Zoology and Marine Biology, Faculty of Biology, University of Athens, Panepistimioupoli Zografou, 6 National Museum, Cirkusova 1740, 19300 Prague, Czech Republic 15784 Athens, Greece 7 Biogeography, University of Basel, Klingelbergstraße 27, 4056 Basel, Switzerland 8 Royal Belgian Institute of Natural Sciences, Rue Vautier 29, 1000 Brussels, Belgium The green lizards of the genus Lacerta (Sauria, Lacertidae) consist of 8 species which are 9 Port Elizabeth Museum, P.O. Box 13147, 6013 Humewood, South Africa almost restricted to the southern European peninsulas. The taxonomic assignment in species 10 Department of Biology, Drexel University, 3245 Chestnut Street, Philadelphia, PA 19104, USA and subspecies level is problematic and their taxonomy is complex and constantly revised. 11Zoology Department, University of the Western Cape, Private Bag X17, 7535 Bellville, South Africa Three species are present in Greece: L. trilineata in continental and insular Greece, L. viridis The biogeography and evolution of biota in the Congo Basin continues to be among the in northern and central continental Greece and some Aegean islands, and L. agilis in northern most enigmatic among the worldwide tropical rain forests. Most likely this vast lowland has Greece. Here, sequence data derived from two mitochondrial (cyt-b and 16S rRNA) and one been substantially impacted by past climatic oscillations leading to dramatic dynamics in nuclear (NKTR) genes, were used to infer the phylogenetic relationships and evaluate the ecosystems over time. We study these events using vertebrates which are generally sensitive to taxonomic status of the genus Lacerta in Greece. All phylogenetic analyses (Maximum Parsi- temperature and humidity: anuran amphibians. In the course of an ongoing collaborative pro- mony, Maximum Likelihood and Bayesian Inference), produced a tree with three major ject, we aim at linking information from phylogenetic and distribution studies using wide- clades; the first clade corresponds to L. trilineata, the second to L. agilis, and the third to a spread species (or species complexes) in the genera Hylarana, Hymenochirus and Hyperolius, complex of L. viridis and L. bilineata, raising questions about the current taxonomy and the showing distinct differences in key elements of their life history (i. e. habitat requirements, distribution of the two latter species. Moreover, L. trilineata appears as a monophyletic group body size, reproductive biology, dispersal capacity, etc.). At the current state, we have been composed of six allopatric lineages. However, the eight L. trilineata subspecies, previously able to investigate mtDNA samples from all over the basin and its peripheries and have devel- defined on the basis of morphology, do not correspond to the different lineages inferred. oped climate-based statistical models on species’ potential distributions. The results enable us Thus, their status should be reconsidered. In conclusion, our data of Greek Lacerta stress the to catch a preliminary glimpse of possible patterns of dispersal and vicariance events. Our need for a reconsideration of the taxonomy of this main reptile group of Greece and high- findings tentatively support a general dispersal tendency from West to East but also the light the difficulties that classical taxonomy is faced with when attempting to infer the re- potential existence of a refuge within the center of the basin and can be used to formulate lationships at species and subspecies levels. further hypotheses.

286 287 MORPHOLOGY, ANATOMY AND DEVELOPMENT MORPHOLOGY, ANATOMY AND DEVELOPMENT

Ovarian cycle of Bothrops atrox, a Neotropical viperid Complex origin of some bones in the grass snake Natrix natrix skull embryogenesis Sávio S. Sant’Anna 1,2, Daniela R. Vasconcellos 1, Ana Paula Croce 1, Cíntia Y. Fugiwara 1, Josiane L. F. Silva 1, Renato Moterani 1, Mayara S. Hidaka 1, Hanna V. Sheverdyukova Bruno M. Costa 1, Rafael V. Fernandes 1, Wilson Fernandes 1, Kathleen F. Grego 1 Schmalhausen Institute of Zoology, NAS of Ukraine, Khmelnitsky Street 15, Kyiv, PA 01601, Ukraine; 1 Laboratory of Herpetology, Butantan Institute, 1500 Vital Brasil Avenue, 05503-900 São Paulo, Brazil; [email protected] [email protected] 2 Department of Pathology, School of Veterinary Medicine and Animal Sciences, University of São Paulo, In order to study the peculiar properties of Natrix natrix skull formation and the way of its 87 Prof. Dr. Orlando Marques de Paiva Avenue, 05508-270, São Paulo, Brazil primordia appearance in embryogenesis 35 embryos of successive stages of development were The reproduction of venomous snakes in captivity has become increasingly necessary. Several investigated. The stages of development were determined in accordance with the table of the species are difficult to obtain or are even threatened with extinction, making studies of their stages of normal development by D.R. Zehr. Pregnant females were caught in their natural poisons very difficult. Knowing the cycle and determining the best reproductive management habitat. Egg clutches were incubated in wetted vermiculite. To obtain embryos at different of these species in captivity increases breeding success. Bothrops atrox is a species found in stages of development several embryos were taken every day. After oviposition the females were northern South America east of the Andes, where it is responsible for most snake bite acci- released in the places of their catch. Serial histological sections and double stained embryos dents that occur in these locations. The aim of this study was to monitor the ovarian cycle of were made. females of B. atrox to determine the best reproductive husbandry of this species. During a As a result it was found that the maxillary and prefrontal bones have two, and post- period of 17 months (March 2010 to July 2011) six females of B. atrox were monitored monthly orbital, parietal, supraoccipital, septomaxillary bones and basisphenoid three centers of ossi- by ultrasound equipment, using a linear transducer of 6 to 13 MHz, to observe the develop- fication. ment of their ovarian follicles. The follicles were divided into four stages according to their According to the classical theory separate centers of ossification in bones are the vestiges of size: stage I (follicles <0.6 cm), stage II (0.6 to 1.1 cm), stage III (1.2 to 2.0 cm) and stage IV independent ancestral bones. The formation of some bones in N. natrix skull from several ( 2.1 cm). The follicles in stages I and II were present in all months of the year, but in a larger centers of ossification confirms their complex origin. We assume that the maxillary bone in ≥ percentage in the months of December to March (summer), follicles in stage III were seen N. natrix is formed as a result of the confluence of maxillary, zygomatic and quadrate–zygo- mainly from April to December (autumn, winter and spring) and stage IV follicles were seen matic bones; septomaxillary—of septomaxillary and supraethmoid bones; prefrontal bone— in the months of June to October (winter and spring). From May to August 2010, groups of of supraorbital, prefrontal and lacrimal bones; postorbital bone—of postorbital and post- two males and one female were released into a heated room with a 24-h monitoring system. frontal bones, supra-occipital—of two supraotic bones, which were independent in ancestors. In June 2010, two females were mated and in September of the same year fertilized eggs could Probably in snakes phylogeny the decreasing number of skull bones was not due to their already be seen in the oviducts. In December 2010, the offspring of both females were born, reduction, but as a result of their merging into monolithic ones. a clutch with 6 and another with 9 healthy babies. It was observed that the month of June had a higher success of mating rate, coinciding with the greatest amount of ovarian follicles in stage III encountered in the females that copulated. Supported by: FAPESP, CNPq and INCTTOX.

288 289 ECOLOGY PRACTICAL CONSERVATION

Effects of thermal spring on elemental concentration of frog toe bones Societas Herpetologica Slovenica’s activities for helping amphibians at road crossings Edina Simon 1, Edvárd Mizsei 2, János Tamás Kundrát 1, István Gyulai 3, Mihály Braun 4, Béla Tóthmérész 1 Maja Sopotnik 1, Tomazˇ Jagar 1, Nino Kirbisˇ 1, Aleksandra Lesˇnik 1,2, Erika Ostanek 1, Iris Petrovicˇ 1, Katja Poboljsˇaj 1,2, Alenka Rozman 1, Lovro Rozman 1, 1 Department of Ecology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary David Stankovic´ 1, Melita Vamberger 1,3, Daniela Vlacˇic´ 1, Anamarija Zagar 1 2 Department of Evolutionary Zoology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary; [email protected] 1 3 Department of Hydrobiology, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Hungary Societas Herpetologica Slovenica, Vecˇna pot 111, 1000 Ljubljana, Slovenia; [email protected] 2 4 Department of Inorganic and Analytical Chemistry, University of Debrecen, Egyetem tér 1, 4032 Debrecen, Centre for Cartography of Fauna and Flora, Antolicˇicˇeva 1, 2204 Miklavzˇ na Dravskem polju, Slovenia Hungary 3 Museum of Zoology (Museum für Tierkunde), Senckenberg Natural History Collections Dresden, Königsbrücker Landstraße 159, 01109 Dresden, Germany The aim of our study was to analyze the effects of thermal spring on elemental concentra- tions on frog toe bones in the Balkan frog (Pelophylax kurtmuelleri). There were collected 60 Road mortality during spring migrations is a major threat to the survival of amphibian pop- adult frogs from five thermal spring and one control pond (10 specimens from each pond) in ulations. In Slovenia, there are more than 1500 amphibian road mortality hotspots. Societas southern Albania in 2011. The control pond was about 20 km from the thermal springs. Herpetologica Slovenica has put much effort into addressing and raising public awareness on Weights and body length of each frog were measured. The following elements were analysed this issue. As a part of this, several projects of helping amphibians at road crossings are carried in toe bones Ba, Ca, Mg, P, S, Sr and Zn, respectively. Our results showed that significantly out annually. Since 2008, mitigating measures are implemented and temporary amphibian smaller frogs were found in the thermal spring than in the control pond, expect one thermal fences of varying length (no fence in 2009) are set up during spring migrations in Ljubljana, spring. Principal component analysis (PCA) showed a weak separation of thermal springs and and from 2010 on also in Maribor. Both locations are set in an urban area. This enables the control pond based on the elemental concentration of frog toe bones. Higher Ca and P con- society to attract a large number of volunteers to help carry the amphibians across the road. centrations were found in frogs from the control pond than in frogs which were collected from The data on the number, species composition and the exact location of amphibians crossing thermal springs. In the cases of other elements such as Fe, S and Ba the highest concentration the roads is gathered yearly. The number of amphibians that were carried over the road in was in frogs from thermal spring. Our results indicated that the bioaccumulation potential Maribor has been increasing each year, together with the increasing number of volunteers was high for frogs from the sediment, because of strong correlation was found between ele- helping with the project. In Ljubljana, the number of volunteers is also increasing, but the mental concentration of sediment and toe bones (R2 = 0.792). While, weakly interaction was number of amphibians varies from year to year. Altogether, more than 11,000 individual between elemental concentration of thermal spring and river water and toe bones (thermal amphibians were carried over the road at the above mentioned locations, most of which spring: R2 = 0.393, river: R2 = 0.180). In summary, the analysis of toe bones demonstrated that belonged to the common toad (Bufo bufo) species. In Maribor, more young B. bufo specimens thermal springs may have effects of frog morphology size and elemental concentration of toe are observed each year—a potential positive effect of the project. bones, and bones are useful to assess the environmental load and the quality of aquatic envi- As an addition to the above mentioned activities, a support page with a “frogphone” tele- ronment. phone number was established in 2011, giving information about amphibians and roads to other amphibian conservation initiatives and to the general public. The attention gained from the public and the media shows that these kinds of activities are a good conservation measure in terms of raising public awareness. The gathered data can serve as the basic information needed for long-term conservation measures, such as permanent fences and tunnels.

290 291 DISTRIBUTION BEHAVIOUR

New distribution data and conservation status of Acoustic communication in two syntopic spadefoot toads Vipera ursinii rakosiensis (Méhely, 1893) in Transylvania, Romania Florina Sta˘nescu 1, Paul Székely 1, Rafael Márquez 2, Dan Coga˘lniceanu 1 Tibor Sos 1, Ioan Ghira 2, Zsolt Hegyeli 1 1 Faculty of Natural and Agricultural Sciences, Ovidius University, Aleea Universita˘¸iit 1, 900470 Constant¸a, 1 “Milvus Group” Bird and Nature Protection Association, 540620 Tîrgu-Mures¸, Romania; [email protected] Romania; [email protected] 2 Babes¸-Bolyai University, Faculty of Biology and Geology, 400084 Cluj-Napoca, Romania 2 Fonoteca Zoológica, National Museum of Natural Sciences (CSIC), José Gutiérrez Abascal 2, 28006 Madrid, Spain

The rare and endangered Vipera ursinii rakosiensis is a small-sized (up to 60 cm long) lowland Although most of the studies in anuran acoustic communication focus on advertisement steppe subspecies of the V. ursinii group. Its distribution is strictly associated with the natural calls, release vocalizations are also an important component of the mate-recognition system, or semi-natural steppe grassland remnants, and it displays a similar insular distribution pattern. by allowing energy conservation in cases of erroneous amplexus during reproduction (e.g. Until now, the only extant population of the subspecies had been recorded in the district of male-male, male-unreceptive female). Release calls are emitted both during and outside the Lopadea Noua˘ locality, Alba County. Previously, V. u. rakosiensis has been recorded from six reproductive season and provide their receivers information about the species, sex and size of localities: Fânat¸ele Clujului, Fa˘get forest and Flores¸ti (all in the vicinity of Cluj-Napoca), the emitter. The genus Pelobates consists of four fossorial and nocturnal species. Two of these between Bont¸ida and Sic, Valea Florilor and Bogata, all in Cluj County. Here we report on species, P. syriacus and P. fuscus, occur in south-eastern Europe, and their ranges overlap in a new locality for the taxon in the administrative territory of Ra˘des¸ti (Alba County) and also the Balkan Peninsula. We quantitatively described the typical release calls (i. e. first order) of reconfirm the presence of the meadow viper outside the borders of the Fânat¸ele Clujului both species and tested for intra- and interspecific differences of the calling pattern. The Nature Reserves and in the hayfields neighbouring Bogata. The newly discovered sites occu- sound samples were obtained under laboratory conditions by amplexus simulation, from 25 pied by meadow vipers at Fâna¸elet Clujului, Ra˘des¸ti and Lopadea Noua˘ have been totally or adults: 12 P. fuscus (8 males, 4 females) and 13 P. syriacus (7 males, 6 females). The animals partially included within the existing Natura 2000 network. One of the most important results were collected from two syntopic populations in Dobrudja region (Vadu, Romania). We used of the 11–12 October 2012 biogeographical seminar in Bucharest was the acceptance of the Raven Pro 1.4 software for sound analysis. We measured a series of temporal and spectral proposal for the Bogata population to be included within a new Natura 2000 site. parameters from 497 calls: call duration, frequency range, dominant frequency and other Although covered by the Natura 2000 network, all known Transylvanian viper populations parameters useful in pattern differentiation. In both species, the release vocalizations consisted are still significantly threatened by habitat loss caused by human agricultural activities. Habitat of series of pulses and harmonic complex tones and the general structure of the call was destruction and fragmentation by ploughing and breaking the natural and semi-natural grass- approximately the same. We found no significant differences in the frequency range and lands for the extension of cultivated fields remains a significant threat. Most existing secondary dominant frequency of the calls, which indicates that the two species share the same acoustic grasslands are still recorded in the official cadastral documents as ploughlands, allowing owners niche. We found significant differences in the temporal and spectral patterns of the calls, to plough them, even when the land has not been ploughed for decades. Overgrazing and between both sexes and species. We suggest that release vocalizations are important in medi- deliberate or accidental burning destroy the vegetation cover. Illegal collecting and harassment ating intra- and interspecific interactions and their communicational potential requires more of snakes, which often end with their death, and associated effects of urban and tourism attention. developments, also play an important role in the population decline. In order to preserve the viper populations in Transylvania, the only suitable solutions would be: (1) to persuade private owners to manage lands traditionally—but with restrictions —through the use of subsidies for Natura 2000 sites, and to encourage them to benefit similar payments under the agro-environmental schemes outside the Natura 2000 network; (2) to create an administrative structure financially supported by the state, in order to ensure proper management of the three large Nature 2000 sites: “Pajis¸tile lui Suciu” (ROSCI0187), “Dealu- rile Clujului de Est” (ROSCI0295) and for the newly proposed “Dealurile Bogatei” site, or (3) for the state or NGOs to lease or purchase these lands in order to implement an appro- priate management regime. Further intensive surveys are needed across the whole region to identify remnant meadow viper populations for immediate protection.

