Workshop on Frugivores & Seed Dispersal FSD 2015

PROGRAMME & ABSTRACTS 6th International Symposium- Workshop on Frugivores & Seed Dispersal FSD 2015

Searching for commonality in frugivory and seed dispersal: merging community ecological dynamics with phylogeny across different biomes 2

6th International Symposium- Workshop on Frugivores & Seed Dispersal FSD 2015

Venue:

Alpine Heath, Northern Drakensberg, South Africa

Dates:

Sunday June 21 arrive, Depart midday Friday 26 June 2015

Organised and Hosted by:

School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg

Website:

http://www.fsd2015.ukzn.ac.za

Contents Chairperson Welcome to FSD 2015 ...... 7 Scientific Committee ...... 8 Local Organising Committee ...... 8 INSTRUCTIONS to FSD 2015 PARTICIPANTS ...... 9 Oral papers ...... 9 Poster presentations ...... 9 Student presentations ...... 9 FSD 2015 PROGRAMME ...... 9 FSD PLENARY ABSTRACTS ...... 17 Böhning-Gaese, K...... 17 Campos-Arceiz, A.1, Kromann-Clausen, A.1, Juwita, W.1, Mahmood, T.1, Solana Mena, A.1,2, Ning, H.1 & Ekanayaka, S.K.K.3 ...... 17 Effiom, E...... 18 Galetti, M...... 18 Jansen, P.A...... 18 Midgley, J...... 19 Nathan, R...... 19 Traveset, A...... 19 Tsuji, Y...... 20 Westcott, D.A...... 20 FSD ORAL ABSTRACTS ...... 21 Acosta Rojas, D.C., Cruz-Tejada, D.M., Páez-Crespo, E.J. & Stevenson, P.R...... 21 Albert-Daviaud, A.1,2, McConkey, K.3, Savini, T.4 & Huynen, M-C.1 ...... 21 Baltzinger, C.1, Albert-Daviaud, A.1,2, Picard, M.1 & Mårell, A.1...... 22 Bello, C*., Galetti, M., Magnago, L.F.S., Rocha, M., Lima, R., Peres, C.A. & Jordano, P...... 22 Bender, I.M.A.*1, Dehling, D.M.2, Hensen, I.3, Kissling, W.D.4, Kühn, I.5, Wiegand, T.6 & Schleuning, M.2 ...... 22 Blendinger, P.G1 & Giannini, N.P.2 ...... 23 Blendinger, P.G.1, Ordano, M.2, Lomáscolo, S.B.1, Chacoff, N.P.1, Núñez Montellano, M.G.3, Jiménez, J.1, Ruggera, R.A.*4, Valoy, M.2, Sánchez, M.S.5 & Martín, E.2 ...... 23 Bogdziewicz, M.*1,2, Steele, M.A.3, Crone, E.E.2 & Zwolak, R.1 ...... 24 Jana, A., Venkateswaran, V. & Borges, R.M...... 24 Bunney, K.1, Bond, W.1 & Henley, M.2 ...... 25 Burns, K.C...... 25 Campos, C.M., Miguel, M.F. & Cona, M.I...... 25 Chen, S-C.* ...... 26

Chibesa, M.*, Downs, C.T., Rouget, M., Taylor, B. & Ramesh, T...... 26 Camargo, P.H.S.A. 1, Martins, M.M. 2 & Christianini, A.V.3 ...... 27 Clausi, B...... 27 Clausi, B...... 27 Culot, L.1, Bello, C.*2 & Galetti M.2 ...... 28 Dracxler, C.M.*1, Pires, A.S.2, Fernandez, F.A.S.3 & Oliveira-Santos, L.G.R...... 28 Escribano-Avila, G.1, Calviño-Cancela, M.2, Pías, B.3, Escudero, A.4 & Virgós, E.4 ...... 29 Falcón, W.*1, Kaiser-Bunbury, C.2, Bunbury, N.3 & Hansen, D.1 ...... 29 Farwig, N., Berens, D.G. & Grass, I...... 30 Fedriani, J.M.1,2,3, Wiegand, T.1, Palomares, F.2 & Grimm, V.1 ...... 30 Forget, P.-M., Albert, A., Boissier, O., Bouiges, A., Mendoza, I. & Feer, F...... 31 Fricke, E.C.*1, Tewksbury, J.J.1,2, Wandrag, E.M.3 & Haldre, S.3 ...... 31 García, D.1, Donoso, I.1, Schleuning, M.2,3 & Fründ, J.4 ...... 31 García, D., Martínez, D., Donoso, I. & Rodríguez-Pérez, J...... 32 González-Varo, J. P. & Jordano, P...... 32 Govender, K.*, Shuttleworth, A., Downs, C.T. & Johnson S.D...... 33 Green, R.J...... 33 Hansen, D.1, Falcón, W.1 & Moll, D.2 ...... 34 Hart, L.A.*, Johnson, S.D., Downs, C.T...... 34 Haurez, B.1,2*, Petre, C-A.1,3,4, & Doucet, J-L.1,2 ...... 34 Ramanantoanina, A.1,2, Ouhinou, A.3 & Hui, C.1,2 ...... 35 Zhang, F., Minoarevilo, H.O. & Hui, C...... 36 Midgley, J.J.1, White J.1, Johnson S.D.2 & Bronner G.1 ...... 36 Jones, L.R.*1, Johnson, D.M.2, Duke-Sylvester, S.M.1 & Leberg, P.L.1 ...... 36 Kaplin, B.A.1, Martino, R.1 & Cordeiro, N.2 ...... 37 Kelly, D., Macfarlane, A.F. & Briskie, J.V...... 37 Kitamura, S. & Sawada, M...... 38 Kromann-Clausen, A.1,2, Dabelsteen, T.1 & Campos-Arceiz, A.2 ...... 38 Kwit, C., Patton, M.T. & Miller, C.N...... 39 Luna, C.A.1, Loayza, A.P.1 & Squeo, F.A.1,2 ...... 39 McConkey, K.R...... 39 Mokotjomela, T.M.*1,2, Hoffmann, J.H.2 & Downs C.T.3 ...... 40 Moran, C.1,2,3, Freeman, A.N.D.4, Catterall, C.P.1, Elgar A.T.1, Freebody, K.5, Montenero, M.4, Shoo, L.P.6 ...... 40 Mueller, T.1, Lenz, J.1, Caprano, T.1, Fiedler, W.2 & Böhning-Gaese, K.1 ...... 41 Neuschulz, E.L.1, Mueller, T.1,3, Bollmann, K.2, Gugerli, F.2 & Böhning-Gaese, K.1,3 ...... 41

Niebuhr, B.S.*1 & Campos, R.P.2 ...... 42 Pedrosa, F.* & Galetti, M...... 42 Pizo, M.A.1, da Silva, F.R.2 & Jacomassa, F.A.F.1 ...... 43 Prasad, S.1 & Westcott, D.A.2 ...... 43 Ramos, D.C.T.L.*1 & Silva, W.R.2 ...... 44 Reynolds, C.* & Cumming, G.S...... 44 Rogers, H., Wandrag, E. & Dunham, A...... 44 Ruggera, R. A.*1,2,3, Blendinger, P.G.1,2 & Gomez, M.D.*1,3 ...... 45 Schleuning, M.1, Fründ, J.2 & García, D.3 ...... 45 Solórzano-Filho, J.A...... 46 Sridhara, S.*1, Prasad, S.2, Westcott, D.3 & Edwards, W.1 ...... 46 Terborgh, J.W.1, Zhu, K.2, Alvarez-Loayza, P.1 & Valverde, F.C.3 ...... 47 Thabethe, V.*, Wilson, A-L.*, Hart, L.A.* & Downs, C.T...... 47 Timóteo, S.1,2, Correia, M.1, Rodriguez, S.E.1 & Heleno, R.1 ...... 48 Tjelele, J.1,2, Ward, D.2 & Dziba, L.3 ...... 48 Türke, M.1,2†, Fischer, C.3, Weisser, W.W.4 & Eisenhauer, N.1,2...... 49 Kaplin, B.1, Carlo-Joglar, T.A.2, Acevedo, J.D.3 & Torres-Caba, J.*1 ...... 49 Trolliet, F.*1, Forget, P-.M.2, Huynen, M-C.1 & Hambuckers, A.1 ...... 49 Tsuji, Y...... 50 Valido, A., Pérez-Méndez, N.1, Jordano, P.1, & García, C2 ...... 50 Wang, B...... 51 White, J.D.M., Midgley, J.J. & Bronner, G.N...... 51 Wilson, A-L.*, Shuttleworth, A. & Downs, C.T...... 52 Wotton, D.M.1,2, McAlpine, K.G.2 ...... 52 Zungu, M.M.* & Downs, C.T...... 52 Zwolak, R., Bogdziewicz, M.* & Wróbel, A...... 53 FSD POSTER ABSTRACTS ...... 54 Burns, K.C...... 54 Cestari, C.1, Pizo, M.A.1 & Loiselle, B.2 ...... 54 Cortinoz, J.1, Traveset, A.2 & Siva, W.3 ...... 54 Dracxler, C.M.*1, Confais, A.2 & Forget, P-.M.1 ...... 55 Ehlers Smith, D.A.*, Ehlers Smith, Y.C. & Downs, C.T...... 55 Escribano-Avila, G...... 56 Falcón, W.*1, García, C.2, Ozgul, A.1 & Hansen, D.1 ...... 56 Jara-Guerrero, A.1, Espinosa, C.I.1, De la Cruz, M.2,3, Méndez, M.3 ...... 57

Kitamura, S.1, Chodapisitkul, S.2, Sooktawee, A.2 & Poonswad, P.3 ...... 57 Kuhlmann, M.*1 & Ribeiro, J.F.2 ...... 58 Martínez-López, V. *1,2, Zapata, V.*1, Smith-Ramírez, C. 3,4 & Robledano-Aymerich, F.1 ...... 58 Mokotjomela, T.M.*1, Downs C.T.2, Knight, J.1 & Esler, K.3 ...... 59 Moran, C.1, 2, 3 & Catterall, C.1 ...... 59 Muñoz, M.C.*1,2, Schaefer, H.M.3, Böhning-Gaese, K.1,2 & Schleuning M.1...... 59 Naoe, S.1, Masaki, T.1 & Sakai, S.2 ...... 60 Pérez-Méndez, N.*1, Jordano, P.1, García, C.2 & Valido, A.1 ...... 60 Ramos, D.C.T.L.*1, Pizo, M.A.2 & Ribeiro, M.C.¹ ...... 61 Schupp, E.W.1, Bennion, L.D.1, Gómez, J.M.2 & Jordano, P.3 ...... 61 Shi, L., Lin, Y. & Wang, Z...... 62 Smith-Ramirez, C.1, Mora, J.P.2 & Zapata, V.M.*3 ...... 62 Strasberg, D.1, Flores, O.1 & Thébaud, C.2 ...... 63 Takahashi, K.1 & Takahashi, K.2 ...... 63 Tsuji, Y...... 63 Vogeler, A.V.1, Bohning-Gaese, K.2,4, Vollstaedt, M.2 & Tschapka, M.1,3 ...... 64 Vollstädt, M.1, Schleuning, M.1, Tschapka, M.2,3, Vogeler, A.2 & Böhning-Gaese, K.1,2 ...... 64 Wróbel, A.* & Zwolak, R...... 65 Zaniratto, C.P.1, Costa, J.C.2 & Silva, W.R.3 ...... 65 Zapata, V.*1, Martínez-López, V.1,2 & Robledano-Aymerich, F.1 ...... 66 Acknowledgements ...... 67 Summary list of delegates ...... 68 How to attend a scientific conference to your best advantage ...... 71 Conferences in 2016 you may be interested in ...... 73

Chairperson Welcome to FSD 2015

It is with great pleasure and enthusiasm that the 6th International Frugivory and Seed Dispersal Symposium (FSD) and Workshop is being hosted in the Drakensberg, South Africa. This is the first time that FSD is held in Africa and is a great opportunity for those interested in frugivory and seed dispersal both in Africa and internationally to meet and share ideas.

With the human population now > 7 billion there is continued and increased pressure on the environment. Land transformation for urban sprawl, agriculture and biofuels continues at high rates. The effects of accelerated climate change further exacerbate the situation. Another factor is the impact of invasive species. Despite all this there is a need to maximise biodiversity and sustain ecosystem function. Fragmentation, degradation, habitat-loss and hunting have cascading effects on wild populations in the form of local extinction, population depletion, behavioural changes or dispersal. Many Protected Areas (PAs) are relatively small and may not provide sufficient protection for species residing outside PAs. Africa is possibly the most affected. Consequently it is most relevant that the FSD is being held here in 2015. One of the other important facets of FSD is its interdisciplinary nature and that it is not taxon specific. We hope that this will generate much fruitful discussion and future collaborative research.

I am most grateful to the plenaries, symposia chairs, funders of the conference, the scientific committee, the local organising committee, the management and staff of Alpine Heath, Andrew de Klerk of Quartex, Prenisha Rajdev, Paige Potter, Mark and Tracey Marshall of Ndaka Safari Lodge, Preshnee Singh, Vicki Hooper and Lliane Bratherton who have all contributed towards the conference implementation.

Best wishes for a stimulating and enjoyable conference

Prof Colleen T. Downs Chairperson: Local Organising Committee School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg campus, South Africa June 2015

Scientific Committee

Surname Name Email Institution

Borges Renee [email protected] Indian Institute of Science, Bangalore , India

Cordeiro Norbert [email protected] Field Museum, Chicago, USA

Downs Colleen [email protected] University of KwaZulu-Natal, Pietermaritzburg campus, South Africa

Farwig Nina [email protected] Philipps University of Marburg , Germany

Forget Pierre-Michel [email protected] Muséum National d'Histoire Naturelle, Brunoy, France

Hansen Dennis [email protected] University of Zurich, Zurich, Switzerland

Shumpei Kitamura [email protected] Ishikawa Prefectural University, Nonoichi, Japan

Local Organising Committee

Surname Name Email Institution Role

Downs Colleen, Prof [email protected] University of KwaZulu-Natal, Pietermaritzburg Chairperson campus, South Africa

Hart Lorinda, Dr [email protected] University of KwaZulu-Natal, Pietermaritzburg Member campus, South Africa

Richardson Dave, Prof [email protected] University of Stellenbosch, Stellenbosch, South Member

Africa

Shuttleworth Amy-Leigh, Dr [email protected] University of KwaZulu-Natal, Pietermaritzburg Member campus, South Africa Singh Preshnee [email protected] University of KwaZulu-Natal, Pietermaritzburg Member campus, South Africa

Ally Eebee [email protected] University of KwaZulu-Natal, Pietermaritzburg Member campus, South Africa Mokotjomela Thabiso [email protected] University of the Witwatersrand, Johannesburg Member South Africa Hooper Vicki [email protected] Venues For Conferences In Africa Member

Bratherton Lliane [email protected] Venues For Conferences In Africa PA

INSTRUCTIONS to FSD 2015 PARTICIPANTS

Oral papers All delegates should give their electronic Powerpoint presentations to the FSD 2015 Secretariat as soon as possible after their arrival and registration. The final hand-in time for each presentation is the evening prior to the day on which the oral takes place. Presenters should make themselves known to the session chairs before the start of the session they are presenting in.