292 293 MONITORING MONITORING

Distribution and status of Rana arvalis in Central Slovenia Amphibians and reptiles in drinking water manholes: a supplementary method for herpetofaunal data collection David Stankovic´ 1, Maja Cipot 2 Márton Szabolcs 1, Szabolcs Lengyel 2, Tamás Zsólyomi 1 1 University of Ljubljana, Biotechnical Faculty, Department of Animal Science, Groblje 3, 1230 Domzˇale, Slovenia; [email protected] 1 Nature Conservation Association of Tokaj, Liget köz 1, 3910 Tokaj, Hungary; [email protected] 2 Centre for Cartography of Fauna and Flora, Antolcˇicˇeva 1, Miklavzˇ na Dravskem polju, 2204, Slovenia 2 Department of Tisza River Research, Hungarian Academy of Sciences, Centre for Ecological Research, Bem tér 18/c, 4026 Debrecen, Hungary Rana arvalis is one of the 19 amphibian species that are found in Slovenia. Until recently it has only been recorded in north-eastern and south-eastern lowlands of Slovenia, parts of the Drinking water manholes with concrete plate cover are ecological traps and act like pitfall Pannonian and Sava Basin. Here we report on the first indisputable record of this species traps for amphibians and reptiles. They get inside by falling down when seek for shelter and from the Central Slovenia, give preliminary results on the size and the distribution of the sub- stuck in. Hence manholes serve as data gathering sources for us. On the other hand they are populations and try to estimate present threats and status of the population and habitat. The a possibly global threat for animals that will die if not rescued. We visited 29 manholes along data presented here was collected between 2009 and 2013. roads in a landscape of varied topography in north-eastern Hungary and counted and res- On the 25th of March 2010 the presence of the moor frog was confirmed for the first time cued fallen amphibians and reptiles. We checked manholes at least once a month for six years in Ljubljansko barje. The closest known population is located about 100 km SE in Krakovski (northern site, river floodplain and hills with forests and vineyards) or for four years (southern gozd near Novo mesto. In the breading season of 2011 an extensive field survey was carried site, mainly croplands, grasslands and wetlands). We found a total of 1726 individuals of ten out. Twelve potential areas where R. arvalis could be breeding were examined and on five amphibian and six reptile species (1455 and 271 individuals, respectively), roughly half (47%) areas breeding was confirmed. Altogether 6,112 spawn clumps of R. arvalis were counted, of the herpetofauna of Hungary. We found every species reported in the literature, or were while additional 3,669 clumps could either belong to R. temporaria or R. arvalis. The highest expected from each area, except one. Bombina bombina was by far the most frequently ecological density was 84.67 adult frogs/ha. encountered amphibian (59%), followed by species of the Pelophylax esculentus complex Major threat for R. arvalis in Central Slovenia is fragmentation and habitat destruction due (14%), Pelobates fuscus (9%), and Triturus dobrogicus (8%). Natrix natrix was the most frequent to urbanization. This area is in the immediate proximity of the capital city and the municipal reptile (63%), followed by N. tessellata (20%) and two Lacerta species (L. agilis, L. viridis, 5% landfill site which is planned for expansion. Because of its close vicinity to the capital this each). Most amphibians (74%) were found at the southern site, whereas most of the reptiles population is under the strong influence of human recreational activities. Here, visitors are (77%) were found at the northern site. Daily capture rates (number of animals trapped divided frequent especially during the spawning time, when they often approach the animals up close by the number of days passed since the last check) of amphibians varied by season (autumn > and scare them. spring > summer) and there was a significant interaction between season and year, whereas at reptiles only depended on site (northern > southern). These results draw attention to the potentially harmful impact of manholes, but also show that counting and rescuing trapped animals in appropriately short time intervals can be a useful tool for monitoring amphibians and reptiles.

294 295 BEHAVIOUR ECOLOGY

Differences in locomotor activity in two syntopic spade- Variation in body size and sexual size dimorphism in two foot toad species (genus Pelobates) Pelobates fuscus populations

1 2 1 1 Diana Székely 1, Mathieu Denoël 2, Paul Székely 1, Dan Coga˘lniceanu 1 Paul Székely , Raluca Ba˘ncila˘ , Diana Székely , Dan Coga˘lniceanu

1 1 Faculty of Natural and Agricultural Sciences, Ovidius University, Aleea Universita˘¸iit 1, 900470 Constant¸a, Faculty of Natural and Agricultural Sciences, Ovidius University, Aleea Universita˘¸iit 1, 900470 Constant¸a, Romania; [email protected] Romania; [email protected] 2 2 Laboratory of Fish and Amphibian Ethology, Behavioural Biology Unit, University of Liège, 22 Quai van Beneden, Institute of Speleology “Emil Racovit¸a˘”, 13 Septembrie Road, No. 13, 050711 Bucharest, Romania 4020 Liège, Belgium The analysis of intra- and interspecific patterns of geographic variation in body size have Understanding the coexistence of related species in ecological communities is a challenge for played and still maintain an important role in evolutionary biology. In amphibians body size ecologists. The spadefoot toads (genus Pelobates) from Dobrudja (Romania) provide a very is often closely related to fitness due to its association with longevity, fecundity, metabolic interesting study-case: the ranges of two species of the genus overlap at the limit of their dis- rate and tolerance to environmental stresses such as starvation and desiccation and is mostly tribution ranges—southern limit for P. fuscus and northern limit for P. syriacus. Both spade- consistent with variation of some characteristics of the environment. Here we investigate the foot toad species are nocturnal, fossorial, and exploit a similar ecological niche. They have variation in body size and sexual dimorphism of common spadefoot toads (Pelobates fuscus) similar sizes at metamorphosis, reach sexual maturity at a similar age but P. syriacus becomes collected from two bioregions in Romania that lie 470 km apart: the Continental (Sa˘licea, significantly larger than P. fuscus. We examined the adaptive value of interspecific variation Transylvania) and the Pontic (Vadu, Dobrudja) bioregion, respectively. Eight morphometric in behavioural patterns, such as level of activity (locomotor behaviour), since any type of measurements, body length (L), head width (Ltc), femur length (F), tibia length (T), tarsus movement includes trade-offs, in terms of energetic costs, exposure to predation and drought, length (Ltars), first toe length (Dp), forearm length (Lma) and mass (G), were recorded for but at the same time provides increased access to food. The nocturnal activity of 6 adult P. 324 (137 females, 187 males) individuals from Sa˘licea and 74 (32 females, 42 males) individu- fuscus and 6 P. syriacus (3 females and 3 males for each) was monitored in a laboratory setting als from Vadu. Significant overall differences between mean trait size for females and males for a period of 30 days. We used webcams to capture time-lapse photographs of the toads’ were found, with a considerable reduction in size from the Sa˘licea, to the Vadu populations. movement, which were then used to obtain tracks for each individual (a total of 48 tracks and Results showed significant differences between populations for F, T, Dp and G (females) and over 260 hours of activity). Our results showed that P. syriacus was significantly more active Ltc, F, T, Dp, and G (males). We found sexual dimorphism in mean Ltc, F, T, Dp and Lma than P. fuscus with almost 2.5 overall distance covered and more than twice the speed size, females being significantly larger than males in both populations. The main implication achieved, but with approximately the same time spent outside burrows. Our preliminary of this study is that the populations differ in the magnitude of geographic variation of the results suggest that the closely related Pelobates species from Dobrudja differ in patterns of body size. The differences are most likely due to the different environmental conditions: foraging behaviour, P. fuscus being relatively sedentary (‘sit-and-wait’) whereas P. syriacus is i) drier and harsher environment, ii) availability of food resources and iii) possible competition more an active (‘widely-foraging’) predator. with the larger Eastern Spadefoot Toad, Pelobates syriacus, in the Vadu population.

296 297 POPULATION GENETICS PRACTICAL CONSERVATION

Genetic connectivity of (meta-) populations in The Active Protection Program of the common toad (Bufo bufo L., 1758) Hyalinobatrachium valerioi with the population’s condition analysis on the migratory routes in Wroclaw, Poland Katharina Trenkwalder, Alexandra Mangold, Walter Hödl, Max Ringler, Eva Ringler Edyta Turniak, Aleksandra Kolanek

Department of Integrative Zoology, University of Vienna, Althanstraße 14, 1090 Wien, Austria; Laboratory of Vertebrate Evolutionary Biology & Conservation, University of Wroclaw, ul. Sienkiewicza 21, [email protected] 50-335 Wroclaw, Poland; [email protected]

The reticulated glass frog Hyalinobatrachium valerioi (Centrolenidae) is a small leaf-breeding Bufo bufo is the most common species of toad in Poland and one of the first active amphib- frog in tropical rainforests of Central and northern South America. The species typically ians in early spring. They usually cross many routes during their migration to the mating sites. occurs in small aggregations of breeding individuals along forest streams. While some studies The Active Protection Program in one of the biggest common toad populations in Wroclaw have investigated the reproductive behavior of this frog species, little information is known has started in 2010. The main goal of our activities is to reduce amphibian mortality caused about individual migration rates and gene flow between such patches. We monitored a pop- by traffic. Migrating individuals are collected from the roadway and transported safely to the ulation of H. valerioi at the tropical lowland stream Quebrada Negra near La Gamba, Costa vicinity of breeding ponds. In 2010, we found 1129 individuals, including 521 males, 301 Rica, during their reproductive season between August and December 2012. Along a 400 m females and 307 dead individuals (sex undefined). Mortality was about 27%. A year later, river transect we identified 10 spatially clustered breeding aggregations, with on average 16 condition of population was similar with total of 1279 individuals collected (701 males, 378 individuals per patch, separated 5 up to 50 meters. We recorded precise spatial locations of all females), mortality decreased to 15.6%. In 2012, a strong reduction of the migrating individuals, frogs and clutches, took dorsal pictures of all frogs for individual identification, and collected was observed, most likely caused by adverse and unusual weather conditions. Even if there was tissue samples from the adults and clutches for subsequent molecular analyses. The integration snow, migration of amphibians started, but temperature dropped below 0 °C again. Migration, of observational field data and molecular analyses of highly polymorphic microsatellite loci normally lasting about two weeks, extended to four weeks. Only 306 toads were found (241 provides valuable information about gene flow between the patches and also the genetic related- alive and 65 dead individuals), so the number of population decreased four times. In 2012 ness among the individuals within single patches in the H. valerioi population under study. action was extended by morphometric studies of carries amphibians. In 2013, data are still collected. This year migration was hampered by a long winter and began about three weeks later than usual.

298 299 ECOLOGY ECOLOGY

Spatial niche segregation among three lacertid taxa living in close The effect of the pesticide Glyphogan Classic® on two common proximity in Bulgaria aquatic predators

Nikolay D. Tzankov 1, Georgi S. Popgeorgiev 2, Atanas P. Grozdanov 3 János Ujszegi 1, Zoltán Gál 2, Zsanett Mikó 3, Attila Hettyey 4

1 National Museum of Natural History, BAS, Tsar Osvoboditel Blvd. 1, 1000 Sofia, Bulgaria “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, 2 Regional Natural History Museum, Hristo G. Danov St. 34, 4000 Plovdiv, Bulgaria Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] 3 Sofia University “St. Kliment Ohridski”, Dragan Tsankov Blvd. 8, 1164 Sofia, Bulgaria; [email protected] Pesticides containing glyphosate and POEA (e. g., Glyphogan®) can have a negative impact The three lacertids subject of this study (Lacerta agilis bosnica, L. a. chersonensis and L. viridis on non-target animals in aquatic habitats. Cladocerans, fishes and larvae of anuran amphib- viridis; 350 individual locations with equal sampling effort among the three taxa) share the ians are commonly used indicator organisms in ecotoxicological studies of pesticides. How- catchment of the Sofia Valley (259,343 ha, altitudinal range between 550 m and 2,290 m ever, little is known about the influence of these chemicals on other taxa inhabiting small a. s. l., western Bulgaria). Our main goals were to calculate the overlap between their spatial water bodies. Our aim was to directly examine the impacts of a frequently applied pesticide niches and to evaluate the dominant factors defining the spatial niches. Environmental pre- (Glyphogan®) on two top predators of temporary ponds. We used adult males of the smooth dictors used for spatial distribution models (SDMs) included climate parameters, geographic newt (Lissotriton vulgaris) and larvae of the dragonfly Aeshna cyanea, which are common in characteristics of the relief and habitats. The dataset was processed with the softwares Maxent central European wetlands. We kept predators individually in outdoors mesocosms containing and NextGIS. The output rasters were then reclassified to four classes: unsuitable, with low either no or a low dose of Glyphogan®. We fed predators with two Rana dalmatina tadpoles suitability, medium suitability, and optimal. every other day. We measured the effect of the pesticide on survival and behaviour of indi- Spatial niche overlap was lowest between L. a. bosnica and L. a. chersonensis (0.28% for low viduals. We also evaluated changes in body mass by measuring predators before and after the and 0.02% for the medium suitability classes); between L. a. bosnica and L. v. viridis was three weeks of the experiment. Finally, we tested for potential effects of Glyphogan® on pre- slightly higher: 2.85% (low suitability), 0.78% (medium suitability) and 0.19% (optimal); dation efficiency by providing predators with 10 R. dalmatina tadpoles and counting sur- between L. a. chersonensis and L. v. viridis was with the highest values, respectively 19.95%, vivors 24 h later. We performed this bioassay both in clear water and in water containing 4.15% and 0.54%. Glyphogan® with all predators in a random order. At the concentration applied and in the The most important for the SDMs for all taxa were altitude and habitats. For each taxa dif- conditions of our experiment, we did not observe significant effects of Glyphogan® on ferent climatic parameters were most important: for L. a. bosnica, maximum temperature of changes in body mass, survival, behaviour or feeding activity either in L. vulgaris or in A. the warmest month, mean temperature of the driest quarter and precipitation of the wettest cyanea. Comparing the two predators, we found that larval dragonflies killed more tadpoles, month that correspond to its mountainous distribution; isothermality and precipitation of than did newts, but this result did not depend on the chronic or the acute exposure to the driest/warmest quarter explain well the present distribution of L. a. chersonensis which is Glyphogan®. From our results it seems that Glyphogan® has no direct effects on the studied confined to the western Sub-Balkan valleys; for the more thermophilous L. v. viridis minimum top predators of temporary ponds. It remains to be seen if the chemical still affects predators temperature of the coldest month and annual precipitation explain best its distribution. via indirect effects, acting through the trophic cascade. In conclusion, the spatial niche segregation between the three taxa is well pronounced and is determined by common geographical factors but different specific climatic requirements.

300 301 BEHAVIOUR DISEASES AND PARASITES

Behavioural consistency and life history in Rana dalmatina tadpoles Distribution of Polystomoides ocellatum (Monogenea: Polystomatidae) in Spain parasitizing the European pond turtle (Emys orbicularis) Tamás János Urszán 1, János Török 1, Attila Hettyey 2, Gábor Herczeg 1 Aitor Valdeón 1,2, Virginia Rada 2, César Ayres 3, Raúl Iglesias 4, 1 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, Luis Alberto Longares 1, Raquel Lázaro 5, Vicente Sancho 6 Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 2 “Lendület” Evolutionary Ecology Group, Research Centre of Agricultural Sciences, Hungarian Academy of Sciences, Nagykovácsi út 26–30, 1029 Budapest, Hungary 1 Department of Geography and Regional Planning, University of Zaragoza, Pedro Cerbuna 12, 50009 Zaragoza, Spain; [email protected] 2 Recent interest in evolutionary behavioural ecology has turned towards understanding the Department of Herpetology, Aranzadi Society of Sciences, Zorroagagaina 11, 20014 Donostia-San Sebastian, Spain 3 Asociación Herpetológica Española, Apdo. de Correos 191, 28911 Leganés, Spain causes and consequences of behavioural consistency, manifesting either as animal personality 4 Laboratorio de Parasitología, Facultad de Biología, Universidad de Vigo, Campus Universitario Las Lagoas, (i. e. consistency in a single behaviour) or behavioural syndromes (i. e. consistency across two 36310 Vigo, Spain or more behaviours). Further, behavioural consistency has been linked to life-history strategies 5 Avda. Islas Baleares, 14 1ºE, 09006 Burgos, Spain 6 currently, possibly forming integrative pace-of-life syndromes (POLS). However, empirical CITMA, Ciutat Administrativa 9 d’Octubre-Torre 1 (3ª planta) c/Castán Tobeñas 77, 46018 Valencia, Spain tests of this theory are extremely scarce. Here, we tested, using agile frog (Rana dalmatina) The genus Polystomoides is an oixenic parasite of freshwater turtles, with more than 30 tadpoles as model, if POLS could be detected in the larval stage of amphibians based on a described species. Polystomoides ocellatum parasitize the buccopharyngeal cavity of the common garden experiment targeting two ontogenetic stages and measuring three behavi- European pond turtle. This species has a wide distribution range, after having been recorded oural traits. Activity was consistent in the early stage and was negatively related to age at in several countries; Old USSR, Ukraine, Romania, Bulgaria, Italy, France, and perhaps metamorphosis. In the later stage, activity, exploration and risk-taking was all consistent, but Tunisia. their relationship with life-history depended on whether the studied individuals were tested In Spain, P. ocellatum was recorded in European pond turtles from Tudela (Navarre, north- in the earlier stage or not. Already tested individuals showed no POLS, but both activity ern Spain) three years ago. Since then, more samplings have been carried out in several popu- (negatively) and risk-taking (positively) was related to age at metamorphosis in their previ- lations of the Ebro basin, and in other locations in Spain. Buccal swabs were used to detect ously untested conspecifics. Individual-level estimate of consistency in exploration (not the adult parasites in the buccopharyngeal area of the pond turtles, and Polystomoides were mean behaviour) was also negatively related to age at metamorphosis in the previously untested located without causing any damage to the turtles. individuals. We only detected one behavioural syndrome: activity and risk-taking was strongly In Cuenca (Central Spain) samplings P. ocellatum was not found, while its presence seemed negatively correlated in the previously tested (i. e. disturbed) group. Our results provide evi- to be widespread in the Emys orbicularis populations from the Ebro basin. Two specimens dence for (i) animal personality in tadpoles, (ii) an activity–metamorphosis-timing POLS were found in a population located in the region of Valencia (eastern Spain), and one in and (iii) an unexpectedly strong effect of moderate disturbance related to standard behavioural Galicia (Northwestern Spain). testing on later behavioural consistency. In this study we show the confirmed presence of P. ocellatum in relation to Emys orbicularis in a distribution map of 10x 10 km UTM squares. More samples are needed from other Iberian and North African populations to elaborate further this hypothesis about the natural history of this species.

302 303 PHYLOGENY BEHAVIOUR

Evaluation of the evolutionary history of Podarcis cretenis (Sauria: Common lizard: microhabitat choice in a subarctic habitat Lacertidae) Balázs Vági 1,2, Tibor Kovács 1,3, Judit Cservenka 4, Tamás Telbisz 5 Alexis Vasilikopoulos 1,2, Agalia Antoniou 3, Antonios Magoulas 3, Petros Lymberakis 1, 1 Efstratios Valakos 4, Nikos Poulakakis 1,2 Behavioural Ecology Group, Department of Systematic Zoology and Ecology, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary; [email protected] 2 “Lendület” Evolutionary Ecology Research Group, Plant Protection Institute, Centre for Agricultural Research, 1 Natural History Museum of Crete, University of Crete, Knosos Avenue, 71409 Irakleio, Greece; [email protected] Hungarian Academy of Sciences, Herman Ottó út 15, 1022 Budapest, Hungary 2 Department of Biology, University of Crete, Vassilika Vouton, 71409 Irakleio, Greece 3 Hungarian Biodiversity Research Society, Hunyadvár utca 43/a, 1165 Budapest, Hungary 3 Institute of Marine Biology and Genetics, Hellenic Centre for Marine Research, Gournes Pediados, 71003 Irakleio, 4 Balaton-felvidéki National Park Directorate, Kossuth utca 16, 8229 Csopak, Hungary Crete, Greece 5 Department of Physical Geography, Eötvös Loránd University, Pázmány Péter sétány 1/c, 1117 Budapest, Hungary 4 Section of Animal and Human Physiology, Department of Biology, University of Athens, Ilissia, 15784 Athens, Greece Common lizard (Zootoca vivipara) is a reptile inhabiting the northernmost parts of the world, Recently, several works have focused on lacertid lizards of the genus Podarcis, revealing cases in Scandinavia and Siberia crossing even the Arctic Circle. In spite of the fact being an of hidden diversity and paraphyly, and offering evidence that suggests the alteration of the ectotherm, it adapted very well to harsh climatic conditions. Thermoregulation in subarctic extant taxonomical arrangements within the genus. The Balkan species P. peloponnesiaca and habitats can have substantial costs, still on the basis of our earlier surveys it is worth for lizards P. erhardii fall in such a case of hidden diversity and paraphyly. To deal with this taxonomic to keep accurate thermal control. Temperature distribution is not the only condition having problem, the populations from the island of Crete and its satellite islets were assigned to a a significant role in thermal control; solar radiation and the choice between various micro- new species (P. cretensis), which is endemic to this island group. On the island of Crete, its habitats are just as important. In our survey conducted at the latitude of the Arctic Circle in distribution pattern is peculiar since the species is present only on the western part, grossly Finland, we wanted to explore to what extent solar radiation and the physical structure of to the west of a N–S axis “cutting” Crete at the town of Rethymno. Here, we inferred vegetation influence the habitat choice of lizards. Only open or semi-open microhabitats of intraspecific relationships, and tested hypotheses for historical patterns of population expan- a meadow abandoned 40 years ago were involved in the survey. We developed a radiation sion, with phylogenetic and phylogeographic approaches using both mitochondrial model and a vegetation map of the lizards’ habitat, and then compared the solar radiation (mtDNA) and nuclear (msats) markers. The phylogenetic relationships supported the mono- and microhabitat distribution of the detection points of lizards and of all the points of the phyly of P. cretensis and suggest that three main haplogroups are present on the island. These habitat. According to our results, the radiation values of the individual microhabitats do not groups have diverged in Pliocene indicating that present distribution of P. cretensis was deter- differ; therefore the character of vegetation is influenced by other ecological variables. How- mined by a combination of dispersal and vicariance. ever, neither because of solar radiation nor on the basis of habitat is the distribution of lizards random. The lizards preferred the points extended to solar radiation for longer periods, and occurred more frequently in areas covered by high grass. According to our results we can state that lizards do have preferences based on solar radiation and microhabitat, and these two choices are independent from each other. Thermoregulation, as well as other physiological constraints, such as food supply and shelter availability can be decisive criteria.

304 305 PHYLOGEOGRAPHY PHYLOGEOGRAPHY

Higher than expected genetic diversity in a small area: Mitochondrial DNA diversity of the Alpine newt, Mesotriton alpestris Zootoca vivipara in the eastern part of Hungary in the Carpathian Basin

Balázs Velekei 1, Ferenc Lakatos 1, Péter Bíró 2, Éva Ács 3, Miklós Puky 3 Judit Vörös 1, Krisztián Szabó 2

1 University of West Hungary, Institute of Silviculture and Forest Protection, Bajcsy-Zs. u. 4, 9400 Sopron, Hungary; 1 Department of Zoology, Hungarian Natural History Museum, Baross u. 13, 1088 Budapest, Hungary [email protected] 2 Institute of Biology, Faculty of Veterinary Science, Szent István University, Rottenbiller u. 50, 1078 Budapest, 2 Hungarian Academy of Sciences, Centre For Ecological Research, Balaton Limnological Institute, Hungary; [email protected] Klebelsberg K. út 1–3, 8237 Tihany, Hungary 3 Hungarian Academy of Sciences, Centre For Ecological Research, Danube Research Institute, Jávorka S. utca 14, The Alpine newt, Mesotriton alpestris occurs with isolated populations in the low-elevation 2131 Göd, Hungary mountains of the Carpathian Basin, surrounded by the Carpathian Mountains and the western Alpine domain. Using 1407 bp of the mitochondrial genome (fragments of the ND4 plus The Eurasian common lizard, Zootoca vivipara (Lichtenstein, 1823), is the lizard with the adjacent tRNAs and 16s genes) we inferred the genetic diversity and evolutionary relation- largest distribution area on Earth. It has several subspecies/clades, some with extremely large, ship of these isolated Carpathian Basin populations. We collected samples from five regions others with more limited distribution areas. Several papers were published recently about of the Hungarian mid-elevation mountains (Ôrség, Bakony Mts., Mátra Mts., Bükk Mts., the phylogenetic relationships of the viviparous and oviparous haplogroups of the species Zemplén Mts.) covering the distribution of the species in Hungary, plus from Mt. Apuseni describing clades living only in Central Europe. Our aim is to describe the phylogeographical in Romania. Our sequences were compared with unpublished sequences representing the rest structure of Z. vivipara in the Carpathian Basin. As a first step we studied Hungarian popu- of the species’ range in Europe. The phylogenetic analyses split our samples into two major lations. Samples were collected from eight locations. Total DNA was extracted from small tis- clades representing the Eastern and Western European clades. The samples from Apuseni sue samples of the tail. Two target genes were selected for the phylogenetic analysis: a partial Mts. are closely related to the Southern Carpathian lineage being part of the Eastern clade. sequence (appr. 300 bp) of the protein encoding cytochrome b (cyt b) and a partial sequence All the Hungarian samples belonged to the Western clade, but with significant substructuring (appr. 500 bp) of the non-protein coding 16s. Sequences of four lacertid species were down- among the Alpine (Ôrség), Western Transdanubian Range (Bakony) and Northern Mountain loaded from the GenBank and were used as outgroups. Maximum likelihood analyses of the Range (Mátra, Bükk, Zemplén) samples. Within this clade Bakony Mts. is a sister taxa of the data revealed a congruent overall topology that is characterized by five main groups: the PA rest of the clade, suggesting that this population might have survived Pleistocene glaciations haplogroup is present in the northwestern, the VH haplogroup in the central part of the within the Carpathian Basin in a separate glacial refugium. country. The other samples collected from the northeastern part of Hungary are in the con- tact zone of VB and VU haplogroups and were divided into three groups. Our study sup- ports the model of four haplogroups being present in the Carpathian Basin, but comparing our results to the previous studies we detected higher presence of the VU haplogroup in the eastern region of the country. Furthermore, the VU haplogroup is divided into two main groups, representing high genetic diversity in a relatively small area.