Poster presentations Posters will be numbered; during the coffee breaks each day, authors assigned to “odd numbers” or “even numbers” should be present at their posters so the delegates wishing to discuss their work can meet them.

Student presentations Student presentations are indicated by an asterisk (*). There will be prizes for the best student oral and poster presentations.

FSD 2015 PROGRAMME

Sunday 21 June Main Reception, Alpine Arrival, Check in hotel. Heath 14.00-17.00 Main Reception, Alpine Registration. Heath 17.00 Boma, Alpine Heath Welcome and opening of FSD 2015- Prof Deo Jaganyi, Deputy Vice-Chancellor & Head of College of Agriculture, Engineering & Science, University of KwaZulu-Natal. Snacks and drinks available. 18.00 Boma, Alpine Heath Zulu Drummers to perform.

Note: Each morning there will be bird ringing at sunrise for those interested- see the noticeboard for further details.

FSD 2015 Programme Monday 22 June

7.45-8.30 Late registration

MAIN HALL

8.30-8.40 Welcome Address

8.40-8.45 Welcome & Introduced by Steve Johnson Plenary introduction

8.45-9.20 Midgley, J.J. Megaherbivore fruits; an African Plenary elephant perspective 9.20-9.25 Questions

9.25-9.30 Plenary Introduced by Dennis Hansen introduction 9.30-10.05 Campos-Arceiz, A. Seed dispersal by elephants and Plenary other rapidly declining megafauna in tropical Asia 10.05-10.10 Questions 10.10-10.45 TEA/ COFFEE BREAK & POSTERS MAIN HALL ADJACENT HALL

10.45-10.50 Symposium: Chair: Kitamura Shumpei Symposium: Chemical Chair: Renee M. Borges & Steve Introduction Ecology & Ecology of Seed Johnson to Evolution Dispersal symposium

10.50-11.05 McConkey, K.R. Not all frugivores are created equal: Borges, R.M. The process of ripening in wild identifying the most effective seed infructescences within a brood-site dispersers in an Asian rain forest pollination system 11.05-11.20 Wang, B. Unequal seed dispersal by seed-size Govender, K.* The role of volatiles in the dispersal of fig sensitive rodents: a case study in seeds (Moraceae) by frugivorous bats in Chinese white pine (Pinus armandii) southern Africa 11.20-11.35 Zwolak, R. Benefits of masting for the European Johnson, S.D. Faecal mimicry by seeds for dispersal by beech: timing of granivore satiation dung beetles and benefits of seed caching support the predator dispersal hypothesis

11.35-11.50 Clausi, B. Frugivory by the bare throated Bogdziewics, M.* Chronic nitrogen pollution alters traits of bellbird in eastern Paraná state , red oak acorns and changes behavior of Brazil : insights for conservation of rodent seed dispersers the Atlantic rainforest

11.50-12.05 Kitamura, S. Dispersal of Ginkgo biloba Steele, M. (by Bogdziewics, The fruit-vs.-seed hypothesis: The (Ginkgoaceae), a large-seeded tree M. or Zwolak, R) adaptive significance of chemical species, in a temperate forest in gradients in acorns for escaping seed Japan predation and promoting seed dispersal

12.05-12.20 Solórzano-Filho, J.A. Big nut for a small mammal: The Zungu, M.M.* Effects of tannins on fruit selection in importance of Proechimys spp. three southern African frugivorous birds (Echimyidae) in Brazil nut seed dispersal 12.20-12.30 Discussion and questions with all presenters Discussion and questions with all presenters

12.30-13.30 LUNCH

MAIN HALL ADJACENT HALL 13.30-13.35 Plenary Introduced by Renee M. Borges introduction 13.35-14.10 Effiom, E. Consequences of bushmeat Plenary hunting in the Afrotropics

14.10-14.15 Questions

14.15-14.20 Symposium: Chair: Nina Farwig Symposium: General Chair: Colleen Downs Interaction Networks 14.20-14.35 Fricke, E.C.* Mutualistic strategy tradeoffs Türke, M. Invasive gastropods and earthworms as determine seed dispersal network potential dispersers of seeds of arable structure and increase robustness to plants coextinction

14.35-14.50 García, D. Towards a functional view of seed White, J.D.M* First use of camera traps to study scatter dispersal networks: a framework hoarding in Cape fynbos based on seed disperser effectiveness and plant fate

14.50-15.05 Ruggera, R.A.* Linking structure and functioning in Wilson, A.L.* Assimilation efficiency of the Red-winged plant-frugivore interactions: Starling Onychognathus morio fed importance of core frugivores and Strelitzia nicolai arils equivalence in their seed dispersal

15.05-15.10 Discussion and questions with all presenters Discussion and questions with all presenters

15.10-15.45 TEA/ COFFEE BREAK & POSTERS

MAIN HALL ADJACENT HALL Symposium: Chair: Nina Farwig Interaction Networks 15.45-16.00 Hui, C. Adaptive interaction switching and phylogeny explains the architecture of mutualistic networks

16.00-16.15 Pizo, M.A. Role of birds and bats in restored seed dispersal networks

16.15-16.30 Bender, I.M.A.* Feeding originality and flexibility of birds in plant-frugivore interaction networks

16.30-16.45 Falcón, W.* (by Giants, birds, and bats: First Hansen, D) approximation of the seed dispersal network of Aldabra Atoll

16.45-16.55 Discussion and questions with all presenters 18h00 DINNER

FSD 2015 Programme Tuesday 23June

MAIN HALL ADJACENT HALL

8.30-8.40 Announcements and address 8.40-8.45 Welcome & Plenary Introduced by Kitamura Shumpei introduction 8.45-9.20 Böhning-Gaese, K. Global change, interaction networks Plenary and seed dispersal services of birds 9.20-9.25 Questions 9.25-9.30 Plenary Introduced by Nina Farwig introduction 9.30-10.05 Traveset, A. Causes and consequences of seed Plenary dispersal disruptions

10.05-10.10 Questions 10.10-10.45 TEA/ COFFEE BREAK & POSTERS MAIN HALL ADJACENT HALL

10.45-10.50 Symposium: Chair: Catherine Moran & Symposium: Seed fate: Chair: Pierre-Michel Forget & Introduction to Patterns and Matthias Schleuning Understanding seed Hazel Chapman symposium Processes in dispersal, seed survival, Frugivore-Plant and plant recruitment in Interactions a changing world

10.50-11.05 Schleuning, M. A trait-based concept to integrate Christianini, A.V. Diplochory increases the effectiveness of structural and functional components seed dispersal of a tropical shrub of plant-frugivore interactions

11.05-11.20 Escribano-Avila, G. Functional complementarity of the Culot, L. Defaunation of large seed dispersers dispersal community leads to forest changes recruitment of a hardwood tree recovery & expansion after land leading to changes in carbon stock abandonment

11.20-11.35 Blendinger, P.G. Community-wide effects of fruit and Loayza, A. Intraspecific variation in seed size affects seed scaling in plants dispersed by the outcome of the interaction between vertebrates in subtropical Andean scatter-hoarding rodents and a threatened Atacama Desert shrub forests

11.35-11.50 González-Varo, J.P. Incorporating time into the seed Timoteo, S. The refaunation of Gorongosa NP and the dispersal effectiveness framework reinstatement of the seed dispersal service

11.50-12.05 Moran, C. Overcoming barriers to seed dispersal Pedrosa, F.* Feral pigs as taxon substitutes: re- and seedling recruitment in retired establishing lost ecosystem functions in tropical pasture defaunated landscapes

12.05-12.20 Jones, L.R.* Animal traits mitigating the negative Forget, P.M. Using trees and fruit ‘to take the consequences of fragmentation on temperature’ of rainforests: a study case spatial patterns of long-distance and along a gradient of anthropogenic disturbances in French contagious seed dispersal Guiana

12.20-12.30 Discussion and questions with all presenters Discussion and questions with all presenters 12.30-13.30 CONFERENCE PHOTOGRAPH FOLLOWED BY LUNCH MAIN HALL ADJACENT HALL

13.30-13.35 Plenary Introduced by Pierre-Michel Forget introduction 13.35-14.10 Westcott, D.A. Seed dispersal processes and their Plenary influence on global change impacts in tropical forests

14.10-14.15 Questions

14.15-14.20 Symposium: Chair: Catherine Moran & Symposium: Anthropogenic Chair: Thabo Patterns and Matthias Schleuning Impacts on Frugivory and Seed Mokotjomela Processes in Dispersal: Mechanisms, Scales Frugivore-Plant & Consequences Interactions 14.20-14.35 Blendinger, P.G. Fruits of Yungas Andean forests as birds Albert-Daviaud, A. The value of disturbance-tolerant see them Cercopithecine monkeys as seed dispersers in degraded habitats

14.35-14.50 Sridhara, S.* Behaviour mediated by habitat Ramos, D.C.T.L.* Vegetation functional and structural traits heterogeneity and not fruiting trees shaping seed rain underneath isolated trees better predicts seed deposition patterns by a tropical ruminant 14.50-15.05 Campos, C.M. Can dryland use modify the assemblage Kaplin, B.A. Matrix effects on frugivory and seed of animals removing Prosopis flexuosa dispersal in tropical montane forest: land (Fabaceae) seeds? use type in the matrix matters

15.05-15.10 Discussion and questions with all presenters Discussion and questions with all presenters 15.10-15.40 TEA/ COFFEE BREAK & POSTERS Symposium: Chair: Catherine Moran& Symposium: Chair: Thabo Mokotjomela Patterns and Matthias Schleuning Anthropogenic Processes in Impacts on Frugivore-Plant Frugivory and Interactions Seed Dispersal: Mechanisms, Scales & Consequences

15.45-16.00 García, D. The Biodiversity-Ecosystem Function Kromann-Clausen, A. Megafauna-caused differences in tree link in Frugivory and Seed Dispersal: distribution between mineral licks and effects across dispersal components random locations in a Malaysian forest and scales

16.00-16.15 Acosta Rojas, D.C. Can Woolly monkeys (Lagothrix Trolliet, F.* Effect of hunting pressure and forest lagotricha lugens) directly affect seed fragmentation on seed dispersal of Staudtia germination at Cueva de los Guacharos kamerunensis (Myristicaceae) in the National Park, Colombia? Western Congolian forest-savanna mosaic

16.15-16.30 Clausi, B. Frugivory by Procnias nudicollis Dracxler, C.M.* Living without seed dispersers: shortening (Cotingidae) in a 12 years study at of seed shadows drastically change eastern Parana, Brazil recruitment curves of three large-seeded palms in defaunated Atlantic Forest fragments

16.30-16.45 Prasad, S. Developing a generic functional Fedriani, J.W. Old-field recolonization under selective classification for seed dispersal defaunation: a spatially-explicit individual- based simulation model

16.45-16.55 Discussion and questions with all presenters Valido, A. (16.45-17.00) Downsized mutualisms: Demographic and genetic consequences of lizard body size reduction for an insular plant species

17.00-17.05 Discussion and questions with all presenters

18h00 DINNER

FSD 2015 Programme Wednesday 24 June

MID CONFERENCE TOUR

FSD 2015 Programme Thursday 25 June

MAIN HALL ADJACENT HALL

8.30-8.40 Announcements and address 8.40-8.45 Welcome & Introduced by Dennis Hansen Plenary introduction 8.45-9.20 Galetti, M. Frugivores and seed dispersal in the Plenary Anthropocene 9.20-9.25 Questions 9.25-9.30 Plenary Introduced by Kitamura Shumpei introduction 9.30-10.05 Tsuji, Y.T. Macaques as seed dispersers in Plenary Asian forests 10.05-10.10 Questions 10.10-10.45 TEA/ COFFEE BREAK & POSTERS

10.45-10.50 Symposium: Chair: Christophe Baltzinger Symposium: Chair: Ran Nathan Introduction to Seed Dispersal Movement Ecology, symposium by Animals as an Dispersal Kernels, Ecological Filter and Genetic Effects

10.50-11.05 Neuschulz, E.L. Seed dispersal by a food-hoarding bird Reynolds , C.* Bigger birds disperse better: seed traits counteracts plant establishment matter less than vector for aquatic plant dispersal

11.05-11.20 Baltzinger, C. Seed dispersal by large ungulates: a Mueller, T. Bird movements reveal seed dispersal multi-stage ecological filtering process networks in fragmented landscapes

11.20-11.35 Kwit, C. Plant advantages in myrmecochory: Niebuhr, B.S.* Effects of animal movement and space on redirecting ‘directed dispersal’ interaction networks in different tropical rainforest successional stages

11.35-11.50 Burns, K.C. A synthesis of seed dispersal by New Hui, C. Spatial assortment of mixed propagules Zealand weta explains the acceleration of range expansion

11.50-12.05 Tjelele, T.J. The effects of seed ingestion by Bunney, K.M. Elephantine carriers: The seed dispersal livestock, dung fertilization, trampling, kernel of the African savanna elephant grass competition and fire on seedling establishment of two woody plant species 12.05-12.20 Tsuji, Y.T. Inter-annual and inter-rank variation in Chibesa, M. GPS telemetry of Trumpeter Hornbill characteristics of endozoochory by wild (Bycanistes bucinator): Habitat use, foraging Japanese macaques (Macaca fuscata) and movement in urban environment of Eshowe, KwaZulu-Natal Province, South Africa

12.20-12.30 Discussion and questions with all presenters Discussion and questions with all presenters

12.30-13.30 LUNCH

MAIN HALL ADJACENT HALL

13.30-13.35 Plenary Introduced by Nina Farwig introduction 13.35-14.10 Jansen, P. Seed dispersal by a seed-predating Plenary rodent 14.10-14.15 Questions 14.15-14.20 Symposium: Chair: Christophe Baltzinger Seed Dispersal by Animals as an Ecological Filter 14.20-14.35 Chen, S.C.* A mammoth mouthful? A test of the idea that big animals disperse big seeds

14.35-14.50 Hansen, D. Frugivory and seed dispersal by chelonians: A systematic review and synthesis

14.50-15.05 Green, R.J. Popularity doesn't always lead to success: why aren't there more Ficus saplings? 15.05-15.10 Discussion and questions with all presenters

15.10-15.40 TEA/ COFFEE BREAK & POSTERS ADJACENT HALL MAIN HALL - preparation for Dinner

15.40-15.45 Symposium: Chair: Katrin Böhning-Gaese Functional Ecology and Conservation of Ecosystem Services

15.45-16.00 Bello, C.* Defaunation of large frugivores impacts carbon storage in tropical rainforests

16.00-16.15 Haurez, B.* Western lowland gorilla seed dispersal: effect of fecal matrix and deposition at nest sites on seedling growth 16.15-16.30 Terborgh, J.W. Seed rain predicts diversity of a tropical floodplain forest in Perú

16.30-16.40 Discussion and questions with all presenters

18h30 for GALA DINNER 19h00

FSD 2015 Programme Friday 26 June

MAIN HALL

8.30-8.35 Announcements and address

8.35-8.40 Welcome & Plenary Introduced by Colleen Downs introduction

8.40-9.15 Nathan, R. Opportunities and challenges in plant dispersal research: how could Kuhn and Plenary Galison run neck and neck in the race for glory? 9.15-9.20 Questions