306 307 PRACTICAL CONSERVATION SYSTEMATICS

Effects of water contamination on site selection by amphibians: Resolving sky island speciation of East African Adolfus alleni experiences from an arena approach with European frogs and newts (Sauria, Lacertidae)

Norman Wagner, Stefan Lötters Philipp Wagner

Trier University, Biogeography Department, Universitätsring 15, 54296 Trier, Germany; [email protected] Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; and Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; Pesticide residues in breeding ponds can cause avoidance by at least some amphibian species. [email protected] So far, outdoor experiments have been performed only with artificial pools in areas where a tar- The genus Adolfus Sternfeld, 1912 currently contains three species from Equatorial Africa. get species occurs and new colonizations of water bodies have been observed. Results of this Two of these occur in widespread, low to mid-elevation, habitats but Adolfus alleni is only kind of study are potentially influenced by natural disturbances and therefore are of limited known from four montane peaks (Aberdares, Mt. Kenya, Cherangani Hills, Mt. Elgon) in comparability. We used an easily-to-establish standardizable arena approach, in which animals Kenya and Uganda. An integrative approach using 58 morphological characters and genetic in reproductive condition for some hours had the choice among pools with different concent- analyses of mitochondrial (16S and cyt b) and nuclear (c-mos and RAG1) DNA sequence rations of a contaminant. Because of an ongoing debate on the potential environmental impacts data revealed differences between these populations, and indicates that A. alleni is a complex of glyphosate-based herbicides, we investigated the impact of glyphosate isopropylamine salt, of at least two cryptic species. Herein, we describe the populations from the Aberdares and Roundup® LB PLUS and glyphosate’s main metabolite aminomethylphosphonic acid (AMPA) Mount Elgon as a new species, and restrict A. alleni to Mount Kenya. This action underscores on individual residence time in water. The following European amphibian species were tested: the importance of conservation strategies to protect these montane peaks, which may harbor common frog (Rana temporaria), palmate newt (Lissotriton helveticus) and Alpine newt (Ichthyo- additional, unique evolutionary lineages. saura alpestris). A total of each 60 pairs of palmate and Alpine newts (n = 240) and 50 male common frogs were sampled in two ponds near Trier, Germany. Experiments were performed in arenas with 10 replicates/night from March 12 to 28. An arena (1.5 x 1.5 m) was defined by an amphibian drift fence and consisted of a hiding place (leaves and planks) and four artificial pools (plastic pans of 34 x 34 cm, each with a 10-L capacity and a 13-cm maximum depth). Beside controls, tested AMPA concentrations were 0.5 (slightly above the European Environmental Quality Standard for AMPA: 0.45 mg/L), 0.05, and 0.005 mg/L; tested glyphosate and Round- up® concentrations were 1.0 (slightly above the German ‘worst-case’ expected environmental concentration: 0.9 mg a. e./L), 0.1 (the European Environmental Quality Standard for glypho- sate), and 0.01 mg a. e./L. To compare the residence time (min) of individuals in the four groups of contamination, we used Kruskal-Wallis tests followed by Wilcoxon rank sum tests (data were not normally distributed). In addition, we tested with generalized linear models (GLMs) if the different concentrations or considered environmental cofactors influenced the percentage resi- dence time of animals of each trial (i. e., percentage residence time of all individuals tested in one night) as proposed for such behavioral studies where uninteresting behavior (here, time spent on land) was excluded from analyses. Considered environmental cofactors were average water tem- perature, nightly minimum ground temperature, precipitation, relative air humidity. The resi- dence time in water was not significantly affected by any concentration. Occasionally, micro- climatic cofactors (nightly minimum ground temperature, water temperature) apparently influ- enced the residence time. The major drawback of such quick arena approach is that results can be transferred to perception and avoidance of contaminated water rather than to site selection by the specimens. Testing for oviposition site selection may require more natural water bodies and a longer study period. Hence, for the development of a standard procedure in risk assessments, an intermediate design between an arena approach, as presented here, and previously performed field studies may be applicable. 308 309 SYSTEMATICS SYSTEMATICS

Island zoogeography at a continental scale An extraordinary tail—integrative review of the agamid genus Xenagama Boulenger, 1895 Philipp Wagner Philipp Wagner Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; and Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; Zoologisches Forschungsmuseum Alexander Koenig, Adenauerallee 160, 53113 Bonn, Germany; [email protected] and Department of Biology, Villanova University, 800 Lancaster Avenue, Villanova, Pennsylvania 19085, USA; [email protected] Africa is well-known for its biodiversity and especially for its endemicity of restricted zoo- geographic areas or single mountain systems. Opposite, little is known about widespread groups Approximately 120 years after its description, this is the first thorough systematic review of occurring across the continent, such as Hemidactylus (Gekkonidae), Trachylepis (Scincidae), the agamid genus Xenagama. Currently, the genus includes two extraordinary species, char- or Boaedon (Colubridae). Lizards in the genus Agama (~45 species) have a pan-continental acterized by strongly discoidal tails. Both species are only known from a few localities in distribution and the evolution of conspicuous sexual dimorphisms accompanied by a vari- northeastern Ethiopia and northern Somalia and are represented by a small number of speci- ability in social systems among colorful versus drab species suggests that these traits may have mens in museum collections. Morphological and mitochondrial (16S) DNA sequence data had a profound impact on lineage diversification. The poster presents a comprehensive time- revealed that X. batillifera is a complex of two cryptic species and the reassignment of calibrated species tree for most African Agama species using five independent loci augmented Acanthocercus zonurus to the genus Xenagama is supported by morphological analyses. Herein, with anchored phylogenomic data from nuclear genes for 23 species to strengthen the back- we present the results of a recent paper in which we described specimens previously re- bone of the phylogeny. The origin, diversity and evolution of African agamas are discussed cognized as a geographic variant of Xenagama batillifera as a new species and compare it to in the light of these new phylogenetic relationships. other whorl-tailed lizards of the genera Xenagama and Acanthocercus from the Horn of Africa, underscoring the significance of this poorly known but important region for agamid evolu- tion. Among other features, the new species is characterized by possessing heterogeneous body scalation, no nuchal crest and no tufts of elongated scales around the ear. Similar to X. batillifera or X. taylori, and distinct to other African agamid lizards, the tail is shorter than the body and head, but the tail base is less discoidal than in X. batillifera or especially X. tay- lori and gradually merges into the terminal filament. The discoidal part of the tail is arranged in whorls with one scale ring each, whereas the filament is not distinctly whorled.

310 311 TROPICAL HERP BIODIVERSITY BEHAVIOUR

Amphibian diversity in Tarutao National Park and capacity-building Fish introduction affects amphibians by increasing avoidance in amphibian research and conservation in peninsular Thailand behaviour and inhibiting sexual activity

Sansareeya Wangkulangkul 1, Paul Bates 2, Chutamas Satasook 1, Tshering Nidup 1, Laurane Winandy, Mathieu Denoël Judit Vörös 3 Laboratory of Fish and Amphibian Ethology, Behavioural Biology Unit, University of Liège, 22 Quai van Beneden, 4020 Liège, Belgium; [email protected] 1 Department of Biology, Faculty of Science, Prince of Songkla University, Hat Yai, Songkhla 90112, Thailand; [email protected] 2 Harrison Institute, Bowerwood House, 15 Botolph’s Road, Sevenoaks, Kent, TN13 3AQ, United Kingdom Amphibians are currently declining worldwide for a variety of reasons. One of the major 3 Hungarian Natural History Museum, Baross u. 13, 1088 Budapest, Hungary causes is the introduction of fish in their natural habitat. Since many amphibian populations do not usually share a common evolutionary history with fish, the detrimental impact of The project seeks to promote anuran conservation in peninsular Thailand and in particular these introductions is understandable. A way for amphibians to persist in such altered envi- in the islands of Tarutao National Park. Our team is led by staff from the Prince of Songkla ronments is to adopt adequate behavioural tactics. However, although it has been shown that University and the Hungarian Natural History Museum and comprises students from the avoidance behaviour may decrease the probability of being detected by a potential predator, Prince of Songkla University and staff of the Tarutao National Park. All are being trained in little is known on fitness consequences, particularly in terms of sexual behaviour. In this anuran identification, ecology and disease, with particular reference to the fungus disease study, we aimed at testing the impact of goldfish (Carassius auratus) on sexual activity and Batrachochytrium dendrobatidis. From the initial field research, 12 amphibian species were mating tactics in adult Alpine newts (Ichthyosaura alpestris) in risky and safe micro-habitats. found, including a new form of Hylarana eschatia and Ingerophrynus parvus—the morpho- To this end, we compared behavioural patterns of newts in function of fish presence in a metrics and ecology of all species were studied. To date, Bd fungus infection has not been replicated laboratory design. Consistently through the whole breeding period, there was a found in Tarutao National Park; however, we continue to monitor for this. We intend to significant effect of fish on the behaviour of newts. Fish presence caused an increased shelter expand this project to other national parks in peninsular Thailand and have already under- use and a decreased sexual activity; this particularly in risky habitats. Moreover in the presence taken a provisional study at Hala Bala Wildlife Research Station. of fish, courtship displays varied and encounters were less successful than in their absence. These results show that fish presence can affect newts in complex ways, including the inhi- bition of their reproduction. They also highlight the importance of integrating behaviour into conservation studies for a better understanding of the interactions between native amphibians and introduced fish.

312 313 MORPHOLOGY, ANATOMY AND DEVELOPMENT

Geometric morphometric analysis of cranial variation in paedomorphic and metamorphic forms of the northern banded newt, List of Participants Ommatotriton ophryticus

Elif Yildirim, Elnaz Najafi-Majd, Ug˘ur Kaya

Ege University, Faculty of Science, Biology Section, Zoology Department, 35100 Bornova-Izmir, Turkey; Ahmadzadeh, Faraham Baptista, Ninda [email protected] Zoologisches Forschungsmuseum Alexander Koenig, Universidade de Lisboa, Biologia Animal, Herpetology Section, Adenauerallee 160, 53113 Bonn, Campo Grande, 1749-016 Lisboa, Portugal; The northern banded newt, Ommatotriton ophryticus has been listed as Near Threatened Germany; [email protected] [email protected] (NT) in the Red List of IUCN. It belongs to the family Salamandridae and can be found in Aowphol, Anchalee Barquero, Marco northwestern Turkey, northwestern Armenia, Georgia, and southern Russia. Kasetsart University, Department of Zoology, Macquarie University, Biological Sciences, An aquatic larval stage is followed by metamorphosis to reach terrestrial juvenile and then Faculty of Science, P.O. Box 1020, 10900 Bangkok, Faculty of Science, 2109 North Ryde, Australia; adult stage in amphibians. However many species of salamanders exhibit paedomorphosis, Thailand; [email protected] [email protected] i. e. retain ancestral juvenile characteristics by adult stages of descendents. Paedomorphic forms Arntzen, Jan W. Bauer, Aaron of O. ophryticus were first described in 2008 from near Karasu, in the vicinity of Sakarya, Naturalis Biodiversity Center, Terrestrial Zoology, Villanova University, Department of Biology, Turkey. Also, there are many species of European newts that show paedomorphic form such Darwinweg 2, 2300RA Leiden, The Netherlands; 800 Lancaster Avenue, Villanova, PA 19085, USA; as Lissotriton vulgaris, Triturus macedonicus, and Ommatotriton vittatus. [email protected] [email protected] Patterns of cranial variation in 27 paedomorphic and metamorphic specimens of O. ophry- Ayres, Cesar Bellakhal, Meher ticus were analyzed using geometric morphometric methods. Dorsal views of the cranium of Asociación Herpetologica Espanola (AHE), Institute Supérieur de Peche et d’Aquaculture de all specimens were photographed with Olympus SZ61 Stereoscope equipped with an Olympus Apdo de Correos 191, 28911 Madrid, Spain; Bizerte, Exploitation des Milieux Aquatiques, digital camera. The analysis was performed on the coordinates of 13 landmarks defined on [email protected] B. P. 15, 7080 Errimel, Tunisia; one side of the cranium of each specimen (tpsDig V.1.40). The relative warps analysis was [email protected] done tpsRelw programe, version 1.42. Patterns of cranial shape change were analyzed by Babocsay, Gergely Mátra Museum of the Hungarian Natural History Bitenc, Katja tpsSplin (tpsSplin V.1.20) to demonstrate cranial shape changes as deformations. Shape analysis Museum, Collection of Vertebrates, Biotechnical Faculty, University of Ljubljana, showed that there was no significant differentiation between paedomorphic and metamorphic Kossuth L. u. 40, 3200 Gyöngyös, Hungary; Department of Biology, Vecna pot 111, 1000 Ljubljana, specimens of O. ophryaticus. [email protected] Slovenia; [email protected]

Bajer, Katalin Blooi, Mark Universidade Federal do Rio Grande do Norte, Ghent University, Faculty of Veterinary Medicine, Department of Botany, Ecology and Zoology, Department of Pathology, Bacteriology and Avian Natal 59078-900, Brazil; [email protected] Diseases, Salisburylaan 133, 9820 Merelbeke, Belgium; [email protected] Bakó, Botond Ministry of Rural Development, Department of Bogaerts, Sergé Rural Development, Kossuth L. tér 1, 1055 Budapest, Lupinelaan 25, 5582CG Waalre, The Netherlands; Hungary; [email protected] [email protected]

Balazˇ, Vojteˇch Böhm, Stephan University of Veterinary and Pharmaceutical Megazoo Brunn, 2345 Brunn, Austria; Sciences Brno, Faculty of Veterinary Hygiene and [email protected] Ecology, Department of Biology and Wildlife Diseases, Palackého tr. 1/3, 61242 Brno,Czech Republic; [email protected]