9.20-9.25 Symposium: Invasives and Chair: David Richardson & Anna Traveset Introduction seed dispersal 9.25-9.40 Farwig, N. Guild-specific shifts in visitation rates of frugivores with habitat loss and plant invasion 9.40-9.55 Torres-Caba, J.* The role of introduced rats (Rattus rattus) as seed dispersers and predators in La Olimpia Forest, Adjuntas, Puerto Rico 9.55-10.10 Mokotjomela, M.T.* The potential for birds to disperse the seeds of Acacia cyclops, an invasive alien plant in South Africa 10.10-10.15 Discussion and questions with all presenters

10.15-10.40 Tea/ Coffee break

10.40-10.45 Symposium: Invasives and Chair: David Richardson & Anna Traveset Introduction seed dispersal to symposium 10.45-11.00 Kelly, D. European blackbirds (Turdus merula) in New Zealand: useful substitutes for lost mid-sized native frugivores, or weed vectors? 11.00-11.15 Thabethe, V.* Effect of invasive and indigenous avian species on seed germination of fleshy-fruited invasive alien plants in South Africa

11.15-11.30 Hart, L. Wahlberg's Epauletted Fruit Bats’ (Epomophorus wahlbergi) effect on alien invasive seed germination and their potential as dispersal agents 11.30-11.45 Wotton, D. Frugivory and seed dispersal of invasive plants in New Zealand

11.45-11.50 Discussion and questions with all presenters 11.50-12.30 CLOSING REMARKS Nina Farwig with the Panel of the Scientific committee Plus a remembering of those deceased since the last conference 12.30-13.30 LUNCH

13.30 Shuttles depart

FSD PLENARY ABSTRACTS

Böhning-Gaese, K. Senckenberg Biodiversity and Climate Research Centre, Senckenberganlage 25, DE-60325 Frankfurt (Main), Germany [email protected] Global change, interaction networks and seed dispersal services of birds Frugivory and seed dispersal is only one step in the regeneration cycle of plants and one of a multitude of ecological processes that take place in ecosystems. With the increasing necessity to record, analyse and project the impact of global change on biodiversity, ecosystem services and human well-being, it is also necessary to ask what roles frugivory and seed dispersal play in this larger context. A meta-analysis on the impact of forest fragmentation and local disturbance on four steps in the life cycle of woody plants, i.e. pollination, seed dispersal, seed predation and regeneration, shows that the steps in the life cycle that depend on mutualistic interaction partners, i.e. pollination and seed dispersal, are the most vulnerable to forest disturbance. A detailed analysis of the movement behaviour of birds in different types of landscapes demonstrates that seed dispersal kernels are extremely variable among bird and plant species, in space and over time. At the same time, modelling studies clarify that the consequences that climate change is projected to have on tree communities are most strongly modified by how realistically we model seed dispersal. Nevertheless, the study of frugivory and seed dispersal offers a unique opportunity to analyse and project the impact of climate change on biodiversity, interaction networks, and ecosystem functions. Fruit resource availability shapes bird community composition and richness, which itself determines seed removal and dispersal. This allows projections of the impact of climate change on interaction networks and the functional diversity of both plants and animals. Frugivory and seed dispersal are not only one of the most vulnerable steps in the life cycle of plants, but a good understanding of seed dispersal is also crucial for modelling the impact of climate change on tree communities. In addition, seed dispersal networks between plants and animals offer a unique opportunity for a better understanding of the impact of global change on biodiversity and ecosystem services.

Campos-Arceiz, A.1, Kromann-Clausen, A.1, Juwita, W.1, Mahmood, T.1, Solana Mena, A.1,2, Ning, H.1 & Ekanayaka, S.K.K.3 1The Management & Ecology of Malaysian Elephants (MEME), University of Nottingham Malaysia Campus, Malaysia, 2Department of Wildlife Management, PMAS - Arid Agriculture University, Pakistan, 3Centre for Conservation and Research, Sri Lanka [email protected] Seed dispersal by elephants and other rapidly declining megafauna in tropical Asia Tropical Asia is home to a complex guild of megafaunal herbivores that are either rapidly declining or functionally extinct. Little is known about their role in seed dispersal processes and the ecological consequences of their decline. Here we report on our ongoing efforts to study the role of Asian megafauna – especially elephants – in seed dispersal. In Malaysia’s tropical rainforests we found that elephants disperse the seeds of a much larger number of tree species than previously reported, including some with very large fruits and seeds such as the genera Mangifera, Durio, Parinari, and Borasodendron, and that megafaunal-syndrome trees are more abundant in patches frequently visited by elephants (mineral licks) than in other parts of the forest. In the dry human-dominated landscapes of Sri Lanka we found that sexual dimorphism affects elephant seed dispersal effectiveness, with males being more effective than females and juveniles dispersing the very large seeds of the palm Borassus flabellifer. Moreover, the high effectiveness of elephant dispersal also results in them acting as vectors of the invasive tree Prosopis juliflora. Tropical Asia’s megafaunal dispersers also include three species of rhinos, the Malayan tapir, and seven species of wild cattle and buffaloes, some of which must have been important forest tree dispersers before their populations collapsed. Tropical Asia is rapidly losing this important guild of dispersers. While it might be too late for some species, tropical Asia’s megafauna offer an opportunity for effective conservation and rewilding programs focusing on the restoration of ecological processes, particularly seed dispersal.

Effiom, E. Department of Biology, Biodiversity unit, Lund University, Sweden [email protected] Consequences of bushmeat hunting in the Afrotropics The largest primates are among the more efficient seed dispersers in the tropics, and hunting of these is a vexing problem around the globe. Through surveys, schematic modeling, manipulative experiments in seedling plots in paired sites with high and low hunting pressure in three areas in Southern Nigeria, we assessed the effects of primate bushmeat hunting on: 1) adult tree, seedling, and animal community compositions, 2) germination and survival among seedlings in association with competition, 3) community composition at multiple trophic levels, 4) leaf nitrogen concentration (LNC), leaf mass per area (LMA) from leaf samples collected from study sites and stem specific density (SSD), and 5) reliance of rural people on forest resources. In hunted sites, primates (ranging in biomass from 4–180 kg) including the Cross River gorilla, Gorilla gorilla diehli, were rarer, while seed predators increased in abundance. Community composition of birds was similar in paired sites. Seedlings and adult tree composition were similar in protected sites, dominated by primate-dispersed species. Abiotically dispersed species dominated in hunted sites having a higher germination rate. Seedling communities were significantly related to mammal communities. LNC and SSD increased with hunting but not LMA. Data from questionnaires administered in four forest-dependent villages around the study areas revealed that more primate dispersed trees have utility for humans compared to trees with other dispersal modes. Our results indicate that altered mammal community composition triggered by the decline of efficient seed-dispersing vertebrates have a major impact on changes in seedling communities and that these changes are largely detrimental to forest ecosystem functioning, biodiversity conservation and human well-being.

Galetti, M. Departamento de Ecologia, Universidade Estadual Paulista, Brazil [email protected] Frugivores and seed dispersal in the Anthropocene We live amidst large-scale change in animal composition and abundance around the world. Deforestation, hunting, habitat loss and pet trade are the major drivers of the Anthropocene defaunation. If on one hand, several species and populations of frugivores are declining or close to extinction, on the other hand, exotic species may replace some extinct species in seed dispersal processes. Here I discuss some ecological and evolutionary consequences of frugivore extinction particularly in a large hotspot biome: the Brazilian Atlantic forest. Based on a large dataset on plant–animal interactions, we predict long-term changes in plant composition, evolution and carbon storage. I argue that although scientists have advanced in many areas of the field of frugivores and seed dispersal, some intensive field-work is still necessary.

Jansen, P.A. Department of Environmental Sciences, Wageningen University, the Netherlands [email protected] Seed dispersal by a seed-predating rodent Many seed-eating rodents hoard large seeds as food supplies. Some do so by scattering seeds over numerous widely scattered shallow caches in the soil surface, a behaviour called scatter hoarding. These rodents tend to cache seeds close to where they find them, and they do so to ultimately recover and consume the seeds. Here, using agoutis in the tropical moist forests of Panama as the principal study system, I discuss how scatter hoarding can nevertheless yield effective seed dispersal for large-seeded plant species. I show that the rate of caching and the spread of caches is enhanced by competition among the rodents, that the rodents steal seeds from each other’s caches and re-cache them elsewhere, and that this theft is reciprocal, that theft produces multi-step dispersal over distances that can far exceed the radius of a typical rodent home range, that re-caching tends to move seeds towards areas with fewer

19 adult trees, that the rodents remember and manage their caches, and that predators may enhance seed survival by eating rodents. After concluding that agoutis are excellent seed dispersers in the study site that I used, I will discuss how this dispersal system is conditional on the relative abundances of seeds, rodents and predators, and how anthropogenic changes may affect dispersal success.

Midgley, J. Department of Biological Sciences, University of Cape Town, South Africa [email protected] Megaherbivore fruits, an African elephant perspective We investigated the interaction between marula (Sclerocarya birrea), torchwood (Balanites maughamii) and elephants. Germination experiments indicate that few seeds germinate from control seeds whereas many germinate from fruits eaten by elephants. We suggest that much of the positive influence of elephants is due to mastication, rather than acid scarification, which cracks open the tough seed coat of these two species. If true, then megaherbivores would be the only legitimate dispersers of these fruits. Both marula and torchwood are dull coloured but are strongly aromatic. Based on this work we suggest three further attributes that may define megaherbivore fruits: strong seed walls, dull coloration and strong scent. We observed that marula seeds in elephant dung are frequently taken and buried by savanna squirrels (Paraxerus cepapi). This suggests a triangular relationship in which scatterhoarded cracked seeds are likely to result in optimum establishment of marula seeds.

Nathan, R. Movement Ecology Lab, Dept of Ecology, Evolution and Behavior, The Hebrew University of Jerusalem, Jerusalem, Israel [email protected] Opportunities and challenges in plant dispersal research: how could Kuhn and Galison run neck and neck in the race for glory? The emerging field of movement ecology largely benefited from the recent development of new tracking technologies, enhanced computation abilities and powerful data analysis tools. Studies of plant dispersal in general, and by frugivores in particular, have utilized those technological advances to better understand dispersal processes and predict dispersal patterns. Such a technology-driven era offers new exciting opportunities but also entails significant challenges. In this presentation, I will summarize the major opportunities and challenges in seed dispersal research, illustrated for various dispersal systems mediated by wind, birds, bats and humans. I will also stress the need to promote a dynamic interplay between advancement of plant dispersal research by new tools (the Galisonian approach) and by new ideas (the Kuhnian approach). Major opportunities include faithful quantification of dispersal patterns and influencing factors, the re-examination of previous dogmas, conceptions and assumptions, and the broader scope and greater coverage enabled by current technologies. Major challenges include the apparent lag in developing new concepts and theories, the management and analysis of big data, and some other long-lasting technological and conceptual gaps, all still await future developments.

Traveset, A. IMEDEA, Spanish Research Council, Mallorca, Balearic Islands, Spain [email protected] Causes and consequences of seed dispersal disruptions Different drivers of global change are influencing seed dispersal processes in a variety of ecosystems worldwide. Among such drivers, habitat loss and fragmentation, defaunation (hunting) and species invasions appear as the most important. Seed dispersal disruptions are increasingly being documented as populations of legitimate animal dispersers decrease or even become locally extinct, although the ecological and evolutionary consequences for plant reproduction are still largely unknown. Little is also known about the patterns of such mutualistic disruptions. Are

20 the disruption consequences occurring mostly at the population level or at the level of the entire community? What are the evidences of community collapse? Is this most likely to happen in simple communities like island ones than in mainland systems? How frequent is functional redundancy, i.e. to what extent can lost seed dispersers be replaced by other (native or alien) frugivores that act as effective dispersers? Can a network approach help us to understand community impacts of seed dispersal disruptions? My talk will address all these questions, and I will present several case studies to illustrate both demographic and evolutionary consequences of disrupted seed dispersal mutualisms in the Mediterranean region.

Tsuji, Y. Primate Research Institute, Kyoto University [email protected] Macaques as seed dispersers in Asian forests In Asia, seed dispersal networks include various kinds of animals, such as primates, birds, carnivores and bats, in tropical regions, while the number of frugivores is limited in temperate regions. The macaques (genus Macaca), comprising about 20 species, inhabit Southeast (e.g., M. leonina, and M. fascicularis), South (e.g., M. mulatta, M. silenus and M. radiata), and East (e.g., M. fuscata and M. mulatta) Asian regions. They are mainly frugivorous, group- living, and large-sized animals, all of which seem contribute to effective seed dispersers in forest ecosystems in this region. Since the 1990s, their roles and effectiveness as seed dispersers, and their potential importance in forest regeneration have been studied. In this plenary talk, I review earlier knowledge on the seed dispersal services performed by the macaques, specifically 1) dispersal distance estimated by both field observation and feeding experiments for captive animals, 2) amount of seeds dispersed, 3) density of seeds after dispersal, 4) effect of environmental fluctuation on the effectiveness of the dispersal, and 5) regional variation (for example, tropical vs. temperate) in characteristics of seed dispersal by the macaques, paying special attention to their relative importance in forest ecosystems. Further, through introducing recent study topics, I would like to point out the advantage of macaque species as subject animals to address spatio-temporal variation in the environment on dispersal characteristics.

Westcott, D.A. CSIRO, Land and Water, Australia [email protected] Seed dispersal processes and their influence on global change impacts in tropical forests There is increasing recognition that projections of biodiversity outcomes under global change must incorporate the outcomes of ecological processes such as dispersal and inter-specific interactions. This is not a simple task. In wet tropical forests, some of the most threatened ecosystems, high diversity and a lack of data makes the task even more daunting. In this talk I outline the results of a collaboration that has explored one path through this problem. We used field survey data and empirical data on seed dispersal processes as the input for macro-ecological modelling of projections of biodiversity outcomes of global change for the distribution and structure of the vertebrate disperser (n=65 species) and plant communities (n=4313 species) of Australia’s tropical rainforests. Our results suggest that climate change will substantially alter the seed dispersal services provided by frugivorous vertebrates, reducing the ability of some plants to track shifts in suitable habitat. The net result is likely to be significant change plant distributions and plant community structure. Our results point to specific management responses that could ameliorate the effects global change on biodiversity outcomes, including the management of specific disperser populations and optimal strategies for habitat restoration. Interestingly, the complexity (or realism) of the dispersal model used in our models had a greater effect on projected biodiversity outcomes than did dramatically different climate change scenarios. This suggests that incorporating ecological processes, such as seed dispersal, can make an important difference in how we understand and respond to the influence of climate change on biodiversity.