314 315 Böhme, Wolfgang Carretero, Miguel Cservenka, Judit Diedericks, Genevieve Zoologisches Forschungsmuseum Alexander Koenig, CIBIO, Centro de Investigação em Biodiversidade e Balaton Uplands National Park Directorate, Stellenbosch University, Botany & Zoology, Herpetology Section, Adenauerallee 160, 53113 Bonn, Recursos Genetico Vairão, Campus Agrário de Vairão, Department of Nature Conservation, Private bag X1, 7602 Matieland, South Africa; Germany; [email protected] Rua Padre Armendo Quintas, 4485-661 Vairão, Cholnoky u. 23/a, 8200 Veszprém, Hungary; [email protected] Portugal; [email protected] [email protected] Böning, Philipp van Diepenbeek, Annemarie Trier University, Department of Biogeography, Cherlin, Vladimir Damas Moreira, Isabel RAVON (Reptile Amphibian Fish Conservation Universitätsring 15, 45259 Trier, Germany; Zoological Park, Exotarium, Aleksandrovskiy park 1, CIBIO, Centro de Investigação em Biodiversidade e Netherlands), P.O. Box 1413, 6501BK Nijmegen, [email protected] 197198 Saint-Petersburg, Russia; [email protected] Recursos Genetico Vairão, Campus Agrário de Vairão, The Netherlands; [email protected] Rua Padre Armendo Quintas, 4485-661 Vairão, Borczyk, Bartosz Cirovic´, Ruza Portugal; [email protected] Djurakic´, Marko Univeristy of Wroclaw, Department of Evolutionary Environmental Protection Agency of Montenegro, University of Novi Sad, Faculty of Sciences, Biology and Conservation of Vertebrates, Department for Monitoring, Analysis and Reporting, Dankovics, Róbert Department of Biology and Ecology, Sienkiewicza 21, 50-335 Wroclaw, Poland; 4th Proleterska No. 19, 81000 Podgorica, Montenegro; Savaria Museum, Department of Natural History, Trg Dositeja Obradovica 2, 21000 Novi Sad, [email protected] [email protected] Kisfaludy S. u. 9, 9700 Szombathely, Hungary; Serbia; [email protected] [email protected] Bosman, Wilbert Clemann, Nick Djuric´, Dragana RAVON (Reptile Amphibian Fish Conservation Arthur Rylah Institute for Environmental Research, Davoli, Francesca Natural History Museum, Department of Netherlands), P.O. Box 1413, 6501BK Nijmegen, Environment and Primary Industries, 123 Brown St., ISPRA, Conservation Genetics Laboratory, Paleontology, Njegoseva 51, 11000 Belgrade, Serbia; The Netherlands; [email protected] Heidelberg, Victoria 3084, Australia; Via Ca’ Fornacetta 9, 40064 Ozzano Emilia [email protected] [email protected] (Bologna), Italy; [email protected] Broeckhoven, Chris Dobay, Gergely Stellenbosch University, Botany & Zoology, Coga˘lniceanu, Dan Degani, Gad Szent István University, Institute of Botany and Private bag X1, 7602 Matieland, South Africa; Ovidius University of Constant¸a, Faculty of Tel Hai Academic College, Biotechnology, Ecophysiology, 2100 Gödöllo˝, Hungary; [email protected] Natural Sciences, Aleea Universita˘¸iit nr. 1, Corp B, P.O. Box 831, Herzel St. 71/2, 110 Kiryat Smona, [email protected] 900470 Constant¸a, Romania; Israel; [email protected] Bulakhova, Nina [email protected] Drakulic´, Sanja Institute of Biological Problems of the North, van Delft, Jeroen Museum für Naturkunde, Invalidenstraße 43, Russian Academy of Sciences, Biocenology, Cortada, Alex RAVON (Reptile Amphibian Fish Conservation 10115 Berlin, Germany; [email protected] Portovay St. 18, 685000 Magadan, Russia; Universitat de Barcelona, Facultat de Biologia, Netherlands), P.O. Box 1413, 6501BK Nijmegen, [email protected] Av. Diagonal 643, 08028 Barcelona, Spain; The Netherlands; [email protected] Dubey, Sylvain [email protected] University of Lausanne, Ecology and Evolution, Bury, Stanislaw Denoël, Mathieu UNIL Sorge Le Biophore, 1015 Lausanne, Jagellonian University, Institute of Environmental Corti, Claudia University of Liège, Behavioural Biology Unit, Switzerland; [email protected] Sciences, Life History Evolution Group, Museo di Storia Naturale dell’Universitá di Firenze, 22 Quai van Beneden, 4020 Liège, Belgium; Gronostaiowa 7, 30-387 Kraków, Poland; Zoological Section, Via Romana 17, 50125 Firenze, [email protected] Eckerlein, Anna [email protected] Italy; [email protected] Trier University, Department of Biogeography, Derus, Natalia Universitätsring 15, 66482 Trier, Germany; Cabido, Carlos Creemers, Raymond Jagiellonian University, Institute of Environmental [email protected] Aranzadi Society of Sciences, Herpetology, RAVON (Reptile Amphibian Fish Conservation Sciences, Life History Evolution Group, Alto de Zorroaga 11, 20014 Donostia–San Sebastián, Netherlands), Volunteers & Data, P.O. Box 1413, Gronostajowa 7, 30-387 Kraków, Poland; Erich, Magdalena Spain; [email protected] 6501BK Nijmegen, The Netherlands; [email protected] University of Vienna, Evolutionary Biology, [email protected] Meyerbeergasse 17, 1210 Wien, Austria; Canessa, Stefano Di Cerbo, Anna Rita [email protected] ARC Centre of Excellence for Environmental Crnobrnja-Isailovic´, Jelka Centro Studi Faunistica dei Vertebrati, Decisions, School of Botany, University of University of Nisˇ, Faculty of Sciences and Italian Society of Natural Sciences, Corso Venezia 55, Erôss, Mariann Melbourne, 3010 Melbourne, Australia; Mathematics, Biology and Ecology, Visˇegradsˇka 33, 20121 Milano, Italy; [email protected] University of Debrecen, Department of Evolutionary [email protected] 18000 Nisˇ, Serbia; [email protected] Zoology and Human Biology, Egyetem tér 1, 4400 Debrecen Hungary; [email protected]

316 317 Famelli, Shirley Fritz, Uwe Golubovic´, Ana Hauswaldt, Susanne Universidade de São Paulo, Escola Superior de Senckenberg Natural History Collections Dresden, Belgrade University, Faculty of Biology, TU Braunschweig, Zoology, Mendelssohnstraße 4, Agricultura Luiz de Queiroz, Ciências Biológicas, Department of Museum of Zoology, Department of Morphology, Systematics and 38106 Braunschweig, Germany; [email protected] Institute for Applied Ecology, Av. Padua Dias 11, Königsbrücker Landstraße 159, 01109 Dresden, Phylogeny of Animals, Studentski trg 16, 13418-700 Piracicaba, Brazil; [email protected] Germany; [email protected] 11000 Belgrade, Serbia; [email protected] Heard, Geoffrey University of Melbourne, School of Botany, Familiar Lopez, Mariel Gabor, Caitlin Gregorovicˇová, Martina Building 122, Parkville, Victoria 3010, Australia; Griffith University, Environmental Future Centre, Texas State University, Biology, 601 University Drive, Charles University, Faculty of Nature, Vinicna 7, [email protected] School of Environment, Parklands Drive G24, 78666 San Marcos, USA; [email protected] 12844 Praha, Czech Republic; [email protected] 4222 Southport, Australia; Heltai, Botond István [email protected] Gál, Zoltán Grillitsch, Heinz Szent István University, Department of Zoology and Plant Protection Institute for Agricultural Research, Natural History Museum Vienna, 1st Zoological Animal Ecology, Páter Károly utca 1, 2100 Gödöllo˝, Farkas, Balázs Hungarian Academy of Science, Department (Herpetology), Burgring 7, 1010 Wien, Hungary; [email protected] Bercsényi u. 21, 2464 Gyúró, Hungary; “Lendület” Evolutionary Ecology Research Group, Austria; [email protected] [email protected] Herman Ottó út 15, 1022 Budapest, Hungary; Herczeg, Dávid [email protected] Gross, Ina University of Debrecen, Department of Evolutionary Fera, Gábor Institute of Pharmacy and Molecular Biotechnology, Zoology and Human Biology, Egyetem tér 1, Plant Protection Institute for Agricultural Research, Gál, Júlia Tünde University of Heidelberg, Department of Biology, 4032 Debrecen, Hungary; [email protected] Hungarian Academy of Science, Szent István University, Department of Ecology, Im Neuenheimer Feld 364, 4. OG, “Lendület” Evolutionary Ecology Research Group, István u. 2, 1078 Budapest, Hungary; 69120 Heidelberg, Germany; [email protected] Herder, Jelger Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] RAVON (Reptile Amphibian Fish Conservation [email protected] Grozdanov, Atanas Netherlands), P.O. Box 1413, 6525ED Nijmegen, Galoyan, Eduard University of Sofia, Faculty of Biology, The Netherlands; [email protected] Ferreira, Mota Joint Russian–Vietnamese Tropical Research and Opalchenska St. 41a, 1233 Sofia; Bulgaria; ICETA, Instituto de Ciências e Tecnologias Agrárias Technological Center of the A.N. Severtsov [email protected] Hero, Jean-Marc e Agro-Alimentares, Rua D. Manuel II, Institute of Ecology and Evolution, 33 Leninskij Griffith University, Environmental Future Centre, Apartado 55142, 4051-401 Porto, Portugal; prosp., 119071 Moscow, Russia; [email protected] Gvozˇdik, Václav School of Environment, Parklands Drive G24, [email protected] University of Basel, Department of Environmental 4222 Southport, Australia; [email protected] Garcia Roa, Roberto Sciences, St. Johanns-Vorstadt 10, 4056 Basel, Ficetola, Gentile Francesco Museo Nacional de Ciencias Naturales, CSIC Switzerland; [email protected] Hettyey, Attila University of Milano-Bicocca, DISAT, Ecologia Evolutiva, José Gutiérrez Abascal 2, Plant Protection Institute for Agricultural Research, Piazza della Scienza 1, 20126 Milano, Italy; 28006 Madrid, Spain; [email protected] Hallermann, Jakob Hungarian Academy of Science, [email protected] University of Hamburg, Zoological Museum, “Lendület” Evolutionary Ecology Research Group, Garin Barrio, Ion Martin-Luther-King-Platz 3, 20146 Hamburg, Herman Ottó út 15, 1022 Budapest, Hungary; Filz, Katharina Aranzadi Society of Science, Herpetology, Germany; [email protected] [email protected] Trier University, Department of Biogeography, Alto de Zorroaga 11, 20014 Donostia–San Sebastián, Universitätsring 15, 54296 Trier, Germany; Spain; [email protected] Halpern, Bálint Hödl, Walter [email protected] MME-BirdLife Hungary, Amphibian and Reptile University of Vienna, Department of Integrative Gent, Antony Conservation Group, Költo˝ u. 21, 1121 Budapest, Zoology, Althanstraße 14, 1090 Wien, Austria; Flecks, Morris Amphibian and Reptile Conservation, Hungary; [email protected] [email protected] Zoologisches Forschungsmuseum Alexander Koenig, 655a Christchurch Road, Bournemouth BH1 4AP, Herpetology Section, Adenauerallee 160, 53113 Bonn, United Kingdom; [email protected] Hamer, Andrew Hölting, Monique Germany; [email protected] Australian Research Centre for Urban Ecology. Senckenberg Natural History Collections Dresden, Gollmann, Günter Royal Botanic Gardens, School of Botany, Department of Museum of Zoology, Foster, James University of Vienna, Theoretical Biology, University of Melbourne, Parkville, Victoria 3010, Königsbrücker Landstraße 159, 01109 Dresden, Amphibian and Reptile Conservation, Althanstraße 14, 1090 Wien, Austria; Australia; [email protected] Germany; [email protected] 655a Christchurch Road, Bournemouth BH1 4AP, [email protected] United Kingdom; [email protected] Hartmann, Timo Horváth, Gergely Zoologisches Forschungsmuseum Alexander Koenig, Eötvös Loránd University, Department of Systematic Herpetology Section, Adenauerallee 160, 53113 Bonn, Zoology and Ecology, Pázmány Péter sétány 1/c, Germany; [email protected] 1117 Budapest, Hungary; [email protected]

318 319 Ibañez, Alex Kielgast, Jos Krasa, Antonin Lorenzo, Marcial Museo Nacional de Ciencias Naturales, CSIC Amphi Consult, Saxogade 54, 1662 Copenhagen, Nature Conservation Agency of the Czech Republic, Salamanca University, Animal Biology, Faculty of Ecologia Evolutiva, José Gutiérrez Abascal 2, Denmark; [email protected] Kaplanova 1, 14800 Praha, Czech Republic; Pharmacy, Campus Miguel de Unamuno s/n, 28006 Madrid, Spain; [email protected] [email protected] 37007 Salamanca, Spain; [email protected] Kieren, Sarah Iglesias-Carrasco, Maider Trier University, Department of Biogeography, Krzysciak-Kosinska, Renata Ludwig, Gerda Aranzadi Society of Science, Herpetology, Universitätsring 15, 54296 Trier, Germany; Bialowieza National Park, Research Unit, University of Innsbruck, Institute of Zoology, Alto de Zorroaga 11, 20014 Donostia–San Sebastián, [email protected] Park Palacowy 11, 17-230 Bialowieza, Poland; Technikerstraße 25, 6020 Innsbruck,Austria; Spain; [email protected] [email protected] [email protected] Kindermann, Christina Ignacio, Martin Griffith University, Environmental Future Centre, Kurali, Anikó Luznik, Martina Polytechnic University of Madrid, School of Environment, Parklands Drive G24, Plant Protection Institute for Agricultural Research, University of Primorska, Faculty of Mathematics, Zoology Department, School of Forestry and 4222 Southport, Australia; Hungarian Academy of Science, Natural Sciences and Information Technologies, Natural Environment Engineering, [email protected] “Lendület” Evolutionary Ecology Research Group, Department for Biodiversity, Glagoljaska 8, Avda. Ramiro de Maezru s/n, 28040 Madrid, Herman Ottó út 15, 1022 Budapest, Hungary; 6000 Koper, Slovenia; [email protected] Spain; [email protected] Kindler, Carolin [email protected] Senckenberg Natural History Collections Dresden, Lymberakis, Petros Ihlow, Flora Department of Museum of Zoology, Kurek, Katarzyna University of Crete, Natural History Museum of Zoologisches Forschungsmuseum Alexander Koenig, Königsbrücker Landstraße 159, 01109 Dresden, Institute of Nature Conservation, Crete, Knossos Av., P.O. Box 2208, 71409 Irakleio, Herpetology Section, Adenauerallee 160, 53113 Bonn, Germany; [email protected] Polish Academy of Sciences, Mickiewicza 33, Greece; [email protected] Germany; [email protected] 31-120 Kraków, Poland; [email protected] Kirschey, Tom Macát, Zdenek Jablonski, Daniel NABU, Herpetology Scientific Board, Lagler, Patricia Palacky University, Department of Ecology & Comenius University, Faculty of Natural Sciences, 16775 Menz, Germany; [email protected] University of Natural Resources and Life Sciences, Enviromental Sciences, 78371 Olomouc, Department of Zoology, Mlynska dolina B-1, Gregor Mendel Straße 33, 1180 Wien, Austria; Czech Republic; [email protected] 84215 Bratislava, Slovakia; Kiss, István [email protected] [email protected] Szent István University, Department of Zoology and MacLeod, Amy Animal Ecology, Páter Károly utca 1, 2100 Gödöllo˝, Lambertz, Markus University of Bielefeld, Department of Animal Joger, Ulrich Hungary; [email protected] Poppelsdorfer Schloss, 53115 Bonn, Germany; Behaviour, Morgenbreede 45, 33615 Bielefeld, Staatliches Naturhistorisches Museum, [email protected] Germany; [email protected] Gausstraße 22, 38106 Braunschweig, Germany; Knorr, Konstantin [email protected] University of Veterinary Medicine, Institute of Lansari, Aziza Maletzky, Andreas Zoology, Tiestestraße 21, 30171 Hannover, Germany; University of Cadi Ayyad, Faculty of Sciences, University of Salzburg, Department of Organismic Jovanovic´, Olga [email protected] Semlalia Biologie, Residence Ajiad 3, Avenue Biology, Hellbrunnerstraße 34, 5020 Salzburg, Josip Juraj Strossmayer University, Department of Yaakoub El Mansour, Gueliz, 40000 Marrakech, Austria; [email protected] Biology, Cara Hadrijana 8/a, 31000 Osijek, Kopena, Renata Morocco; [email protected] Croatia; [email protected] Museo Nacional de Ciencias Naturales, CSIC Manenti, Raoul Ecologia Evolutiva, José Gutiérrez Abascal 2, Leeb, Christoph Univeristá degli Studi di Milano, Karameta, Emmanouela 28006 Madrid, Spain; [email protected] University of Vienna, Department of Integrative Dipartimento di Bioscienze, Via Celoria 26, Interno University of Athens, Biology, University Campus, Zoology, Althanstraße 14, 1090 Wien, Austria; 7b, 20133 Milano, Italy; [email protected] Ilisia, 15701 Athens, Greece; [email protected] Kovács, Tibor [email protected] MME-BirdLife Hungary, Amphibian and Reptile Mangold, Alexandra Katona, Patrik Conservation Group, Költo˝ u. 21, 1121 Budapest, Lisicic´, Duje University of Vienna, Department of Integrative University of Debrecen, Department of Evolutionary Hungary; [email protected] University of Applied Health Studies, Zoology, Althanstraße 14, 1090 Wien, Austria; Zoology, Egyetem tér 1, 4032 Debrecen, Hungary; Department of Biology, Mlinarska cesta 38, [email protected] [email protected] Kovács, Dániel 10000 Zagreb, Croatia; [email protected] Szent István University, Department of Zoology and Marshall, Jonathon Kaufmann, Peter Animal Ecology, Páter Károly utca 1, 2100 Gödöllo˝, Lopez, Pilar Weber State University, Zoology, 2505 University University of Salzburg, Department of Organismic Hungary; [email protected] Museo Nacional de Ciencias Naturales, CSIC Circle, 84408 Ogden, USA; [email protected] Biology, Hellbrunnerstraße 34, 5020 Salzburg, Ecologia Evolutiva, José Gutiérrez Abascal 2, Austria; [email protected] 28006 Madrid, Spain; [email protected]