FSD ORAL ABSTRACTS

Acosta Rojas, D.C., Cruz-Tejada, D.M., Páez-Crespo, E.J. & Stevenson, P.R. Universidad de los Andes [email protected] Can Woolly Monkeys (Lagothrix lagotricha lugens) directly affect seed germination at Cueva de los Guacharos National Park, Colombia? The effect of gut treatment on seed germination is an important determinant of seed dispersal effectiveness. The main objective of this study was to evaluate the effect of gut treatment by Woolly Monkeys on seed germination of three highland plant species (i.e. Hedyosmum cuatrecazanum, Tapirira guianensis subandina, Guettarda hirsuta) at Cueva de los Guácharos National Park, Colombia. We performed germination and permeability tests to examine whether the gut treatment effect was the result of pulp removal or an increase in seed permeability. In addition, we evaluated whether the initial conditions of seeds were different between treatments and if physical and physiological traits could explain differences in germination. Pulp removal by woolly monkeys was the main factor that enhanced seed germination of H. cuatrecazanum and T. g. subandina. The overall effect of gut passage on seed germination varied among species. For H. cuatrecazanum and G. hirsuta, we found similar germination percentages between dispersed and control seeds, but for T. g. subandina, control seeds had greater germination success than dispersed seeds. However, the initial morphological conditions of T. g. subandina seeds were different between treatments, and explained germination probability. Our study demonstrates a clear interaction between the initial conditions of seeds and the evaluation of the effect of frugivores on dispersed seeds in T. g. subandina. Therefore, we suggest that the results of experiments evaluating the effect of animal vectors on seed germination should be carefully interpreted, and that these types of experiments require an accurate quantification of seed initial conditions.

Albert-Daviaud, A.1,2, McConkey, K.3, Savini, T.4 & Huynen, M-C.1 1University of Liège, Belgium, 2National Museum of Natural History, France, 3National Institute of Advanced Studies, Indian Institute of Science Campus, India, 4King Mongkut's University of Technology Thonburi, Thailand [email protected] The value of disturbance-tolerant cercopithecine monkeys as seed dispersers in degraded habitats Habitat disturbance has caused a global decline in populations of frugivores, with critical consequences for seed dispersal. Large-seeded plants are especially threatened as they depend on a restricted number of large-bodied dispersers that are vulnerable to extinction and cannot maintain populations in most disturbed habitats. Cercopithecine monkeys are potentially key seed dispersers in disturbed habitats, because of the robustness of some species to disturbance and their ability to disperse large seeds. However, the potential ecological roles of the more disturbance-tolerant species are rarely discussed. This review evaluates the seed dispersal role of cercopithecines in disturbed habitats by investigating their ability to tolerate habitat disturbance, their seed dispersal abilities, and the threats to species survival. Cercopithecines are characterized by ecological flexibility, most species adjust their diet, group size, home range size and, often, feeding methods according to resource availability and habitat structure. Consequently, 79% of species are tolerant of varying degrees of habitat disturbance. Cercopithecines are often inconsistent seed dispersers, but they have the capacity to disperse many seeds, large seeds, and to disperse them across large distances. They may be among the most important frugivores in altered environments in Asia and Africa. However, many disturbance-tolerant cercopithecine species are targeted by local people as pests, which poses a major threat to their conservation. In conclusion, the management of all disturbance-tolerant cercopithecine species should be re-evaluated given their importance in the regeneration of degraded Asian and African habitats.

Baltzinger, C.1, Albert-Daviaud, A.1,2, Picard, M.1 & Mårell, A.1 1Irstea, Domaine des Barres, 45290 Nogent-sur-Vernisson, France, 2National Museum of Natural History, France [email protected] Seed dispersal by large ungulates: a multi-stage ecological filtering process Multi-stage ecological filtering mechanisms including seed dispersal, abiotic constraints and biotic interactions among plants and across trophic levels act upon regional species pools to form dynamic transient plant assemblages. Large ungulates disperse seeds (zoochory), modify the environment (ecosystem engineering) and consume plants (herbivory), whereby they shape plant assemblages by intervening at different levels in the ecological filtering process. The ecological processes through which large ungulates affect the environment make ecosystems more susceptible to invasion by non-resident species, we therefore include the role of ungulates in shaping plant assemblages within the framework of the theory of invasibility proposed in 2000 by Davis and co-workers. Here, we specifically highlight how complementary zoochorous processes in a guild of large ungulates affect the composition and spatial distribution patterns of plant assemblages. First, we use a trait-based meta-analysis to show how endozoochory and hoof- and hair-epizoochory filter the regional species pool differently. Then, we show how interactions among vectors, seeds and plant traits intervene in the seed dispersal cycle (emigration, transfer and immigration). Finally, we highlight how closely ungulates combine dispersal with abiotic filters. We show that large ungulates, as selective and daily long-distance seed dispersal vectors, play a role in determining the spatial and temporal dynamics of meta-communities. These animals are likely to affect plant responses to global changes such as habitat fragmentation and climate warming, and may have contributed to plant migration patterns since the last glaciation.

Bello, C*., Galetti, M., Magnago, L.F.S., Rocha, M., Lima, R., Peres, C.A. & Jordano, P. Universidade Estadual Paulista (UNESP) [email protected] Defaunation of large frugivores impacts carbon storage in tropical rainforests Carbon storage is a widely acknowledged and increasingly valued function of tropical forest ecosystems. Forest size reductions, logging, edge effects and climate change have shown significant impacts on tropical carbon stocks, however an elusive and yet undetected cause of change in carbon storage is the effect of defaunation of large frugivores. Since many large tropical trees, that make a large contribution to carbon stocks, rely on large frugivores for seed dispersal, we wanted to quantify the potential indirect effect of defaunation of large-bodied seed dispersers on carbon storage. We show that the extinction of large frugivores could lead to drastic long-term erosion of carbon storage in tropical rainforests. Analyzing a large dataset of tree species traits of a tropical biodiversity hotspot, the Brazilian Atlantic forest, we found a significant correlation (r=0.27, P <0.001) between traits that limit dispersion (seed diameter) and traits that influence carbon storage (wood density). Also, when we simulated the extinction of trees that depend on large frugivores in 25 tree communities, we found that defaunation erodes carbon storage by 0.5 Mg hectare-1 representing a total carbon loss effect of 3.32 Tg for the largest fragments of the Atlantic rainforest. Inter-governmental programs to reduce emissions of carbon dioxide have been mostly focused on deforestation and forest degradation, but our results demonstrate that defaunation of large frugivores also poses a serious risk for the maintenance of the carbon storage ecosystem service.

Bender, I.M.A.*1, Dehling, D.M.2, Hensen, I.3, Kissling, W.D.4, Kühn, I.5, Wiegand, T.6 & Schleuning, M.2 1German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Germany, 2Senckenberg Biodiversity and Climate Research Centre, Germany, 3Institute of Biology, Geobotany and Botanical Garden, Martin-Luther- University Halle-Wittenberg, Germany, 4Institute for Biodiversity and Ecosystem Dynamics (IBED), University of Amsterdam, the Netherlands, 5Helmholtz Centre for Environmental Research (UFZ) Leipzig-Halle, Germany, 6Helmholtz Centre for Environmental Research (UFZ) Leipzig-Halle, Germany

23 [email protected] Feeding originality and flexibility of birds in plant–frugivore interaction networks A fundamental ecosystem function in tropical forests is seed dispersal by frugivorous birds. Bird species differ fundamentally in their feeding niches, but little is known about the variability of these niches over time. We use two plant–frugivore networks in the tropical Andes to examine seasonal variation in the foraging preferences of frugivorous birds, employing a trait-based analysis of interaction networks. To quantify the foraging preferences of frugivorous birds, we projected interactions between birds and plants onto a multi-dimensional plant trait space and calculated the seasonal and overall interaction centroids for each bird species in plant trait space. We then measured functional originality (the distance of the interaction centroid of a bird species to the community centroid) and functional flexibility (the distance of a bird's seasonal interaction centroid to its overall centroid). We found that functional originality and flexibility varied systematically throughout the year, but were phylogenetically conserved. There was a positive relationship between originality and flexibility, suggesting that the functionally most original species were also the most flexible in their foraging choices. Our results suggest that species that forage in a more original area of the plant trait space tend to be most flexible in switching to alternative fruit resources. This finding challenges the notion that specialized species have a low capacity to respond to changes in functional community composition, for instance due to climate change. The finding is also important for inferring unknown interactions in potential future communities of plants and frugivores.

Blendinger, P.G1 & Giannini, N.P.2 1CONICET & Instituto de Ecología Regional, Universidad Nacional de Tucumán, Argentina, 2CONICET & Facultad de Ciencias Naturales e IML, Universidad Nacional de Tucumán, Argentina [email protected] Community-wide effects of fruit and seed scaling in plants dispersed by vertebrates in subtropical Andean forests An adaptive basis exists by which plants and their dispersers may commit to a continuing relationship over time, leaving traceable patterns of plant–animal coevolution. An adaptive explanation of the variation in quantitative traits of whole fruit, pulp, and seeds would imply combinations of those values that help accomplish both the dispersal and nutrition functions of the interaction. However, an adaptive explanation of inter-specific patterns may emerge only after considering other options that may account for variation. With the aid of dimensional analysis, we dissected inter-specific effects of scaling in fleshy-fruited plants (121 species, 46 families) of subtropical Andean forests (Yungas). We contrast expected to empirically obtained relationships to test 15 specific predictions based on scaling, developmental, phylogenetic, and interactive hypotheses for quantitative relationships of the fruit, pulp and seed relevant to the plant–frugivore interaction. Phylogenetic relationships explained a minor fraction of the variability in fruit traits. We found strong support for most predicted relationships between fruit traits relevant for animal dispersal, community-level variation in dimensions, quantities and forms of fruits and seeds can be mostly explained by allometry, dimensionality, modularity and development, while adaptive explanations can account for deviations observed in individual species rather than for community-level responses. Additional aspects should be taken into account before judging whether the varying dimensions of the fruit, seeds, and pulp respond adaptively to forces shaping the plant–frugivore interaction. We conclude that adaptive responses to forces shaping the plant– disperser mutualism cannot explain the strong variability observed in the dimensions of fruits, seeds and pulp.

Blendinger, P.G.1, Ordano, M.2, Lomáscolo, S.B.1, Chacoff, N.P.1, Núñez Montellano, M.G.3, Jiménez, J.1, Ruggera, R.A.*4, Valoy, M.2, Sánchez, M.S.5 & Martín, E.2 1CONICET & Instituto de Ecología Regional, Universidad Nacional de Tucumán, Argentina, 2 CONICET & Fundación Miguel Lillo, Argentina, 3CONICET & IBIGEO, Argentina, 4CIT Jujuy CONICET, Argentina, 5CONICET & Instituto de Biología Subtropical, Universidad Nacional de Misiones, Argentina [email protected] Fruits of Yungas Andean forests as birds see them

In seed dispersal mutualisms, natural selection should promote strategies of fruit display that maximize disperser attraction. Plants might increase seed dispersal profitability by increasing fruit conspicuousness, understood as the ease with which fruits are detected by dispersers. Because strategies to maximize fruit conspicuousness are various, and most are costly, we would expect that individual plant species invest primarily in only one. We explored this prediction using fleshy-fruited species of subtropical Andean forests (Yungas) and a large dataset on frugivore diets. As mechanisms of advertising enhancement, we considered the chromatic contrast between diaspores and foliage, fruit size and architecture (exposure and aggregation) of fruit display. Towards this goal, we (1) characterized the tetrahedral colorspace of fruits and the variation in chromatic contrast (just noticeable difference (JND)) using avian vision models, (2) assessed the relationship between fruit traits determining conspicuousness, and (3) analyzed the importance of fruit conspicuousness on fruit consumption by birds. The colorspace of Yungas fruits was considerably larger than the average reported for tropical forests of South America. Fruit chromatic contrast did not covary with other traits determining conspicuousness (fruit size, exposure and aggregation), although it tended to decrease with fruit size. Fruit consumption was explained by chromatic contrast depending on fruit size, exposure and aggregation. Within the diversity of fruit displays observed in the Yungas, plant species might be using display strategies that combine chromatic contrast with morphological and architectural fruit traits.

Bogdziewicz, M.*1,2, Steele, M.A.3, Crone, E.E.2 & Zwolak, R.1 1Adam Mickiewicz University, Poznań, Poland, 2Tufts University, Medford, USA, 3Wilkes University, Wilkes-Barre, USA [email protected] Chronic nitrogen pollution alters traits of red oak acorns and changes behavior of rodent seed dispersers The increase in anthropogenic inputs of reactive nitrogen caused by fossil fuel combustion is a recent stress for both terrestrial and aquatic ecosystems worldwide. In our research we studied how chronic nitrogen deposition (> 20 years) alters traits of red oak (Quercus rubra) acorns and how these changes influence dispersal of acorns by rodents. We found that chronic nitrogen deposition decreased tannin levels in red oak acorns while lipid levels were not affected. Moreover, acorn weight was slightly (but significantly) elevated in acorns collected from trees growing in fertilized plots (N acorns). Distance of acorn dispersal by rodents was higher in N acorns than in control acorns. On the other hand, rodents preferred to both remove and eat control rather than N acorns. Finally, proportion of cached acorns that survived till next spring was three times lower in N acorns than in the control. We conclude that increased dispersal distance was probably caused by the larger size of N acorns. Lower concentration of tannins in N acorns was most likely responsible for reduced removal and caching by rodents. Germination of red oak acorns is extremely low without burial, and in the eastern United States, acorns are buried mostly by rodents. Therefore, nitrogen deposition might impede natural regeneration of red oaks through its effects on the caching behavior of rodents.

Jana, A., Venkateswaran, V. & Borges, R.M. Centre for Ecological Sciences, Indian Institute of Science, Bangalore 560 012, India [email protected] The process of ripening in wild infructescences within a brood-site pollination system The process of ripening can influence fruit odours during the developmental process of the fruit. In the brood-site pollination system of Ficus (Moraceae), some flowers develop into seeds within an enclosed infructescence called a syconium, the development of other flowers, however, can be subverted by fig wasp gallers whose progeny develop within galled flowers. While the legitimate fig wasp pollinators of the system are flower gallers, there are several illegitimate galler fig wasp species that can gall floral primorida as well as syconium wall tissue. Working with a model wild fig system Ficus racemosa, about which basic information is already available on the syconium volatilome, we investigate how the presence of these different gallers influences the ripening process of the syconium and the volatile profile of the syconium. We experimentally alter the galling and ripening process and investigate the impact of these manipulations on the volatiles produced by the developing infructescence. We also

25 examine the volatilome of syconia which have been experimentally manipulated to produce only seeds. Our work furthers the understanding of the relationship between pollination, galling and the timing of these processes in the development of a brood-site whose dual function is to ensure successful reproduction of its legitimate occupants, i.e. pollinator fig wasp progeny and plant seeds.

Bunney, K.1, Bond, W.1 & Henley, M.2 1Biological Sciences Department, University of Cape Town. Cape Town, South Africa, 2Save the Elephants, Transboundary Elephant Research Programme, South Africa [email protected] Elephantine carriers: The seed dispersal kernel of the African savanna elephant Elephant seed dispersal studies have largely focused on African and Asian forest elephants. These studies have firmly established forest elephants as prolific seed dispersers, consuming more seeds from more species than any other taxon of large vertebrate disperser. African forest elephants are also the principle, and in some cases, the sole extant, seed dispersal agent for a number of large-fruited and large-seeded forest trees, thus filling a very specific ecological niche. Savanna or bush elephants are morphologically distinct from the elephants of the tropical forests of Africa. Despite this, little is known of the savanna trees that rely on elephants for their dispersal or the spatial scale at which these seeds are dispersed. We combined information from feeding trials conducted on sanctuary elephants with field telemetry data collected over an 8-year period in Timbavati/Kruger National Park to assess the seed dispersal service provided by savanna elephants. This study provides the first detailed account of the spatial scale at which African savanna elephants disperse seeds. Our mechanistic model predicts that 50% of seeds are carried over 2500 m and distances up to 65km are achievable in maximum gut passage time. These findings establish the savanna elephant as one of the longest distance vertebrate disperser yet investigated. Maintaining their ecological role as a seed disperser may prove to be a significant factor both in the conservation of large-fruited tree diversity within the savannas and in facilitating shifts in biome boundaries in response to global climate change.

Burns, K.C. Victoria University of Wellington [email protected] A synthesis of seed dispersal by the New Zealand weta New Zealand is home to gigantic, flightless crickets known as wetas, which are arguably the world’s most unusual seed disperser. Although insects generally do not consume fleshy fruits and disperse seeds after gut passage, recent work has shown that a diverse range of weta species disperse seeds in a variety of habitats, from temperate rainforests to alpine scrublands. Here, I will (1) review the evidence for seed dispersal by wetas, (2) highlight the controversy and debate surrounding the phenomenon, (3) derive a simple conceptual model to explain and predict variation in seed dispersal effectiveness among different-sized wetas, and (4) discuss how seed dispersal by the weta fits within the context of the ‘island syndrome’ or a suite of repeated patterns in the evolution of island biotas.

Campos, C.M., Miguel, M.F. & Cona, M.I. Instituto Argentino de Investigaciones de las Zonas Áridas (CONICET-CCT Mendoza), Argentina [email protected] Can dryland use modify the assemblage of animals removing Prosopis flexuosa (Fabaceae) seeds? The fruits of Prosopis flexuosa are indehiscent pods, with a mesocarp rich in sugars and protein and palatable to dispersing animals. The aim of this study was to analyse the role of seed removers at sites with different land uses (protected area with grazing exclusion: Ñacuñán Reserve, protected area with domestic animals included: Ischigualasto Park, unprotected area under continuous grazing pressure:adjacent to Ñacuñán Reserve). Fruit removal field trials are conducted using camera traps (Moultrie M-990i). At each site, we offered 20 pods (250 seeds) under 20 trees for 48h. The main results allow identifying and quantifying seed removal by animals that disperse seeds by

26 endozoochory (native species: Lycalopex griseus, Dolichotis patagonum, Rhea americana, Lama guanicoe, exotic species: Lepus europaeus, cow, horse, donkey) and scatter-hoarding (Microcavia asutralis), and animals that predate seeds (Graomys griseoflavus). In the protected area without livestock and in the unprotected area, the main species removing seeds were G. griseoflavus (41.75±1.4, mean ± SE), L. griseus(13.12±0.9), and M. australis (11.4± 00.9). In the protected area with domestic animals, endozoochorous mammals were the most important dispersers: donkey, L. griseus, cow, horse, and D. patagonum. This is the first study assessing the extent of P. flexuosa seed removal by different animals in drylands under different land use conditions. In the long term, grazing exclusion produced changes in habitat structure and a decrease in the abundance of some dispersers, such as D. patagonum and M. australis, promoting seed removal by sigmodontine rodents that mainly act as seed predators.

Chen, S-C.* University of New South Wales, Australia [email protected] A mammoth mouthful? A test of the idea that big animals disperse big seeds It has been widely assumed that large seeds generally require large dispersers to ingest and disperse them, but this relationship has only been quantified in single animal groups (e.g. birds) and in a few communities. We made the first global-scale study across all vertebrate groups (fish, amphibians, reptiles, birds and mammals), to test the prediction that large animals are able to ingest both large and small seeds while small animals ingest small ones. Literature data were compiled for over 12,000 seed dispersal interactions from over 300 study sites globally. Quantile regression showed a triangular relationship between seed size and disperser size (p < 0.001 for both 95th and 5th quantiles). The slope and significance of the relationship between body weight and ingested seed size varied between animal groups, presumably because of their diversified foraging behaviours, movement modes and digestive characteristics. Both reptiles and birds showed a significantly positive relationship, whereas mammals showed the opposite, indicating that absence of some dispersers differentially affected seed dispersal of large- and small-seeded species. In conclusion, the upper-bound size of ingested seed size was positively correlated with the disperser’s body weight. Our broad-scaled study confirmed that the size of ingested seeds depended on the body size and taxonomic group of the animal feeding on it, and indicated that elimination of large-bodied animals by hunting or habitat loss might impose restrictions on the regeneration of large-seeded plants.

Chibesa, M.*, Downs, C.T., Rouget, M., Taylor, B. & Ramesh, T. School Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] GPS telemetry of the Trumpeter Hornbill (Bycanistes bucinator): Habitat use, foraging and movement in urban environments of Eshowe, KwaZulu-Natal Province, South Africa Despite the many challenges posed by urbanisation to the existence and persistence of wildlife in human-dominated habitats, some species of wildlife still utilise the urban environment. The Trumpeter Hornbill (Bycanistes bucinator) is one of a few large avian frugivores that has been observed to utilise the urban environment. Understanding how wildlife persists and responds to urban life is critical to biodiversity conservation and urban planning. We used Global Positioning System (GPS) transmitters to record the diurnal movements of Bycanistes bucinator in Eshowe to better understand how this large frugivore is utilising the urban environment. GPS transmitters were attached to five individuals captured in Eshowe town between 7th March and 10th March, 2014. The species was monitored for a period of 1 year and yielded useful data. Majority of the locations were in urban environments (trees in gardens and streets) and the nearby intact Dlinza Forest. Our findings indicate that key food resources (such as figs) are vital in controlling movements and attracting Bycanistes bucinator to the urban environment as all the species tagged exhibited repeated foraging movements. Dlinza Forest acts as a primary food resource and roosting location. These findings are important in supporting policy formulation, decision making and species conservation in human- dominated environments.

Camargo, P.H.S.A. 1, Martins, M.M. 2 & Christianini, A.V.3 1Programa de Pós-Graduação em Diversidade Biológica e Conservação, Universidade Federal de São Carlos, Brazil, 2Universidade de Santo Amaro, Brazil, 3Departamento de Ciências Ambientais, Universidade Federal de São Carlos, Brazil [email protected] / [email protected] Diplochory increases the effectiveness of seed dispersal of a tropical shrub Plants often have complex seed dispersal patterns performed by different dispersers such as birds and ants in successive phases (i.e. diplochory). Yet, the contribution of each vector to seed dispersal effectiveness (SDE) is poorly understood. We investigated the contribution of birds and ants to SDE of Erythroxylum ambiguum in the Atlantic forest of Southeast Brazil between 2012–2014. We estimated transition probabilities (TP) from seed to one-year seedling for seeds dispersed by birds (from plant crown, Phase 1) and by ants (after the seeds reached the ground, Phase 2) with the aid of seed removal experiments, observations of places of seed deposition and seedling survival. We used the cumulative TP to estimate the SDE for each dispersal phase. We recorded 12 species of birds feeding on fruits and dispersing 26% of the fruit crop. The rest (74%) fell beneath the parental plant, 67% as viable seed. Twenty species of ants rapidly removed fallen diaspores (whole or embedded in bird droppings), thus redistributing the seed shadow generated by Phase 1. Rodents preyed on a third of seeds on the ground. Birds and ants increased germination percentage and rate similarly compared to controls. Birds dispersed seeds farther, where seedling survival was enhanced, but seedlings growing in ant nests showed higher one-year survival than controls. When the combined contribution of birds and ants in diplochory was considered, SDE was significantly increased compared to a single dispersal phase effected by birds. The ecological and evolutionary implications of diplochory will be discussed.

Clausi, B. Myrtlau, Brazil [email protected] Frugivory by the bare throated bellbird in eastern Paraná state, Brazil: insights for conservation of the Atlantic rainforest The bare throated bellbird is endemic to the Atlantic rainforest of South America. It is endangered in the greater part of its range. I focused on two main study sites, one in lowland and the other in highland forests to determine what the bellbirds are eating throughout the year. Several other sites were also sampled throughout the study, which started in 2005. A total of 132 tree species were classified as bellbird food, with only three species overlapping between the two sites. I concluded that the bellbirds make a complex altitudinal migration in the region and its conservation will only be achieved with a network of connected reserves. It is indeed a major disperser of seeds from many tree species and only the other cotingid in the area Carpornis cucullatus and the cracid Penelope obscura may have a similar impact on the dispersal of these fruits. However, in the highland site Carpornis is absent, possibly due to a local extinction. The bellbird might therefore be unique for the forest survival, as its dispersal seed rain is more directed than that of Penelope.

Clausi, B. Myrtlau, Brazil [email protected] Frugivory by Procnias nudicollis (Cotingidae) in a 12 years study at eastern Parana, Brazil The bellbirds are notorious for their impact in bird dispersed seeds, being amongst the most frugivorous of all neotropical birds. I checked what the species Procnias nudicollis is consuming to survive the year in eastern Parana state, south Brazil. Since 2003, my main study sites comprised: QU at 100 meters and BA at 900 meters of altitude. Proved fruits eaten 99 species, however estimated at least 200 species. Plant families total 20 with distribution in order of importance: Myrtaceae 45 species, Lauraceae 22, Moraceae 08, Sapindaceae 05, Arecaceae 02,

Symplocaceae 02, Myrsinaceae 02, other 13 families with only one specie. Myrcia lajeana case study shows more than 20 times more recruitment in male bellbird perches than in random at the forest.

Culot, L.1, Bello, C.*2 & Galetti M.2 1Departamento de Zoologia, Laboratório de Primatologia (LaP), Universidade Estadual Paulista (UNESP), Brazil, 2Departamento de Ecologia, Laboratório de Biologia da Conservação, Universidade Estadual Paulista (UNESP), Brazil [email protected] Defaunation of large seed dispersers changes recruitment of a hardwood tree leading to changes in carbon stock The loss of seed dispersers can alter plant recruitment and consequently the composition of future plant communities. Since wood density positively correlates with carbon storage, reduced seed dispersal and/or seedling recruitment of hard wood species is likely to decrease the carbon storage capacity of future forests. We examined the possible loss of carbon stock resulting from the loss of large seed dispersers (muriquis, howler monkeys and jacutingas) of one plant species having one of the highest carbon storage capacity in the Atlantic forest, Cryptocarya mandioccana (Lauraceae, 0.72g/cm3). We determined the carbon stock resulting from the presence of adult Cryptocarya trees and their recruitment success through the study of seed fate from fruit production to 1-year seedling survival. We investigated whether compensation effects followed the loss of Cryptocarya seed dispersers by comparing their contribution in three areas along a disperser-defaunation gradient. We found a partial compensation effect in the dispersal service of jacutingas in the absence of muriquis and howler monkeys, and of howler monkeys in the absence of muriquis. The future carbon storage capacity of Cryptocarya in our most disperser-defaunated site (only jacutinga present) is already potentially reduced by 52%, being equivalent to a loss of 1.2 tons of carbon/ha. The loss of all Cryptocarya dispersers in our non-defaunated site would potentially reduce the storage capacity by 84% (-0.9 tons/ha). Based on a one-species simple example, our study shows that the impact of large disperser defaunation can thus severely affect the carbon ecosystem services provided by tropical forests.

Dracxler, C.M.*1, Pires, A.S.2, Fernandez, F.A.S.3 & Oliveira-Santos, L.G.R. 1Muséum National d’Histoire Naturelle, Département Ecologie et Gestion de la Biodiversité, France, 2Departamento de Ciências Ambientais, Universidade Federal Rural do Rio de Janeiro, Brazil, 3Departamento de Ecologia, Universidade Federal do Rio de Janeiro, Brazil, 4Departamento de Biologia Animal, Universidade Federal de Mato Grosso do Sul, Brazil [email protected] Living without seed dispersers: shortening of seed shadows drastically change recruitment curves of three large- seeded palms in defaunated Atlantic Forest fragments Defaunation can disrupt or alter animal–plant interactions. Since vertebrates are usually more affected by hunting and forest fragmentation, we expect that vertebrate seed dispersal and predation may be more affected than invertebrate seed predation. Those responses, however, may depend on plant vulnerability to defaunation, which can be initially predicted by the set of mutualistic partners. We expected that regeneration of plants interacting with a higher number of dispersers would be less affected by defaunation, especially due to escape from seed predation through seed dispersal events. We utilized the palms Astrocaryum aculeatissimum, Attalea humilis and Euterpe edulis — whose coterie of dispersers increases in the order above — as a model to investigate how defaunation can affect seed shadow, seed predation and seedling recruitment curves in defaunated and non-defaunated Atlantic Forest fragments. While number of seeds dispersed decreased with distance from parents for all palm species in defaunated fragments, seed predation patterns were more variable, depending both on predator and palm species. Of more concern was that seedling recruitment curves were drastically affected for all palm species, with higher probabilities of recruitment near parents in defaunated areas, regardless of differences in seed dispersal and seed predation patterns. Our study highlights that the regeneration of large-seeded plants can be rendered vulnerable even by small imbalances between mutualistic and antagonistic interactions. In scenarios of increasing defaunation, the combination of reduced regeneration of vulnerable species and the dominance of resilient plant species can

29 radically change compositional patterns, reducing biodiversity and favoring a homogenization of Atlantic Forest fragments.

Escribano-Avila, G.1, Calviño-Cancela, M.2, Pías, B.3, Escudero, A.4 & Virgós, E.4 1Institute of Ecology, Technical Particular University of Loja, Ecuador, 2Ecology and Animal Biology Department, Vigo University, Spain, 3Plant Biology Department, Complutense University of Madrid, Spain, 4Biodiversity and Conservation Area, Rey Juan Carlos University, Spain [email protected] Functional complementarity of the dispersal community leads to forest recovery and expansion after land abandonment Land abandonment due to increasing depopulation of rural areas is an ongoing trend in developed countries worldwide. Abandoned lands represent an opportunity for ecosystem recovery in which seed dispersal provided by animals is a key feature. Different dispersers may differentially contribute to plant recruitment under varied ecological conditions, leading to complementary or redundant dispersal services. We studied seed dispersal effectiveness (recruited seedlings/m2) resulting from the dispersal activity of the main dispersers of the Spanish juniper (Juniperus thurifera L.) which are small- to medium-sized strongly frugivorous birds (thrushes) and medium- to large-sized carnivorous mammals (red fox and stone marten), in two ecological contexts derived from ecosystem recovery after a historical period of degradation: remnant forests and old fields. Results showed a clear shift in the contribution to plant recruitment between these guilds, resulting in complementary and non-redundant dispersal services. Thrushes were the main contributors to plant recruitment in forests (73%), leading to population growth but with a reduced impact on the colonization of old fields where carnivores contributed to 80% of recruitment (42% red fox, 38% stone marten). The combination of short-distance, strongly frugivorous dispersers (e.g. passerine birds) together with big generalist frugivores with long-distance movements (e.g. carnivorous mammals) maintained a) effective seed dispersal services in remnant forests, and b) the connectivity between patches promoting old field colonization and forest expansion. By performing effective dispersal services across an array of habitat types, functionally diverse disperser communities assist natural restoration of human-impacted ecosystems all over the world replacing costly human- mediated restoration projects.