320 321 Martel, An Mester, Béla Nagy, Zoltán Tamás Poboljsˇaj, Katja Ghent University, Faculty of Veterinary Medicine, University of Debrecen, Department of Ecology, Royal Belgian Institute of Natural Sciences, Centre for Cartography of Fauna and Flora, Department of Pathology, Bacteriology and Avian Egyetem tér 1, 4032 Debrecen, Hungary; JEMU, Rue Vautier 29, 1000 Brussels, Belgium; Ljubljana office, Klunova 3, 1000 Ljubljana, Diseases, Salisburylaan 133, 9820 Merelbeke, [email protected] [email protected] Slovenia; [email protected] Belgium; [email protected] Mészáros, Boglárka Najafi-Majd, Elnaz Popgeorgiev, Georgi Martin, José Eötvös Loránd University, Department of Systematic Ege University, Faculty of Science, Biology Section, Regional Natural History Museum, Vertebrates, Museo Nacional de Ciencias Naturales, CSIC Zoology and Ecology, Pázmány Péter sétány 1/c, Zoology Department, Ege Üniversitesi Kampüsü Hristo G. Danov St. 34, 4000 Plovdiv, Bulgaria; Ecologia Evolutiva, José Gutiérrez Abascal 2, 1117 Budapest, Hungary; [email protected] 35040, 35100 Bornova-Izmir, Turkey; [email protected] 28006 Madrid, Spain; [email protected] [email protected] Miaud, Claude Popovic´, Nina Martínez-Freiría, Fernando Ecole Pratique des Hautes Etudes, UMR CEFE Ogoanah, Sylvia Karlovac University of Applied Sciences, CIBIO, Centro de Investigação em Biodiversidade e Montpellier, Biogeography and Vertebrate Ecology, University of Benin, Department of Animal and Gamekeeping and Environmental Protection, Recursos Genetico Vairão, Campus Agrário de Vairão, 1919 route de Mende, 34293 Le Bourget-du-Lac, Environmental Biology, 30001 Benin, Nigeria; Trg J.J. Strossmayera 9, 47000 Karlovac, Croatia; Rua Padre Armendo Quintas, 4485-661 Vairão, France; [email protected] [email protected] [email protected] Portugal; [email protected] Mikó, Zsanett Ortega, Jesus Poulakakis, Nikos Matos, Catia Plant Protection Institute for Agricultural Research, Museo Nacional de Ciencias Naturales, CSIC University of Crete, Department of Biology and CICGE, Research Center for Geo-Spatial Sciences Hungarian Academy of Science, Ecologia Evolutiva, José Gutiérrez Abascal 2, Natural History Museum of Crete, Knossos Av., University of Porto, Vila Nova de Gaia, “Lendület” Evolutionary Ecology Research Group, 28006 Madrid, Spain; [email protected] P.O.Box 2208, 71409 Irakleio, Greece; 4430-146 Porto, Portugal; [email protected] Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] [email protected] Ortiz Rocha da Costa, Carolina Mayer, Michael ESALQ-USP, Ciências Biológica, Av. Pádua Dias 11, Psonis, Nikolaos Trier University, Department of Biogeography, Mikulícˇek, Peter Caixa postal 9, 13418900 Piracicaba, Brazil; University of Crete, Department of Biology and Universitätsring 15, 54296 Trier, Germany; Comenius University, Faculty of Natural Sciences, [email protected] Natural History Museum of Crete, Knossos Av., [email protected] Department of Zoology, Mlynska dolina B-1, P.O.Box 2208, 71409 Irakleio, Greece; 84215 Bratislava, Slovakia; [email protected] Pasmans, Frank [email protected] Mebert, Konrad Ghent University, Faculty of Veterinary Medicine, Siebeneichenstraße 31, 5634 Merenschwand, Mizsei, Edvárd Department of Pathology, Bacteriology and Avian Pujol-Buxó, Eudald Switzerland; [email protected] University of Debrecen, Department of Evolutionary Diseases, Salisburylaan 133, 9820 Merelbeke, Universitat de Barcelona, Departament de Biologia Zoology, Egyetem tér 1, 4032 Debrecen, Hungary; Belgium; [email protected] Animal, Facultat de Biologia, Av. Diagonal 643, Mechura, Tímea [email protected] 08028 Barcelona, Spain; [email protected] Eötvös Loránd University, Department of Systematic Pasˇukonis, Andrius Zoology and Ecology, Pázmány Péter sétány 1/c, Molnár, Tamás University of Vienna, Department of Cognitive Puky, Miklós 1117 Budapest, Hungary; [email protected] Kaposvár University, Department of Nature Biology, Mariahilferstraße 150/9, 1150 Wien, Austria; MTA Centre for Ecological Research, Danube Protection, Guba Sándor út 40, 7400 Kaposvár; [email protected] Research Institute, Jávorka S. u. 14, 2131 Göd, Meikl, Magdalena [email protected] Hungary; [email protected] University of Salzburg, Department of Organismic Pásztor, Katalin Biology, Hellbrunnerstraße 34, 5020 Salzburg, Molnár, Orsolya Plant Protection Institute for Agricultural Research, Rannap, Riinu Austria; [email protected] Dartmouth College, Biological Sciences, Hungarian Academy of Science, University of Tartu, Institute of Ecology and Earth 78 College Street, 03755 Hannover, USA; “Lendület” Evolutionary Ecology Research Group, Sciences, Vanemuise 46, 51014 Tartu, Estonia; Meilink, Willem [email protected] Herman Ottó út 15, 1022 Budapest, Hungary; [email protected] VUB & Naturalis, Pleinlaan 2, 1080 Brussel, [email protected] Belgium; [email protected] Mozaffari, Omid Rato, Catarina Aria Herpetological Institute, No. 1 Parastoo 4 St., Petzold, Alice CIBIO, Centro de Investigação em Biodiversidade e Mermod, Claude Shahrak-e-Homa, Ashraf Esfehani Highway, 14647 Senckenberg Natural History Collections Dresden, Recursos Genetico Vairão, Campus Agrário de Vairão, Institute of Biology, Rue du Jura 15d, 1422 Grandson, Tehran, Iran; [email protected] Department of Museum of Zoology, Rua Padre Armendo Quintas, 4485-661 Vairão, Switzerland; [email protected] Königsbrücker Landstraße 159, 01109 Dresden, Portugal; [email protected] Germany; [email protected]

322 323 Rausch, Anna Rosioru, Daniela Mariana Samoila, Ciprian Sillero, Neftali University of Vienna, Department of Integrative National Institute for Marine Research and Ovidius University of Constant¸a, Faculty of CICGE, Research Center for Geo-Spatial Sciences Zoology, Althanstraße 14, 1090 Wien, Austria; Development, Ecology and Environment Protection, Natural Sciences, Aleea Universita˘¸iit nr. 1, Corp B, University of Porto, Vila Nova de Gaia, [email protected] Mamaia Bd. 300, 900581 Constanta, Romania; dro- 900470 Constant¸a, Romania; 4430-146 Porto, Portugal; [email protected] [email protected] [email protected] Rautenberg, Julia Silva-Rocha, Iolanda Trier University, Department of Biogeography, Roussos, Stephen Sant’Anna, Sávio Stefanini CIBIO, Centro de Investigação em Biodiversidade e Universitätsring 15, 54296 Trier, Germany; Department of Biological Sciences, Texas Tech University of São Paulo, School of Veterinary Recursos Genetico Vairão, Campus Agrário de Vairão, [email protected] University, MS 43131, Lubbock, TX 79409, USA; Medicine and Animal Sciences, Av. Vital Brasil 1500, Rua Padre Armendo Quintas, 4485-661 Vairão, [email protected] 05503-900 São Paulo, Brazil; [email protected] Portugal; [email protected] Reinhardt, Timm Helmholtz Centre for Environmental Research Rovelli, Valentina Scali, Stefano Siroky´, Pavel GmbH, UFZ River Ecology, Brueckstraße 3a, 39114 Roma Tre University, Department of Science, Museo di Storia Naturale di Milano, Herpetology, University of Veterinary and Pharmaceutical Magdeburg, Germany; [email protected] Viale Guglielmo Marconi 446, 00146 Roma, Italy; Corso Venezia 55, 20121 Milano, Italy; Sciences Brno, Faculty of Veterinary Hygiene and [email protected] [email protected] Ecology, Department of Biology and Wildlife Reniers, Jane Diseases, Palackého tr. 1/3, 61242 Brno, KU Leuven, Biology Department, Laboratory of Rozínek, Roman Schmitz, Andreas Czech Republic; [email protected] Aquatic Ecology, Evolution and Conservation, NaturaServis Ltd., Ricˇarˇova 66, Natural History Museum Geneva, Department of Ch. Deberiotstraat 32, 3000 Leuven, Belgium; 50301 Hradec Králové, Czech Republic; Herpetology, Route de Malagnou 1, 1208 Geneva, Sopotnik, Maja [email protected] [email protected] Switzerland; [email protected] Societas Herpetologica Slovenica, Presernova 20, 1000 Ljubljana, Slovenia; [email protected] Ringler, Max Ruben, Iosif Schulte, Ulrich University of Vienna, Department of Integrative Ovidius University of Constant¸a, Faculty of Trier University, Department of Biogeography, Sos, Tibor Zoology, Althanstraße 14, 1090 Wien, Austria; Natural Sciences, Aleea Universita˘t¸ii nr. 1, Corp B, Universitätsring 15, 54296 Trier, Germany; Milvus Group, Bird and Nature Protection [email protected] 900470 Constant¸a, Romania; [email protected] Association, Márton Áron St. 9/b, 540620 Targu- [email protected] Mures, Romania; [email protected] Ringler, Eva Schulte, Lisa Maria University of Vienna, Department of Integrative Rujirawan, Attapol Trier University, Department of Biogeography, Sparreboom, Max Zoology, Althanstraße 14, 1090 Wien, Austria; Kasetsart University, Department of Zoology, Universitätsring 15, 54296 Trier, Germany; Naturalis Biodiversity Center, [email protected] Faculty of Science, P.O. Box 1020, 10900 Bangkok, [email protected] Reijnier Vinkeleskade 38 II, 1071SV Amsterdam, Thailand; [email protected] The Netherlands; [email protected] Rödder, Dennis Secondi, Jean Zoologisches Forschungsmuseum Alexander Koenig, Sacchi, Roberto University of Angers, GECCO, 40 rue de Rennes, Spitzen-van der Sluijs, Annemarieke Herpetology Section, Adenauerallee 160, 53113 Bonn, University of Pavia, Dipartimento di Scienze 49045 Angers, France; [email protected] RAVON (Reptile Amphibian Fish Conservation Germany; [email protected] della Terra e dell’Ambiente, Via Taramelli 24, Netherlands), P.O. Box 1413, 6501BK Nijmegen, 27100 Pavia, Italy; [email protected] Sfenthourakis, Spyros The Netherlands; [email protected] Roessler, Daniela University of Cyprus, Biological Sciences, Trier University, Department of Biogeography, Sagonas, Konstantinos University Campus, P.O. Box 20537, 1678 Nicosia, Spitzweg, Cäcilia Universitätsring 15, 54296 Trier, Germany; National and Kapodistrian University of Athens, Cyprus; [email protected] Senckenberg Natural History Collections Dresden, [email protected] Department of Biology, Panepistimioupoli Zografou, Department of Museum of Zoology, 15784 Athens, Greece; [email protected] Sheverdyukova, Hanna Königsbrücker Landstraße 159, 01109 Dresden, Romanazzi, Enrico Institute of Zoology NAS of Ukraine, Vertebrate Germany; [email protected] Science Museum of Trento, Zoologia dei Vertebrati Salleh, Sarahaizad Mohd Evolutionary Morphology, Vul. B. Khmelnytskogo 15, Corso del Lavoro e della Scienza 3, 38123 Trento, Universiti Sains Malaysia, School of Biological 01601 Kyiv, Ukraine; [email protected] Stanescu, Florina Italy; [email protected] Sciences, Sungai Dua, 11050 Penang, Ovidius University of Constant¸a, Faculty of Malaysia; [email protected] Shine, Rick Natural Sciences, Aleea Universita˘¸iit nr. 1, Corp B, Rosa, Gonçalo M. University of Sydney, Biological Sciences, 900470 Constant¸a, Romania; Durrell Institute of Conservation and Ecology, Salvidio, Sebastiano Heydon-Laurene Building A08, Sydney, NSW [email protected] University of Kent, Canterbury CT2 7NR, Università di Genova, DISTAV, C. so Europa 26, 2006, Australia; [email protected] United Kingdom; [email protected] 16132 Genova, Italy; [email protected]