Falcón, W.*1, Kaiser-Bunbury, C.2, Bunbury, N.3 & Hansen, D.1 1Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Switzerland, 2Department of Biology, Technical University of Darmstadt, Germany, 3Seychelles Island Foundation, La Ciotat Building, Mont Feluri, Mahé, Republic of Seychelles [email protected] Giants, birds, and bats: First approximation of the seed dispersal network of Aldabra Atoll Animal-mediated seed dispersal is an important ecological function in many ecosystems worldwide. Obtaining a detailed understanding of this process at a community level in species-rich ecosystems, based on experimental evaluations of all pairwise interactions, is a Herculean task. To reduce these complexities, ecologists have a long tradition of using isolated islands as field laboratories. We use the vertebrate frugivores and fleshy-fruited plants on Aldabra Atoll as a model system to investigate the seed dispersal network (SDN). Aldabra is one of the few islands with an intact frugivore fauna, spanning small lizards, pigeons, fruit bats, and tortoises. The atoll is the last place on Earth where giant tortoises occur in high densities. We employ three methodologies to construct a robustly informed SDN: observations, camera traps, and fecal analysis. We present preliminary network structure analyses revealing a significantly nested pattern in the SDN, compartmentalization, and asymmetries of interaction strengths. Blue pigeons and giant tortoises are frequent frugivores of the sampled plant species, and thus, potential drivers of SDN structure. In addition, one of the most connected plants is Solanum aldabrense, which encourages focusing on this plant to assess the effects of endozoochory on intra and inter-island dispersal by using genetic techniques. We are collecting more data to elucidate the structure of the SDN, and — focusing on giant tortoises — to assess effects

30 of gut passage on seed germination, and to predict animal-mediated seed deposition by constructing a mechanistic, individual-based simulation model.

Farwig, N., Berens, D.G. & Grass, I. University of Marburg, Faculty of Biology - Conservation Ecology, Germany [email protected] Guild-specific shifts in visitation rates of frugivores with habitat loss and plant invasion Habitat loss and plant invasions lead to a loss of biodiversity and alter species interactions, which may imperil vital ecosystem processes such as seed dispersal. Reponses of frugivores to disturbance are often linked to their specialization on certain habitats or resources. Yet, it is poorly understood how habitat loss and plant invasion structure interactions between plants and different habitat or feeding guilds. We therefore investigated whether visitation rates of frugivores change guild-specifically with increasing habitat loss and invasion level in a heterogeneous subtropical landscape. Visitation rates of forest specialists decreased with increasing habitat loss, but not with changes in invasion level. In contrast forest generalists and forest visitors were unaffected by either driver. Similarly, obligate frugivores that overall showed a generalized fruit choice were unaffected by habitat loss and changes in invasion level. On the contrary, visitation rates of specialized partial and opportunistic frugivores decreased with higher invasion level. Importantly, the negative effect of plant invasion on partial frugivores was more pronounced as habitat loss in the same study site increased, indicating a synergistic effect of the two drivers. First, frugivores respond guild-specifically to habitat loss and plant invasion. Consequently, forest dependency is mainly related to habitat loss, and degree of frugivory is mainly related to plant invasion. Forest generalists and obligate frugivores in turn may play a key role for forest regeneration in disturbed forest landscapes. Second, frugivores with a specialized fruit choice may be, in particular, threatened by synergistic effects between habitat loss and plant invasion.

Fedriani, J.M.1,2,3, Wiegand, T.1, Palomares, F.2 & Grimm, V.1 1Department of Ecological Modelling, Helmholtz Centre for Environmental Research GmbH – UFZ, Germany, 2Department of Conservation Biology, Estación Biológica de Doñana (EBD - C.S.I.C.), Spain, 3Technical University of Lisbon, Institute of Agronomy, Centre for Applied Ecology, Tapada da Ajuda, Portugal [email protected] Old-field recolonization under selective defaunation: a spatially-explicit individual-based simulation model Land use changes and defaunation are two major drivers of global change whose combined effect is poorly understood. In particular, increased levels of land abandonment worldwide are leading to noticeable changes in landscape cover. Whether and how old fields are colonized depend firstly on the arrival of seeds which, in turn, is contingent on the density and identity of seed vectors. In animal-dispersed plants, the strength and spatial pattern of the seed rain can be strongly impacted by selective defaunation. Our study site, located within the Doñana National Park (SW Spain), was used for intensive cow grazing until 1996, when the land was expropriated, the cows removed and, afterwards, protected by the National Park Service. Based on large datasets collected during the last two decades (concerning disperser movements, disperser fruit consumption rates, fruiting plant distribution, and crop sizes), we built an individual-based spatially explicit simulation model to evaluate the effect of rarefaction of different seed dispersers (frugivorous mammals) in the recolonization of this old field by the endozoochorous Iberian pear Pyrus bourgaeana. Simulation experiments indicate that selectivity of defaunation has a major effect on different traits of the seed dispersal kernel. Because seed disperser attraction to the old field can be readily manipulated through undemanding management, we performed similar simulation experiments and found that the level of attraction to old fields can strongly affect the pace of woody recolonization but in a manner rather different to the predicted one. Our study may have implications in mitigating some detrimental effects of global change.

Forget, P.-M., Albert, A., Boissier, O., Bouiges, A., Mendoza, I. & Feer, F. Muséum National d’Histoire Naturelle, France [email protected] Using trees and fruit ‘to take the temperature’ of rainforests: a study case along a gradient of anthropogenic disturbances in French Guiana Ecological processes in tropical forests are being affected at unprecedented rates by human activities. Yet, the continuity of ecological functions like seed dispersal is crucial for forest regeneration. Using a new method that we developed to rapidly assess the health status of the rainforest we studied the effect of anthropogenic impact on fruit consumption and seed removal of Virola michelii (Myristicaceae) between 2009–2015. The five chosen forest sites illustrate a gradient of habitat degradation in French Guiana. We counted fallen fruits, fruit valves, and seeds of each focal fruiting trees (N = 100) in 1 m2 quadrats, and calculated two indices: the proportion of fruits opened and the proportion of seeds removed by arboreal and terrestrial mammals. Our results showed that seed removal rates gradually decrease with increasing habitat degradation. Our indices also confirm that the activity of birds that cannot open maturing dehiscent fruit is positively related to increased global forest disturbances. Small body-sized consumers in the most impacted habitats are unlikely to efficiently compensate for the disappearance of the large- body sized frugivores and seed dispersers contributing to high density of undispersed seeds beneath parents. In addition to low seed removal, little post-dispersal seed predation by invertebrates and terrestrial frugivores exacerbates the overall saturation effect contributing to possible seedling recruitment beneath trees.

Fricke, E.C.*1, Tewksbury, J.J.1,2, Wandrag, E.M.3 & Haldre, S.3 1Department of Biology, University of Washington, USA, 2Luc Hoffmann Institute, WWF International, Gland, Switzerland, 3Department of Ecology and Evolutionary Biology, Rice University, USA [email protected] Mutualistic strategy tradeoffs determine seed dispersal network structure and increase robustness to co- extinction The structure of seed dispersal networks, and the position of species within those mutualistic networks, determines their robustness to co-extinction. Yet, the mechanisms currently used to explain network assembly and to predict co- extinction do not consider how species vary in their dependence on the mutualism for reproduction and survival. Rather, studies that assess network stability or predict co-extinctions usually assume that all species are obligate mutualists. For plant species in a highly resolved seed dispersal network, we show that a tradeoff between mutualistic and non-mutualistic strategies explains investment in and benefits of the mutualism, as well as the disperser diversity of plant species and dispersal frequency. Using estimates of the portion of fruit in bird diets, we show that birds present in empirical networks exhibit a positive relationship between the degree of frugivory and their partner diversity and interaction frequency. This tradeoff strongly increases network robustness to co- extinction, reducing co-extinctions simulated in empirical networks relative to predictions assuming obligate mutualists. The existence of the tradeoff — causing a positive relationship between intrinsic dependence on the mutualism and partner diversity and interaction frequency — confers greater robustness to co-extinction than do simple reductions in intrinsic dependence. Our results show that mutualistic strategy tradeoffs underlie the assembly of nested network architecture, reorient predictions for the species most vulnerable to co-extinction, and buffer seed dispersal networks from co-extinction.

García, D.1, Donoso, I.1, Schleuning, M.2,3 & Fründ, J.4 1Depto. Biología de Organismos y Sistemas, Universidad de Oviedo, Asturias, Spain, 2Biodiversity and Climate Research Centre (BiK-F), 3Senckenberg Gesellschaft für Naturforschung, Frankfurt (Main), Germany, 4Dept. of Integrative Biology, Univ. of Guelph, Canada [email protected] / [email protected]

Towards a functional view of seed dispersal networks: a framework based on seed disperser effectiveness and plant fate Seed dispersal (SD) networks are thought to represent the complexity of interactions between plants and frugivores. However, networks based on frequencies of seed removal may say little about the ultimate functional effects of seed dispersal. To overcome this limitation, seed fate (i.e. the probability that a seed recruits to a new individual plant) must be incorporated into SD networks. For that, both disperser and plant species traits are relevant. Depending on species traits, seed dispersers may differ in the quantity of seed removal and the quality of seed treatment and seed deposition, and plants may differ in demographic fate. We present a conceptual framework to predict changes in network structure from SD to recruitment, depending on the correlations between the quantitative and qualitative roles of frugivore and plant species. These correlations are contingent on species traits and environmental conditions. When correlations between quantity and quality are negative, interaction evenness (i.e. homogeneity of species’ contributions) is expected to increase from SD to recruitment networks. The sign of quantity–quality correlation may depend on the occurrence of trait-mediated trade-offs (e.g. body size and population size in dispersers, seed size and seed number in plants), disperser–plant trait matching (disperser body size–seed size) and the sign of the effects of plant traits (e.g. seed size) on seed fate. Different functional outcomes are expected for networks mostly shaped by the relative abundances of dispersers and plants compared to those mostly shaped by trait-matching between interaction partners.

García, D., Martínez, D., Donoso, I. & Rodríguez-Pérez, J. Depto. Biología de Organismos y Sistemas, Universidad de Oviedo, Asturias, Spain [email protected] / [email protected] The biodiversity–ecosystem function link in frugivory and seed dispersal: effects across dispersal components and scales The positive, causal link between biodiversity and ecosystem function is a current ecological paradigm, but it has been seldom tested in frugivory and seed dispersal. Here, we evaluate the relationship between frugivore biodiversity (abundance, richness, diversity) and the quantity and the quality of seed dispersal function. We focused on fleshy-fruited woody plants dispersed by frugivorous birds in the fragmented forests of the Cantabrian range (N Spain), covering different spatial scales and years. On a local scale, we found that the abundance and the richness of frugivorous thrushes affected positively the quantity (density of dispersed seeds) and the quality (spatial distribution of seeds, colonization of deforested habitat) of tree seed dispersal. Nevertheless, the abundance of thrushes mostly affected the quantity of seed dispersal, whereas the quality depended on frugivore richness. The shape of the relationship between frugivore richness and seed dispersal changed across years, from linear to quadratic. On a regional scale, the abundance of frugivores affected all the components of the seed dispersal of woody plants, and increases in frugivore diversity led to higher densities and wider spatial distributions of dispersed seeds. These regional effects were independent of habitat cover availability. Frugivore abundance and richness affected positively the quantity and the quality of seed dispersal function, but these effects depended on the dispersal component, as well as on the spatio-temporal scale. The link between frugivore diversity and seed dispersal may be explained by sampling effects, spatial complementarity between frugivores and inter-specific facilitation.

González-Varo, J. P. & Jordano, P. Estación Biológica de Doñana (EBD-CSIC) [email protected] Incorporating time into the seed dispersal effectiveness framework Since the development of the seed dispersal effectiveness (SDE = Quantity × Quality) framework, researchers have placed a huge emphasis on the spatial location of animal-dispersed seeds. The fates of seeds and seedlings often differ among microhabitats, and that is the reason why space has been a major target when evaluating the quality of seed dispersal services provided by animals. The quantitative component has been usually assessed by combining

33 information on microhabitat use by different animal vectors and the magnitude of seed arrival in different microhabitats. The qualitative component has been usually assessed by experimentally evaluating seed survival and seedling establishment in different microhabitats at a given time. However, the temporal dimension within the SDE framework has been virtually overlooked. The fruiting period of many (sub-) tropical fleshy-fruited plants can last for several months. Both biotic and abiotic factors affecting recruitment processes can strongly vary across the seasons. Thus, the timing of dispersal could be more decisive for recruitment than the seed deposition sites. Besides, the interactive effects ‘space × time’ could shift the relative quality of microhabitats across seasons. Furthermore, the assemblage of animal vectors can also be temporally structured, as many frugivores and seed dispersers are migratory animals. In this contribution, we incorporate time into the SDE framework, using as case study a Mediterranean shrub with a fruiting phenophase that lasts for seven months.

Govender, K.*, Shuttleworth, A., Downs, C.T. & Johnson S.D. School Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] The role of volatiles in the dispersal of fig seeds (Moraceae) by frugivorous bats in southern Africa Olfaction is a key sense for mammals and it has been shown that fruit-eating bats use olfactory cues during foraging. However, little has been documented about the volatiles used by bats to locate fruits and determine their palatability. Ripe figs (Ficus spp.) are an essential source of food for frugivorous bats and they in turn disperse fig seeds. This study aimed to determine the volatile organic compounds (VOCs) emitted by fig syconia (Moraceae) as they develop from unripe to ripe figs and to determine the responses of bats to these compounds. We found significant changes in VOCs in the scent of unripe, ripe and over-ripe fig fruits across day and night using the dynamic headspace sampling and gas chromatography-mass spectrometry. Spectral analysis also revealed colour changes as figs ripen and analysis using a generic animal colour vision model suggests that bats can perceive these changes. Field-based choice experiments indicated that Wahlberg’s Epauletted Fruit Bat (Epomophorus wahlbergi) prefer ripe fruits over green ones. These bats also responded to synthetic compounds matching those identified from headspace samples of different fig developmental stages. Determining the responses of fruit-eating bats to the volatiles and colour of fig fruits is important for understanding the evolution of fruit characteristics in Ficus and also for understanding the foraging behaviour and seed dispersal role of bats.

Green, R.J. Environmental Futures Research Institute, Griffith University, Brisbane, Australia [email protected] Popularity doesn't always lead to success: why aren't there more Ficus saplings? Field observations of Australian subtropical rainforest trees show relatively high visitation rates by a variety of potential avian dispersers to certain Ficus species. Ficus trees generally have very large crops, and each fig contains many seeds. Yet counts of saplings in natural and semi-cleared habitats indicate a very low percentage of Ficus relative to various other local fleshy-fruited species producing much smaller crops and visited by far fewer dispersers. The aim of this (on-going) research is to determine the necessary conditions for germination and persistence of local Ficus spp. by observations of conditions (e.g. % foliage cover, dominant ground cover) under which saplings are currently growing, comparison of the same with observations of where local avian dispersers sit when not feeding on fruit (and thus likely to deposit seed), and field and greenhouse experimental plantings. Germination trials indicated that many viable seeds pass through local bird species, and that Ficus seeds germinate very readily and a high percentage continue to survive under greenhouse conditions both in local soils and a commercial seed-raising mix, so lack of viability is unlikely as an explanation. Seeds of Ficus spp. and other local species (including genera such as Jagera, Breynia and Maclura) were planted in the field under a variety of treatments (e.g. shade versus no shade, protection from vertebrate and invertebrate folivores, competition with

34 grasses and weedy species) as part of an on-going experiment on conditions for germination and subsequent survival. Results so far suggest folivory and brief drought periods as major factors.

Hansen, D.1, Falcón, W.1 & Moll, D.2 1Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Switzerland, 2Biology Department, Missouri State University, Springfield, USA [email protected] Frugivory and seed dispersal by chelonians: A systematic review and synthesis Frugivory and seed dispersal (FSD) has been much studied in most of the major vertebrate taxa, with a chief focus on birds and mammals. However, in recent years it has become clear that FSD by chelonians—turtles and tortoises—is much more common than previously thought. Yet, a systematic review and synthesis is lacking. Here, we present the results of a systematic review of published and unpublished records of chelonian FSD. A substantial proportion of the world’s aquatic and terrestrial turtles and a major part of testudinid tortoises include fruits in their diet. For some species, overall or in certain seasons, fruit may even form the largest part of their diet. Most importantly, contrary to the other major reptilian seed dispersers, i.e. lizards, chelonian FSD is not an island phenomenon in terms of geographic distribution. Nevertheless, on islands especially, tortoises are often among the largest native terrestrial vertebrates—or were, until humans got there. We synthesize the lessons learned so far, and discuss 1) fruitful avenues for future chelonian FSD research, and 2) the relevance of our findings for conservation and restoration, especially in relation to rewilding with large and giant tortoises.

Hart, L.A.*, Johnson, S.D., Downs, C.T. DST-NRF Centre for Invasion Biology, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] The effect of Wahlberg's Epauletted Fruit Bats (Epomophorus wahlbergi) on alien invasive seed germination and their potential as dispersal agents The spread of invasive alien plants into natural habitats is of growing global concern. Several studies have investigated the role that avian frugivores play in the dispersal of these seeds and their effects on germination success. Fruit bats have however received little attention as important dispersal agents of invasive alien plants, despite their recognized role as long distance dispersal agents of various native flora. We investigated whether Wahlberg’s Epauletted Fruit Bats, Epomophorus wahlbergi, would positively influence the germination of seeds of invasive alien plants. These fruit bats were fed fruits of four invasive alien plant species, i.e. Psidium guajava, Melia azedarach, Eriobotrya japonica, and Morus alba. Epomophorus wahlbergi were able to process more fruit per gram body mass than birds. Spat and de-pulped control seeds had similar germination success and germinated at approximately the same time for most species. While seeds retained in whole fruit had significantly less germination success than spat seeds for all species, except M. azedarach, they mostly germinated at approximately the same time. Epomophorus wahlbergi can swallow small seeds (<2 mm), while seeds larger than this are generally spat out. Large fruit are usually carried away to feeding roosts where seeds are dropped, thereby dispersing seeds and fruits which are too large for some bird species to ingest. Epomophorus wahlbergi should not be underestimated as dispersers of these invasive alien plants as they consume proportionally large amounts (0.62 ± 0.09 to 0.99 ± 0.11 g-1 body mass) of fruit, except for M. azedarach, and affect their seed germination rates positively.

Haurez, B.1,2*, Petre, C-A.1,3,4, & Doucet, J-L.1,2 1Laboratory of Tropical Forestry, University of Lièg, Belgium, 2École Régionale Post-Universitaire d’Aménagement et de Gestion Intégrés des Forêts et Territoires Tropicaux (ERAIFT), Kinshasa, Democratic Republic of Congo, 3Centre for Research and Conservation, Royal Zoological Society of Antwerp (RZSA) - Belgium, 4Conservation Biology Unit - Section Education and Nature - Royal Belgian Institute of Natural Sciences, Belgium

35 [email protected] Western lowland gorilla seed dispersal: effect of fecal matrix and deposition at nest sites on seedling growth The Western lowland gorilla is a key dispersal agent that disperses viable seeds of various size. In consequence of its nesting behavior, the gorilla deposits half of its feces, and therefore about half of dispersed seeds, at nest sites. This study was conducted in a logging concession of southeast Gabon where 78 % of the observed nest sites (n=183) were installed in open canopy terra firma forest. The aim of this research was to assess if seed deposition by gorillas 1) in fecal matrix and 2) at nest sites is advantageous for seedling growth. To assess the effect of fecal matrix, seeds of Santiria trimera (Burseraceae), Chrysophyllum lacourtianum (Sapotaceae) and Plagiostyles africana (Euphorbiaceae) collected in gorilla feces were sown with and without fecal matrix in a nursery at the study site. The impact of seed deposition at nest sites on seedling growth was evaluated in situ. Seedlings of Santiria trimera and Dacryodes normandii (Burseraceae) were planted at nest sites and at closed canopy terra firma forest sites. For both type of trials, seedling growth was monitored. A manuring effect of fecal matrix on seedling growth was only observed for Plagiostyles africana. A positive effect of deposition at nest sites on seedling growth was observed, mostly explained by a more open canopy at nest sites. Seedling growth was positively correlated to canopy openness for both tested species. Therefore, the gorilla provides directed dispersal by depositing seeds most frequently in open canopy sites. Its role in forest regeneration is thought to be essential.

Ramanantoanina, A.1,2, Ouhinou, A.3 & Hui, C.1,2 1Centre for Invasion Biology, Department of Mathematical Sciences, Stellenbosch University, South Africa, 2Mathematical and Physical Biosciences, African Institute for Mathematical Sciences, South Africa, 3Department of Mathematics, Faculty of Sciences and Technology, University of Sultan Moulay Slimane, Beni-Mellal, Morocco [email protected] Spatial assortment of mixed propagules explains the acceleration of range expansion The ability to estimate the rate of spread of an invasive species is important for the success of its management and control. Range expansion of spreading organisms has been found to follow three types: (1) linear expansion with a constant rate of spread, (2) bi-phase expansion with a faster linear expansion following a slower linear expansion, and (3) accelerating expansion with a continuously increasing rate of spread. To date, no overarching formula exists that can be applied to all three types of range expansion. Early theories suggested that the velocity at which a species expands its range depends on the population growth and dispersal rates. Models based on partial differential equations, specifically the reaction-diffusion model, assume a normal distribution of species’ dispersal distances (i.e. dispersal kernel) and yield a widely used formula which depicts a constant rate of spread (c = 2(rD)1/2, where r and D denote the intrinsic growth and diffusion rates, respectively). However, a growing body of evidence suggests that the rate of spread for most species may not be constant. We investigated how propagule pressure, i.e. the initial number of individuals and their composition in terms of dispersal ability, affects the spread of a population. A system of integrodifference equations was then used to model the spatiotemporal dynamics of the population. We studied the dynamics of dispersal ability as well as the instantaneous and asymptotic rate of spread. We found that individuals with different dispersal abilities were spatially sorted with the stronger dispersers situated at the expanding range front, causing the velocity of expansion to accelerate. The instantaneous rate of spread was found to be fully determined by the growth and dispersal abilities of the population at the advancing edge of the invasion. We derived a formula for the asymptotic rate of spread under different scenarios of propagule pressure. The results suggest that data collected from the core of the invasion may underestimate the spreading rate of the population. Aside from better managing of invasive species, the derived formula could conceivably also be applied to conservation management of relocated, endangered or extra-limital species.

Zhang, F., Minoarevilo, H.O. & Hui, C. 1Centre for Invasion Biology, Department of Mathematical Sciences, Stellenbosch University, South Africa; 2Department of Mathematics, Gansu Agricultural University, China, 3Theoretical Ecology Group, African Institute for Mathematical Sciences, South Africa [email protected] Adaptive interaction switching and phylogeny explains the architecture of mutualistic networks Ecological interaction networks, such as those describing the mutualistic interactions between plants and their frugivores, exhibit non-random structural properties that cannot be explained by simple models of network formation. Nested architecture is distinctive in plant–animal mutualistic networks. However, to date an integrative and quantitative explanation has been lacking. It is evident that species often switch their interactive partners in real world mutualistic networks. By incorporating an interaction switch into a novel multi-population model, we show that the nested architecture rapidly emerges from an initially random network. Another factor affecting the formation and eventual structure of such a network is its evolutionary history. Empirical studies of interaction networks along with the phylogenies of the interacting species have demonstrated significant associations between phylogeny and network structure. We present a model describing the evolution of pairwise interactions as a branching Markov process, drawing on phylogenetic models of molecular evolution. The model allowing interaction switches between partner species produced predictions which fit remarkably well with observations from 81 empirical networks. Thus, the nested architecture in mutualistic networks could be an intrinsic physical structure of dynamic networks and the interaction switch is likely a key ecological process that results in nestedness of real-world networks. In contrast, the phylogenetic model yielded a significantly better fit to 21% of real-world mutualistic networks, a substantial minority of cases. Adaptive interaction switching and coevolutionary history, together, could be the driver behind network emergence.

Midgley, J.J.1, White J.1, Johnson S.D.2 & Bronner G.1 1Department of Biological Sciences, University of Cape Town, South Africa, 2DST-NRF Centre for Invasion Biology, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] Faecal mimicry by seeds for dispersal by dung beetles Mimicry, deception and sensory exploitation are controversial in plants, especially for seed dispersal. We investigated the dispersal of Ceratocaryum argenteum, an endemic Restionaceae plant species with particularly large round brown nuts. Large nuts in the Cape fynbos are often scatterhoarded by indigenous small mammals, but those of C. argenteum are unusual in having a strong foetid odour. We ruled out scatterhoarding of nuts of C. argenteum and show that they are dispersed by native Epirinus flagellatus dung beetles. The nuts emit an odour chemically similar to that of antelope droppings, particularly bontebok and eland which both occur naturally with C. argenteum plants. In conclusion the C. argenteum-E. flagellaris interaction is a case of seed dispersal deception and dung mimicry. The rewardless seeds of C. argenteum successfully exploit the visual and scent sensory perception of the dung beetle to ensure dispersal and burial.

Jones, L.R.*1, Johnson, D.M.2, Duke-Sylvester, S.M.1 & Leberg, P.L.1 1University of Louisiana Lafayette, United States, 2Virginia Commonwealth University, United States [email protected] Animal traits mitigating the negative consequences of fragmentation on spatial patterns of long-distance and contagious seed dispersal Large animals are thought to be high quality seed dispersers because they move long distances and exhibit high seed retention times, providing long-distance seed dispersal and potentially reducing clumped (contagious) spatial patterns. However, large animals often decline or disappear in the face of habitat loss and fragmentation. In their absence, some traits of smaller dispersers may produce similar spatial patterns of seed dispersal. We tested the

37 hypotheses that increasing habitat fragmentation decreases long-distance and increases clumped seed dispersal. We also hypothesized that high levels of some animal traits, specifically movement distance, seed retention times, and movement frequency, can mitigate these negative consequences of fragmentation. To test these hypotheses, we constructed an individual-based mathematical model in R and created simulations of an animal dispersing seeds in a theoretical forest landscape of 0–90% habitat loss across a wide range of values of the above animal traits. Increasing fragmentation decreased long-distance seed dispersal by up to 22% and increased clumping by up to 41%, supporting our hypotheses on fragmentation. Among mitigating variables, increasing animal movement distance resulted in moving seeds up to three times further from the parent plant and other seeds than increasing movement frequency or seed retention times. However, at low animal movement distances, intermediate and high values of movement frequency and seed retention times improved long-distance seed dispersal and reduced clumping beyond critical thresholds. In the absence of large dispersers that move long distances, smaller animals retaining seeds longer and moving frequently may also mitigate the negative consequences of fragmentation.

Kaplin, B.A.1, Martino, R.1 & Cordeiro, N.2 1Antioch University New England, NH, USA, 2The Field Museum, Chicago, IL, USA [email protected] Matrix effects on frugivory and seed dispersal in tropical montane forest: land use type in the matrix matters Frugivory and seed dispersal in tropical forest can be affected by matrix type and edge effects, yet little research has explored how matrix affects ecological processes such as seed dispersal. Our aim was to examine how matrix type (tea plantations (hard) vs. pine plantations (soft)) affects seed dispersal effectiveness and vegetation structure and composition in adjacent montane tropical forest. Our hypothesis was that soft or low contrast matrix types will better support ecological processes and biodiversity than hard, high contrast matrix. Research was conducted in Nyungwe National Park, Rwanda, in the Albertine Rift, a biodiversity hotspot. Over a 12-month period, we conducted focal tree observations and sampled vegetation. We found significant differences in frugivore assemblages and forest structure adjacent to hard vs. soft matrix types. Large birds made more visits to focal trees and removed more fruit in forest edges adjacent to soft, low contrast pine plantation, while primates made significantly more visits to focal trees and removed more fruit in forest edge adjacent to hard, high contrast tea plantations. Forest adjacent to tea plantations showed a deeper penetration of edge effects resulting in a greater proportion of small seeded, shade intolerant tree species than in edges adjacent to pine plantations. These findings demonstrate how land use around tropical forests affects seed dispersal processes and forest composition, and how matrix can contribute to ecological integrity in systems facing increasing pressures from human land use in the surrounding matrix.

Kelly, D., Macfarlane, A.F. & Briskie, J.V. Biological Sciences, University of Canterbury, Christchurch, New Zealand [email protected] European blackbirds (Turdus merula) in New Zealand: useful substitutes for lost mid-sized native frugivores, or weed vectors? There is worldwide interest in how exotic frugivores integrate into fruit dispersal networks in their introduced range. The New Zealand avifauna has lost numerous bird species, leaving few birds capable of dispersing large fruits. In theory, the relatively large-bodied introduced European blackbird (Turdus merula) and song thrush (T. philomelos) could function as replacement dispersers for larger fruits. However, there is also concern that they may serve to disperse exotic woody weeds, perhaps more so than endemic and native birds. Recent studies have come to opposite conclusions about the numerical contribution of blackbirds to the dispersal of native fruits in New Zealand: a very small contribution in one study (Kelly et al. 2006) but the second largest contribution in a second study (Burns 2012). Here we identify seeds from faecal samples gathered during mist-netting in Kowhai Bush (South Island) to test whether blackbirds and song thrushes cover the large-native-seed disperser gap or just move weeds. We also review

38 data on blackbird densities in various habitats to see if differences in the density of blackbirds can account for the differences in blackbird importance in the different studies. We conclude that blackbirds are important dispersers of native seeds but differences in their importance reflect habitat differences, with these birds being more important in urban areas. In Kowhai Bush, the main contribution of blackbirds and song thrushes which was not already well covered by native bird species was moving large-seeded woody weeds.