324 325 Stankovic´, David Takacs, Zoltan Üveges, Bálint de Vries, Wouter University of Ljubljana, Biotechnical Faculty, World Toxin Bank Initiative, ToxinTech, Inc., Plant Protection Institute for Agricultural Research, Asociación AMBOR, P.O. Box 30, Department of Animal Science, Domzale Groblje 3, P. O. Box 6266, New York, NY 10022, USA; Hungarian Academy of Science, 41450 Constantina, Spain; [email protected] 1230 Ljubljana, Slovenia; [email protected] [email protected] “Lendület” Evolutionary Ecology Research Group, Herman Ottó út 15, 1022 Budapest, Hungary; Wagner, Norman Starnberger, Iris Tóth, Zoltán [email protected] Trier University, Department of Biogeography, University of Vienna, Department of Integrative Plant Protection Institute for Agricultural Research, Universitätsring 15, 54296 Trier, Germany; Zoology, Althanstraße 14, 1090 Wien, Austria; Hungarian Academy of Science, Üzüm, Nazan [email protected] [email protected] “Lendület” Evolutionary Ecology Research Group, University of Adnan Menderes, Faculty of Science Herman Ottó út 15, 1022 Budapest, Hungary; and Art, Biology Department, Kepez, 09010 Aydin, Wagner, Philipp Steinfartz, Sebastian [email protected] Turkey; [email protected] Zoologisches Forschungsmuseum Alexander Koenig, University of Bielefeld, Department of Animal Herpetology Section, Adenauerallee 160, 53113 Bonn, Behaviour, Morgenbreede 45, 33615 Bielefeld, Toxopeus, Bert Vági, Balázs Germany; [email protected] Germany; [email protected] ITC, University of Twente, Department of Natural Plant Protection Institute for Agricultural Research, Resources, Hengelosestraat 99, 7514AE Enschede, Hungarian Academy of Science, Wahlgren, Richard Stoev, Pavel The Netherlands; [email protected] “Lendület” Evolutionary Ecology Research Group, International Society for the History and Pensoft Publishers Ltd., National Museum of Herman Ottó út 15, 1022 Budapest, Hungary; Bibliography of Herpetology, Prennegatan 23b, Natural History, Prof. G. Zlatarski St. 12, Trenkwalder, Katharina [email protected] 22353 Lund, Sweden; [email protected] 1700 Sofia, Bulgaria; [email protected] University of Vienna, Department of Integrative Zoology, Althanstraße 14, 1090 Wien, Austria; Valdeón, Aitor Wangkulangkul, Sansareeya Szabó, Krisztián [email protected] Aranzadi Society of Science, Herpetology, Prince of Songkla University, Faculty of Science, Szent István University, Institute of Biology, Alto de Zorroaga 11, 20014 Donostia–San Sebastián, Department of Biology, Hat Yai, 90112 Songkhla, Department of Ecology, Rottenbiller u. 50, Turniak, Edyna Spain; [email protected] Thailand; [email protected] 1077 Budapest, Hungary; [email protected] University of Wroclaw, Laboratory of Vertebrate Evolutionary Biology & Conservation, Vamberger, Melita Weiperth, András Szabolcs, Márton Sienkiewicza 21, 50335 Wroclaw, Poland; Senckenberg Natural History Collections Dresden, MTA Centre for Ecological Research, Danube Nature Conservation Association of Tokaj, [email protected] Department of Museum of Zoology, Research Institute, Jávorka S. u. 14, 2131 Göd, Liget köz 1, 3910 Tokaj, Hungary; Königsbrücker Landstraße 159, 01109 Dresden, Hungary; [email protected] [email protected] Ujszegi, János Germany; [email protected] Plant Protection Institute for Agricultural Research, Werner, Philine Székely, József-Paul Hungarian Academy of Science, Vélez Olalde, Yaiza Trier University, Department of Biogeography, Ovidius University of Constant¸a, Faculty of “Lendület” Evolutionary Ecology Research Group, EUIT Forestal y del Medio Natural (Universidad Universitätsring 15, 54296 Trier, Germany; Natural Sciences, Aleea Universita˘t¸ii nr. 1, Corp B, Herman Ottó út 15, 1022 Budapest, Hungary; Politécnica de Madrid), Zoology, c/Juan Ramón [email protected] 900470 Constant¸a, Romania; [email protected] [email protected] Jiménez 18 1 A, 28100 Alcobendas (Madrid), Spain; [email protected] Wielstra, Ben Székely, Diana Unglaub, Bianca Naturalis Biodiversity Center, Terrestrial Zoology, Ovidius University of Constant¸a, Faculty of University of Bielefeld, Department of Animal Verkovich, Olga Darwinweg 2, 2300RA Leiden, The Netherlands; Natural Sciences, Aleea Universita˘t¸ii nr. 1, Corp B, Behaviour, Morgenbreede 45, 33615 Bielefeld, Zoological Park, Aleksandrovskiy park 1, [email protected] 900470 Constant¸a, Romania; [email protected] Germany; [email protected] 197198 Saint-Petersburg, Russia; [email protected] Wilkinson, John Szymura, Jacek M. Ursenbacher, Sylvain Amphibian and Reptile Conservation, Jagiellonian University, Comparative Anatomy, University of Basel, Department of Environmental Virkki, Diana 655a Christchurch Road, Bournemouth BH1 4AP, Gronostajowa 9, 30-387 Kraków, Poland; Sciences, St. Johanns-Vorstadt 10, 4056 Basel, Griffith University, Environmental Future Centre, United Kingdom; [email protected] [email protected] Switzerland; [email protected] School of Environment, Parklands Drive G24, 4222 Southport, Australia; [email protected] Winandy, Laurane Takács, Gábor Urszán, Tamás János University of Liège, Behavioural Biology Unit, Ferto˝-Hanság National Park, Nature Conservation Eötvös Loránd University, Department of Systematic Vörös, Judit 22 Quai van Beneden, 4020 Liège, Belgium; Department, Rév-Kócsagvár, 9435 Sarród, Hungary; Zoology and Ecology, Pázmány Péter sétány 1/c, Hungarian Natural History Museum, [email protected] [email protected] 1117 Budapest, Hungary; [email protected] Department of Zoology, Herpetological Collection, Baross u. 13, 1088 Budapest, Hungary; [email protected]

326 327 Winkler, Martina Zamfirescu, Stefan Paris Lodron University of Salzburg,Organismic Alexandru Ioan Cuza University of Ias¸i, Biology, Oberdorfer Straße 13, Faculty of Biology, St. Carol I, nr. 20a, Author Index 5303 Thalgau, Austria; [email protected] 700505 Ias¸i, Romania; [email protected]

Wolf, Silas Zinenko, Oleksandr Trier University, Department of Biogeography, The Museum of Nature at V.N. Karazin, Universitätsring 15, 54296 Trier, Germany; Kharkiv National University, Trinkler St. 8, [email protected] 61058 Kharkiv, Ukraine; [email protected] Abd Mutalib, A.H. 84 Berninger, U. G. 268 Channing, A. 203 Dikic, D. 137 Yildirim, Elif Znari, Mohammed Ács, É. 306 Bert, W. 59 Chiers, K. 59 Dittrich, C. 217 Ege University, Faculty of Science, Biology Section, University of Cadi Ayyad, Faculty of Sciences, Adriaensen, C. 61 Bertoluci, J. 220, 277 Choleva, L. 181 Djordjevic´, S. 115 Zoology Department, Ege Üniversitesi Kampüsü Semlalia Biologie, Residence Ajiad 3, Avenue Adriano, L.R. 220 Beukema, W. 61, 95 Cipot, M. 78, 294 Djurakic´, M. 214 35040, 35100 Bornova-Izmir, Turkey; Yaakoub El Mansour, Gueliz, 40000 Marrakech, Ahmadzadeh, F. 85, 86, Biaggini, M. 100 Civiˇ,s P. 56 Djuric´, D. 215 [email protected] Morocco; [email protected] 164, 200, 201 Biancardi, C. M. 205 Clare, F. 57 Dobay, G. 216 Al-Jaidah, N. 210 Bíró, P. 306 Clemann, N. 120 Doglio, S. 268 Alkuwary, A. 210 Bitenc, K. 195, 206 Cmiel, A.M. 134 Drakulic´, S. 137, 217 Yodthong, Siriporn Zollinger, Ronald Alpagut-Kerskin, N. 181 Blooi, M. 57, 59, 61 Cogalniceanu, D. 75, 210, Drechsler, A. 183 Kasetsart University, Department of Zoology, RAVON (Reptile Amphibian Fish Conservation Ananjeva, N. 87, 197 Bogaerts, S. 61, 95 242, 285, 293, 296, 297 Du Preez, L. 279 Faculty of Science, P.O. Box 1020, 10900 Bangkok, Netherlands), P.O. Box 1413, 6501BK Nijmegen, Andjelkovic´, M. 115 Bogicˇevic´, K. 215 Conradie, W. 286 Duarte, A. 62 Thailand; [email protected] The Netherlands; [email protected] Angulo, A. 109 Bologna, M. A. 167 Cornette, R. 214 Duarte, R. 98 Angulo, I. 265, 266 Bonardi, A. 109, 174 Cortada, A. 99 Ducatelle, R. 59 Zagar, Anamarija Zuffi, Marco Antoniou, A. 304 Bonato, L. 141, 165 Corti, C. 100, 131, 174 Dufresnes, Ch. 141 CIBIO, Centro de Investigação em Biodiversidade e University of Pisa, Museum of Natural History, Aowphol, A. 88, 168, 194 Bonnet, X. 115 Costa, B. M. 288 Dvoráková, N. 176 Recursos Genetico Vairão, Campus Agrário de Vairão, Via Roma 79, 56011 Calci (Pisa), Italy; Argaña, E. 98, 267 Borczyk, B. 207 Cotrel, N. 50 Dzˇukic´, G. 243 Rua Padre Armendo Quintas, 4485-661 Vairão, [email protected] Arnell, A.P. 45, 81 Bosch, J. 62, 110 Creemers, R. 66, 101, 174 Eckerlein, A. 218 Portugal; [email protected] Arntzen, J. W. 34, 46, 89, Bosman, W. 58, 59, 122 Crnobrnja-Isailovic´, J. Eikelmann, D. 119 192, 202, 255 Bossuyt, F. 59 102, 157, 174, 211 El Mouden, E. H. 257 Arsovski, D. 115 Bovero, S. 61 Croce, A. P. 288 Engler, J. O. 86, 126, 164, Astrin, J. 284 Böhm, S. 42 Crochet, P.A. 174 201 Audebaud, B. 50 Böhme, W. 35, 85, 86, 118, Crottini, A. 129 Erich, M. 162, 219 Ayres, C. 40, 303 131, 200, 201, 234, 284 Czarnoleski, M. 213 Ernst, R. 238 Babiasz, R. 134 Böning, Ph. 193 Cservenka, J. 305 Fabre, A.-C. 214 Babik, W. 182 Branch, W.R. 279 Csillag, Á. 186 Famelli, Sh. 220 Babocsay, G. 41, 90 Braun, M. 290 Damas Moreira, I. 103 Familiar Lopez, M. 107 Bajer, K. 91, 92, 151, 271 Brendonck, L. 282 Dambach, J. 126, 164 Feldhaar, H. 217 Bakó, B. 74 Brito, J. C. 144 Daniels, S. R. 106 Fera, G. 221 Balázˇ, V. 49, 56, Broeckhoven, Ch. 96 Dankovics, R. 53 Fernandes, R. V. 288 Balogh, S. B. 248 Brooks, D. R. 236 Davoli, F. 167 Fernandes, W. 171, 288 Ba˘ncila˘, R. 202, 285, 297 Buckley, D. 225 De Pous, Ph. 129, 174 Fernández Cardenete, J. R. Baptista, N. 203 Bulakhova, N. 208 de Vries, W. 80 268 Barej, M. F. 178 Burger, M. 286 Deák, G. 212 Ferraz, K. M. P. 277 Barquero, M. 93 Bury, S. 134 Degani, G. 104 Ferreira, M. 108 Barta, Z. 236 Bülbül, U. 181 Dehling, M. 286 Ficetola, G. F. 44, 105, Bas, G. 134 Cabido, C. 209, 226, 240 Dejean, T. 44, 50, 122 109, 142, 174 Bassu, L. 100 Campos, J. 174 Denoël, M. 105, 142, 174, Filz, K. 222 Bates, P. 312 Canessa, S. 97 176, 296, 313 Fischer, D. 49 Beja, P. 108 Carretero, M. A. 51, 85, Densmore III, L. D. 166 Fisher, M. C. 57, 59, 61, 110 Bell, R. 286 86, 98, 99, 103, 159, 195, Derus, N. 213 Fleck, J. 284 Bellakhal, M. 94, 204 200, 201, 206, 224 Desgranges, S. 50 Flecks, M. 85, 86, 126, Benavides Sánchez de Carvalho, M. 171 Di Cerbo, A. R. 100, 205 164, 201, 284 Molina, J. 268 Carvalho, S. B. 159 Di Francesco, N. 100 Fonseca-Pinto, A. C. 171 Benedek, I. 273 Castilla, A. M. 210 Di Piazza, S. 228 Foster, J. 45, 81 Benkovic, V. 137 Castley, G. 188 Di Tizio, L. 100 Freitas, S. N. 85, 200 Berman, D. I. 208 Catão-Dias, J. L. 171 Diedericks, G. 106 Fresi, C. 100

328 329 Fritz, U. 131, 177, 187, 279 Harris, D. J. 85, 86, 103, Kaya, U. 152, 314 239 Mesquita, B. 224 Pálinkás, I. 273 Represas, A. G. 268 Schulte, L. M. 87 Fugiwara, C. Y. 288 159, 200, 201 Kéri, A. 247 Lorenzo, M. 145, 259, 260 Mester, B. 70, 147, 247, Parmakelis, A. 287 Richer, R. 210 Schulz, S. 178 Gabor, C. R. 110 Hartel, T. 75 Kielgast, J. 77, 286 Lorigados, C. 171 270 Pasmans, F. 57, 58, 59, 61, Rijks, J. M. 63 Scillitani, G. 100 Gaebele, T. 190 Hartmann, T. 118, 126, Kieren, S. 129 Lovász, Zs. 261 Mészáros, B. 271 63, 95, 97 Ringler, E. 136, 160, 162, Scroggie, M. P. 120 Gál, János 273 164, 234, 256, 284 Kik, M. 63 Loyau, A. 57 Metallinou, M. 131 Pastorino, M. V. 169 163, 219, 264, 298 Secondi, J. 50 Gál, Júlia Tünde 111 Hastrup, M. 77 Kindermann, Ch. 130 Lötters, S. 146, 193, 218, Meyer, L. 279 Pasˇukonis, A. 155, 162 Ringler, M. 136, 162, 163, Seiler, M. 217 Gál, Z. 148, 223, 301 Hauswaldt, S. 119 Kindler, C. 131 222, 281, 286, 308 Miaud, Claude 44, 50, 60, Pásztor, K. 133, 221, 278 219, 264, 298 Seligmann, H. 90 Galoyan, E. A. 112 Havlíková, B. 56 Kirbisˇ, N. 291 Ludwig, G. 139 285 Péchy, T. 117 Roberts, D. M. 156 Seng, R. 234 Garcia-Diaz, P. 145, 260 Heard, G. W. 120 Kiss, I. 212, 216, 235, 248, Luzˇnik, M. 262 Mihaljevic, Z. 137 Pellinger, A. 53 Rocha, A. 283 Sfenthourakis, S. 244, 258 García-Muñoz, E. 99, 224 Hegyeli, Zs. 292 251, 261 Lymberakis, P. 116, 157, Mikó, Zs. 148, 223, 301 Pelster, B. 139 Rodríguez, A. 174 Sheverdyukova, H. 289 García-Parra, C. 140 Heltai, B. 235 Knudsen, J. 178 174, 244, 287, 304 Mikulícˇek, P. 47, 243, 246 Penev, L. D. 156 Rodríguez de Rivera, O. Shine, R. 36 García-Roa, R. 225 Herczeg, D. 121, 236 Koch, G. 50 Macale, D. 167 Milcic, E. 137 Penn, D. 237 265, 266 Shiryaev, K. 197 Garin-Barrio, I. 226 Herczeg, G. 91, 92, 151, Koch, V. 140 Macˇát, Z. 263 Milojkovic´, D. 211 Pereira, F.P.A. 220 Romanazzi, E. 165, 268 Sillero, N. 174, 267 Garner, T.W.J. 56, 57, 61 250, 271, 302 Kolanek, A. 299 MacLeod, A. 140 Milosevic´, D. 102 Pereira Almeida, C. 48 Romano, A. 100 Silva, J. L. F. 288 Garriga, N. 48, 227 Herder, J. 122 Kopena, R. 132, 250 Macura, B. 211 Mizsei, E. 149, 150, 290 Perera, A. 103, 159, 200 Rondinini, C. 109 Silva-Rocha, I. R. 51 Geiser, C. 113 Hero, J.-M. 107, 123, 130 Kovács, D. 251 Magoulas, A. 304 Mohd Sah, S. A. 170, 272 Perry, S.F. 135 Rosa, G. M. 62 Simon, E. 290 Geissler, P. 118, 234, 256 Herrel, A. 137, 214 Kovács, T. 186, 245, 252, Mahony, M.J. 67 Mohd Salleh, S. 170, 272 Peters, R. 93 Rosioru, D. 285 Sindaco, R. 174 Gémes, D. 70 Hesselsøe, M. 77 261, 305 Mahr, K. 184 Molnár, M. 273 Petrov, B. 157 Roussos, S. A. 166 Sinsch, U. 139 Gent, T. 45, 81 Hettyey, A. 148, 184, 186, Krasa, A. 253 Mahroo, B. S. 241 Molnár, O.R. 91, 92, 151, Petrovan, S. 68, 69 Rovelli, V. 167 Siroky´, P. 131, 176 Georges, A. 220 221, 223, 237, 301, 302 Król, W. 134 Maia, J. P.M. C. 103 271 Petrovicˇ, I. 291 Rozínek, R. 49, 56 Skidmore, A. K. 116 Georgiev, T.A. 156 Hidaka, M. S. 288 Krysˇtufek, B. 262 Maletzky, A. 141, 246 Molnár, T. 273 Petzold, A. 279 Rozman, A. 291 Slimani, T. 257 Ghaffari, H. 241 Hofman, S. 181 Krzys´ciak-Kosin´ska, R. 254 Manenti, R. 142 Moravec, J. 243 Philipp, D. 114 Rozman, L. 291 Smith, V.S. 156 Gherghel, I. 157, 196 Hofmeyr, M. D. 279 Kukushkin, O. V. 157 Mangiacotti, M. 172 Moterani, R. 288 Pitt, A. 268 Rozylowicz, L. 75 Solarz, W. 52 Ghira, I. 292 Hoi, H. 184, 186 Kundrát, J. T. 290 Mangold, A. 264, 298 Mozaffari, O. 85 Plaiasu, R. 285 Rödder, D. 85, 86, 118, Sopotnik, M. 291 Giannoccari, Ch. A. 172 Horváth, G. 92 Kurali, A. 133, 278 Mantilla, M. X. G. 233 Murphy, R. 197 Plummer, M. V. 241 126, 159, 164, 200, 201, Sos, T. 292 Gollmann, G. 114, 136, 255 Hödl, W. 124, 155, 160, Kurek, K. 52, 134 Marco, A. 80, Müller, S. 286 Poboljsˇaj, K. 78, 291 234, 241, 284 Sós, E. 117 Golubovic´, A. 115 162, 163, 178, 219, 264, Kusamba, Ch. 274 Mariaux, J. 116 N Rosely, N.F.N. 84 Polly, P.D. 164 Rödel, M.-O. 178, 217 Sotgiu, G. 61 Gomes, V. 103 298 Kutuzovic´ Hackenberger, B. Markus, M. 158 Nagy, Z.T. 274, 286 Pompanon, F. 44 Rössler, D. 286 Sousa, F. 174 Gonçalves, J. 174 Hölting, M. 238 128 Marosán, M. 273 Naimi, M. 153, 275 Popgeorgiev, G. S. 300 Rujirawan, A. 88, 168 Sousa, P. 174 Gonçalves-Seco, L. 174 Hörnes, D. 164 Kuzmin, S. 174 Márquez, R. 225, 293 Najafi-Majd, E. 152, 314 Popovic´, N. 280 Rulík, M. 263 Souza, F. L. 220 Gorynia, M. 207 Ibáñez, A. 125, 239 Kvicerová, J. 176 Marshall, J. C. 143 Najbar, B. 134 Poth, D. 178 Sacchi, R. 172 Sparreboom, M. 95 Gosá, A. 210 Iglesias, R. 303 Lagler, P. 255 Martel, A. 57, 58, 59, 61, Najberek, K. 134 Potoczek, M. 134 Sagonas, K. 258, 287 Spatz, T. 217 Govedic´, M. 78 Iglesias-Carrasco, M. 240 Lakatos, F. 306 63, 97 Namous, S. 153, 275 Potyó, I. 190 Sahlean, T. C. 196 Spitzen-van der Sluijs, A. Goverse, E. 76, 80 Ihlow, F. 118, 126, 164, 234, Lambertz, M. 135, 256 Martín, I. 265, 266 Narayan, E.J. 123, 130 Poulakakis, N. 157, 244, Saifelnasr, E. O. 210 58, 59, 63 Grasselli, E. 228 241, 284 Lammers, J. W. 43 Martín, J. 125, 132, 138, Nasir, N. 84 287, 304 Salvi, D. 51, 103 Spitzweg, C. 177 Greenbaum, E. 286 Ikovic´, V. 115 Langton, T. 67 154, 209, 239, 250, Nawal, H. 153, 275 Poyarkov, N. A. 112, 118 Salvidio, S. 61, 169, 228 Spreyboeck, J. 174 Grego, K. F. 171, 288 Ilgaz, Ç. 86, 201, 244 Lansari, A. 257 Martínez-Freiría, F. 144, Nekum, S. 126 Psonis, N. 157 Sály, P. 212, 235, 261 Sta˘nescu, F. 293 Gregorovicˇová, M. 229 Ilic´, M. 211 Lawing, A. M. 164 145, 259, 260, 283 Newell, D. 107 Pujol-Buxó, E. 48, 227 Samoila, C. 242, 252 Stankovic´, D. 291, 294 Griffiths, R. 62 Iosif, R. 242 Lázaro, R. 303 Martínez-Saura, C. 226 Nidup, Tsh. 312 Puky, M. 70, 71, 147, 190, San Sebastián, O. 48 Starnberger, I. 178 Griggio, M. 186 Ivanovic, A. 202 Leeb, Ch. 136 Martínez-Solano, I. 89 Nilson, G. 197 247, 270, 306 Sancho, V. 303 Steinfartz, S. 119, 140, 183 Grillitsch, B. 230 Iversen, L. L. 77 Lehmann, A. 113 Marzal, A. 239 Novarini, N. 141 Rada, V. 303 Sandera, M. 56, 181 Stiels, D. 164 Grillitsch, H. 230, 244 Jablonski, D. 131, 243 Lengyel, Sz. 74, 150, 270, Mas, P. 210 Nulchis, V. 100 Rajaei, H. 126 Sannolo, M. 172 Stoev, P.E. 156 Groen, T. 116 Jagar, T. 291 295 Mastropasqua, F. 100 Ogoanah, S.O. 276 Ramsey, D.S.L. 120 Sant’Anna, S. 171, 288 Stojadinovic´, D. 102 Grone, A. 63 Jáger, E. 248 Lesˇnik, A. 78 Masucci, G. D. 198 Ohler, A. 89 Randi, E. 167 Santos, X. 283 Stöck, M. 141 Gross, I. 232 Jandzˇik, D. 131, 243 Lewis, T. 188 Matos, C. 267 Oldenburg, S. 77 Rannap, R. 158 Sá-Rocha, L. C. 171 Strugariu, A. 196 Grozdanov, A. P. 300 Jehle, R. 160 Lisicic, D. 137 Mayer, M. 146 Oliveira, M. A. 174 Rato, C. 159, 224 Satasook, Ch. 312 Stuckas, H. 187 Gruber, B. 220 Jelic´, D. 111, 243 Litsi-Mizan, V. 258 Mazanaeva, L. 197 Oneto, F. 61, 169 Rausch, A. 160 Satta, M. G. 100 Stumpel, A. 179 Guti, G. 190 Joger, U. 127, 197 Lizana, M. 145, 259, 260 McAtear, J. 89 Orlov, N. 197 Rautenberg, J. 281 Scali, S. 172 Stümpel, N. 197 Gümü¸,s Ç. 185, 232 Jovanovic´, O. 128 Ljubisˇavljevic´, K. 243 McLaughlin, P.J. 286 Orlowska, M. 254 Ray, N. 113 Schedl, H. 114 Sylvain, D. 180 Gvozˇdík, V. 131, 233, 243, Kaart, T. 158 Llorente, G. A. 48, 227 McLeod, D. S. 88, 256 Ortega, J. 138, 154 Razzetti, E. 205 Schidelko, K. 164 Szabó, K. 307 274, 286 Kaczkó, D. 216 Llusia, D. 225 Mechura, T. 70, 247 Ortiz, C. 277 Rebelo, R. 62, 203 Schiesari, L.C. 230 Szabolcs, M. 150, 295 Gyulai, I. 290 Kaliontzopoulou, A. 103 Loader, S. 233 Meikl, M. 268 Ostanek, E. 291 Reckendorfer, W. 114 Schleicher, A. 279 Szalai, M. 270 Habel, J. Ch. 164 Karameta, E. 244, 258 Lollback, G. 107 Meilink, Willem 46 Ottonello, D. 61, 169 Recuero, E. 89 Schmeller, D. S. 57 Szederkényi, M. 184, 186 Halpern, B. 117 Katariya, V. 109 Loncaric´, Z. 128 Menegon, Michele 233 Padoa-Schioppa, E. 109 Reinhardt, T. 161 Schmidt, B. R. 79, 183 Székely, D. 296, 297 Hamer, A. 67 Katona, P. 150, 186, 245 Longares, L. A. 303 Mermod, C. 269 Pafilis, P. 244, 258, 287 Reiter, A. 263 Schmitz, A. 118 Székely, P. 293, 296, 297 Handschuh, M. 234 Kaufmann, P. 246 López, P. 125, 132, 138, 154, Meshcheryakova, E. N. 208 Pagano, A. 257 Reniers, J. 282 Schöttler, T. 284 Sztatecsny, M. 160

330 331 Szymura, J. M. 181 Uchedike, E. 276 Veith, M. 129, 218 Weiperth, A. 190 Taberlet, P. 44 Ujszegi, J. 148, 186, 223, 301 Velekei, B. 306 Weitere, M. 161 Tadic, Z. 137 Unglaub, B. 183 Vélez, Y. 265, 266 Werner, Ph. 191 Takacs, Z. 19 Ursenbacher, S. 79, 113 Velo-Antón, G. 144 Whiting, M. 93 Takács, G. 53 Urszán, T.J. 302 Vences, M. 119, 174, 178 Wielstra, B. 46, 192, 255 Tavares, L. 62 Üveges, B. 149, 184 Venegas, P.J. 193 Wierzbanowski, P. 134 Telbisz, T. 305 Üzüm, N. 86, 185, 201, 232 Viana, L. 283 Wild, C. H. 130 Tessa, G. 61 Vági, B. 41, 186, 245, 305 Viertel, B. 218 Wilkinson, J. W. 45, 81 Thomsen, Ph. F. 77 Valakos, E. D. 258, 287, 304 Vieites, D. R. 174 Winandy, L. 313 Thonhauser, K. 237 Valdeón, A. 210, 303 Vignoli, L. 167 Woeltjes, A. 59, 61 Tomé, B. 103 Valentini, A. 122 Virkki, D. 188 Wolf, S. 193 Tomovic´, L. 115 Vamberger, M. 177, 187, Vojar, J. 56 Yildirim, E. 152, 314 Tóth, J. P. 149 291 Vörös, J. 111, 121, 236, 248, Yobe, M. 170, 272 Tóth, M. 247 van Alphen, J. 89 307, 312 Yodthong, S. 194 Tóth, Z. 133, 182, 237, 278 Van Buskirk, J. 182, 237 Vrezec, A. 195, 206 Zagar, A. 195, 206, 291 Tóthmérész, B. 290 van Delft, J. 43, 46, 76, 101 Vujovic´, A. 125 Zamfirescu, O. 196 Toxopeus, A. G. 116 van den Broek, J. 63 Wagner, N. 308 Zamfirescu, S. R. 196 Toxopeus, B. 174 van der Ree, R. 67 Wagner, Ph. 189, 309, 310, Ziecik, A. 134 Török, J. 91, 92, 151, 271, van Diepenbeek, A. 66 311 Ziermann, J. M. 89 302 van Rooij, P. 61 Walsh, Sh. 256 Zinenko, O. 127, 197 Tran, C. 188 Vanschoenwinkel, B. 282 Walzer, Ch. 117 Znari, M. 153, 275 Trenkwalder, K. 264, 298 Vasconcellos, D.R. 288 Walzl, M. 178 Zollinger, R. 43, 76, 101, Trillmich, F. 140 Vasilikopoulos, A. 304 Wandycz, A. 181 179 Tuniyev, B. 197 Vassilieva, A. B. 112 Wangkulangkul, S. 312 Zotti, M. 228 Turniak, E. 299 Vecchione, G. 228 Waringer-Löschenkohl, A. Zuffi, M. 100, 198 Tzankov, N. D. 243, 300 Végvári, Zs. 236 114 Zsólyomi, T. 295

332