Kitamura, S. & Sawada, M. Ishikawa Prefectural University, Japan [email protected] Dispersal of Ginkgo biloba (Ginkgoaceae), a large-seeded tree species, in a temperate forest in Japan Numerous plants have seed-dispersal mechanisms that are associated with animals. We investigated the dispersal of a large-seeded tree species, Ginkgo biloba, in a temperate forest of Ishikawa Forest Experiment Station in Japan. Although its smelly seeds are one of its most well-known and distinctive features, we know very little about how seed dispersal works in living ginkgo, Ginkgo biloba (Ginkgoaceae). By using camera-trapping observations of fallen seed consumption on the ground, we identified the frugivore assemblage that foraged on the seeds of G. biloba and assessed their role in seed dispersal and seed predation. Our results showed that the frugivores dispersing seeds were carnivorous mammals, including Raccoon Dog Nyctereutes procyonoides, Eurasian Badger Meles anakuma, Masked Palm Civet Paguma larvata, and Asiatic Black Bear Ursus thibetanus, whereas the large Japanese field mouse Apodemus speciosus predated most seeds, but scatter-hoarded some on the ground. High seed removal rates and defecation of intact seeds by these carnivorous mammals lead us to the conclusion that they show high effectiveness, in terms of quantity, in dispersal of this tree species. Ginkgo biloba seeds are confirmed to adapt primarily to mammalian endozoochory, a mutualistic association similar in function to fleshy pulp.

Kromann-Clausen, A.1,2, Dabelsteen, T.1 & Campos-Arceiz, A.2 1Department of Biology University of Copenhagen, Copenhagen, Denmark, 2School of Geography, University of Nottingham Malaysia Campus, Selangor, Malaysia [email protected] Megafauna-caused differences in tree distribution between mineral licks and random locations in a Malaysian forest Tropical rainforest landscapes contain resource patches distributed in a heterogeneous manner in both time and space, from fruiting trees available frequently in the short term, to rarer mineral deposits with long term availability. Asian elephants (Elephas maximus) utilize these deposits to supplement their diet with minerals. Due to their rarity, mineral licks attract elephants from a greater area than other resource patches (e.g. food). Elephants are effective seed dispersers due to their size and ability to ingest seeds that other species are incapable of. We expect the rare megafauna-syndrome tree species (i.e. those with large seeds and fruits) to be more common at mineral licks frequently visited by elephants. In northern Peninsular Malaysia, we compared the difference in abundance of six species, with varying dependence on elephants for dispersal, at mineral licks and similar random locations. At each site we compared tree abundance in elephant-made trails, and non-trail transects. We sampled eight 500 m transects (4 trails and 4 non-trails) per site at a total of 15 sites (8 mineral licks and 7 random locations). We found that significantly more megafaunal-syndrome trees could be found at mineral deposits (df=1, F-value = 11,9790, p= 0.0005), but there were not significantly more trees at trails compared to non-trail areas at mineral deposits (df = 1, F-value = 0,0926, p = 1.24929). Our results indicate that the search behavior by megafauna for rare nutritional resources has great implications for forest structure, tree distribution, and forest conservation in Southeast Asia. It is therefore important to take mineral deposits into consideration in forest conservation.

Kwit, C., Patton, M.T. & Miller, C.N. University of Tennessee, USA [email protected] Plant advantages in myrmecochory: redirecting ‘directed dispersal’ Myrmecochory, the dispersal of elaiosome-bearing seeds by ants, is typically cast as a mutualistic relationship. Benefits of this putative mutualism include nutritive rewards for in-nest ant larvae, and safe and nutrient-rich microsites conferring enhanced seed survival, germination and seedling establishment. To evaluate this from the plant perspective, we conducted a literature review and found 1) natural history-related findings noting the prevalence of re-dispersal of seeds out of ant nests, 2) lack of consensus on effects of elaiosome removal on seed predation and germination, 3) lack of research on ant-produced antimicrobial secretions within nests, and 4) a mammal-centric approach to myrmecochorous seed mortality. We argue that this ant–plant interaction is improperly cast as a prime example of ‘directed dispersal’ and pose a more relevant framework to test whether ants are critically important dispersers in such systems. This should include characterization of relevant abiotic and biotic metrics in areas near ant nests, and a better incorporation of microbial seed depredation and the effectiveness of ant seed treatments. Thorough tests of ‘directed dispersal’ in myrmecochorous systems should include a combination of empirical, experimental, and modeling efforts better documenting the dynamics of myrmecochorous seeds and their microbiota through the dispersal process. Such approaches are applicable in other systems exhibiting disperser seed-treatment, and will transform the conceptual framework of ‘directed dispersal.’

Luna, C.A.1, Loayza, A.P.1 & Squeo, F.A.1,2 1Universidad de La Serena, Instituto de Ecología y Biodiversidad (IEB), Chile, 2Centro de Estudios Avanzados en Zonas Áridas (CEAZA), Chile [email protected] Intraspecific variation in seed size affects the outcome of the interaction between scatter-hoarding rodents and a threatened Atacama Desert shrub Myrcianthes coquimbensis is a large-seeded, fleshy-fruited shrub with no present-day dispersers, its only vectors of seed dispersal are three species of scatter-hoarding rodents: Octogon degus, Phyllotis darwini and Abrothrix olivaceus. These rodents forage M. coquimbensis seeds ranging from 0.5 to 15 g, and frequently only partially consume them, discarding seed fragments at the hoarding sites. We assessed whether the role of rodents as predators or dispersers depends on the size of seeds. Additionally, we examined whether seed size affected emergence probabilities of seeds with simulated partial predation. For all rodent species, seed size was positively related to the probability of being hoarded. Hoarded seeds were three times larger than non-hoarded ones, therefore, large seeds were more frequently dispersed. Additionally, smaller seeds had higher probabilities of in situ consumption. Phyllotis darwini and A. olivaceus hoarded bigger seeds than O. degus, the larger rodent. The latter also consumed more seeds in situ than the smaller species, and was more likely to consume hoarded seeds, thus, compared to the other two species, it acts more as a predator than a disperser. Seed size influenced the probability of emergence from seeds with simulated partial predation: larger seeds were more likely to emerge than smaller ones. We conclude that scatter-hoarding rodents are selecting for large seeds. Moreover, large seeds are more likely to recruit after partial consumption than smaller ones, which may explain the maintenance of the large seed size of M. coquimbensis in a desert ecosystem, where having large, recalcitrant seeds would otherwise reduce recruitment probabilities.

McConkey, K.R. National Institute of Advanced Studies, Indian Institute of Science Campus, Bangalore, India [email protected] Not all frugivores are created equal: identifying the most effective seed dispersers in an Asian rain forest

Frugivores clearly differ in their respective abilities at dispersing seeds and researchers often make assumptions of disperser effectiveness based on observations of fruit and seed handling. From this animal-based perspective, gibbons (small Asian apes) are considered to be among the most important seed dispersers in tropical Asian rainforests. Gibbons are generalist frugivores, eating a wide range of fruit-types and dispersing seeds of most species intact in their scats. However, gibbons are often unable to consume the quantity of fruits that more abundant (but usually more wasteful) frugivores, such as macaques and squirrels, can. Without an understanding of disperser effectiveness from a plant perspective it is not clear whether having high quality dispersers (such as the gibbon) is ultimately more important than having high quantity dispersers (such as the macaque). We assessed the comparative effectiveness of gibbons and other frugivores in the seed dispersal of 10 plant species that feature commonly in the gibbon diet. We measured seed dispersal effectiveness by evaluating the early recruitment dynamics, beginning with fruit consumption through to one- or two-year seedling survival. Our results provide interesting insights into the relationships between high-quality seed dispersal vs. high-quantity dispersal and the relative importance of post-dispersal events.

Mokotjomela, T.M.*1,2, Hoffmann, J.H.2 & Downs C.T.3 1University of the Witwatersrand, South Africa, 2Department of Biological Sciences, University of Cape Town, Rondebosch, South Africa, 3School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] The potential for birds to disperse the seeds of Acacia cyclops, an invasive alien plant in South Africa Rooikrans Acacia cyclops is an aggressive invasive tree that threatens natural resources in South Africa. The seeds of A. cyclops have a prominent aril which attracts birds that ingest the seeds and disperse them endozoochorously. Two biological control agents, a Seed Weevil Melanterius servulus and a Flower-galling Midge Dasineura dielsi, were released on A. cyclops in 1991 and 2002, respectively. Together these agents have substantially reduced seed production and generally far lower numbers of seeds are now available to birds. A consequence of this transition from historically bounteous quantities of seeds to scanty seed availability is that birds may no longer associate with the trees and seed dispersal may be disproportionately reduced. To assess whether this has happened, seed attrition was measured by comparing the amount of seeds that disappeared from two groups of branches, one available to birds and the other enclosed in bird netting. Other types of granivores (mainly field mice) were excluded from both groups of branches with a plastic funnel placed around the stems. Mature seeds were also harvested and fed to caged bird species to determine gut retention times and germination rates of ingested seeds. Attrition rates of seeds showed that birds continue to remove seeds but that only a proportion of the crop is taken. Only two frugivorous species (Knysna Turaco Tauraco corythaix and Red-winged Starling Onychognathus morio) and two granivorous species (Red-eyed Dove Streptopelia semitorquata and Laughing Dove Streptopelia sengalensis) ingested A. cyclops seeds during feeding trials. Ingestion by birds enhanced seed germination except for those ingested by Laughing Doves. There were no apparent effects of length of gut passage time and avian body size on seed germination rates. Despite the diminished seed resource due to biological control agents, birds continue to disperse A. cyclops seeds.

Moran, C.1,2,3, Freeman, A.N.D.4, Catterall, C.P.1, Elgar A.T.1, Freebody, K.5, Montenero, M.4, Shoo, L.P.6 1Environmental Futures Research Institute, School of Environment, Griffith University, Brisbane, Australia, 2College of Marine and Environmental Sciences, James Cook University, Cairns, Australia, 3Biodiversity and Ecosystem Knowledge Services, CSIRO, Atherton, Australia,4The School for Field Studies, Centre for Rainforest Studies, Yungaburra, Australia; 5Tablelands Community Revegetation Unit, Tablelands Regional Council, Malanda, Australia, 6School of Biological Sciences, The University of Queensland, St Lucia, Australia [email protected] Overcoming barriers to seed dispersal and seedling recruitment in retired tropical pasture While restoration of tropical forests on cleared land is widely considered to be important for conservation, a major challenge lies in achieving restoration at scales that are ecologically meaningful. This is largely due to the high costs

41 of directly replanting cleared areas. This work aims to understand whether natural seed dispersal and forest regeneration processes can be facilitated by installing bird-attracting features in abandoned pasture in tropical Australia. We established an experimental trial using three, 0.64 ha plots, each with nine artificial bird-attracting perches, and nine water troughs which functioned as bird attractants as well as seed-collecting traps. We also controlled pasture grasses in experimental plots. We monitored use of plots by seed dispersing birds, and measured seed rain and seedling recruitment around perches over two years. A subset of the regional seed-dispersing bird assemblage was observed using plots; three of these are known from earlier work to have a high potential to disperse rainforest seeds, including large seeds (>10mm). All of the seeds captured in seed rain beneath perches are known to be bird-dispersed. Here, we examine links between the seed-dispersing bird assemblage, and the composition of the seed rain and seedling recruitment around artificial perches. We discuss the potential to facilitate bird-mediated seed dispersal in assisted regeneration designs.

Mueller, T.1, Lenz, J.1, Caprano, T.1, Fiedler, W.2 & Böhning-Gaese, K.1 1Senckenberg Biodiversity and Climate Research Centre, Frankfurt (Main), Germany, 2Max-Planck Institute for Ornithology, Germany [email protected] Bird movements reveal seed dispersal networks in fragmented landscapes Spatially explicit mapping of ecosystem functions remains a challenge and methodological hurdles have limited our understanding of dispersal pathways on the landscape scale. We demonstrate how movement data of frugivores can be used for spatially explicit mapping of seed dispersal networks in fragmented landscapes. We combined movement data of 30 Trumpeter Hornbills (Bycanistes bucinator) with gut-passage times and high-resolution habitat data in a fragmented forest landscape in South Africa. We identified potential seed dispersal paths and distinguished whether potential seed transport happened between forest patches, within the same patch, or into the habitat matrix. To quantify functional landscape connectivity we identified all possible between-patch connections and used graph networks to estimate landscape connectivity provided by hornbills. Although potential between-patch dispersal events were rare (on average 7% of all dispersal paths), hornbills could cover distances of up to 15 km. Hornbills visited over 100 forest patches and connected a habitat network with an extent of about 50 km which increased the potential functional connectivity of the landscape more than two-fold. We identified habitat patches that were critical stepping stones for seed dispersal pathways. Without these stepping-stones the network would likely disintegrate into separated components and lead to isolation of forest fragments. We showed that large frugivorous birds can greatly improve functional connectivity for fleshy-fruited plants across broad scales. Combining high-resolution movement and landscape data into graph networks allows the identification of seed dispersal pathways and critical stepping stones in fragmented landscapes, which could be widely incorporated in reserve design and landscape-level conservation planning.

Neuschulz, E.L.1, Mueller, T.1,3, Bollmann, K.2, Gugerli, F.2 & Böhning-Gaese, K.1,3 1Senckenberg Biodiversity and Climate Research Centre (BiK-F), Frankfurt (Main), Germany, 2Swiss Federal Research Institute for Forest, Snow and Landscape Research (WSL), Birmensdorf, Switzerland, 3Department of Biological Sciences, Johann Wolfgang Goethe-Universität Frankfurt, Frankfurt (Main), Germany [email protected] Seed dispersal by a food-hoarding bird counteracts plant establishment Seed-hoarding by animals is an important mechanism for plant regeneration. The storage of seeds at sites with specific environmental conditions is, however, only little understood. One explanation for such a behavior is the avoidance of pilferage by other animals. Another hypothesis is that animals choose locations that hamper the perishability of stored food, allowing the consumption of unspoiled food items over long time periods. We tested whether pilferage avoidance or seed perishability drives the selection of caching sites of Spotted Nutcrackers (Nucifraga caryocatactes). We studied the caching behavior of Spotted Nutcrackers in the Swiss Alps that store seeds of Swiss Stone Pine (Pinus cembra). We observed seed deposition by nutcrackers for three years across a 500 m

42 elevational gradient. We also experimentally deployed seed caches to unveil seed pilferage and seedling establishment. We used seedling establishment as an inverse measure of seed perishability, since established seedlings can no longer be consumed by nutcrackers. We found a significant mismatch in environmental conditions between microsites of high seedling establishment and microsite of seed caching. Caching by nutcrackers decreased, but seedling establishment increased, with increasing canopy openness and vegetation cover. Thus, nutcrackers cached seeds at sites of low seed perishability. We could not relate caching behaviour to pilferage-avoidance. Our study highlights the importance of seed perishability as a mechanism for seed-caching behaviour. The regeneration potential of plants dispersed by seed-caching animals may strongly decrease if animals cache seeds at sites that favour seed perishability rather than seedling establishment.

Niebuhr, B.S.*1 & Campos, R.P.2 1Spatial Ecology and Conservation Lab, Universidade Estadual Paulista - UNESP Rio Claro, Brazil, 2Laboratory of Plant Ecology, Universidade Federal do Paraná, Curitiba, Brazil [email protected] Effects of animal movement and space on interaction networks in different successional stages in tropical rainforest In tropical forests, ecological processes such as frugivory and seed dispersal are greatly affected by forest successional stage. In order to understand mechanistically the differences in species interactions, one must look at species richness, abundance, and composition, but also at the spatial distribution of fruit plants and at behavioral and movement features of frugivorous animals. Here we developed a spatially explicit stochastic agent-based model to simulate animal movement and frugivory process and recreate mutualistic bird–plant interaction networks. We used data on community structure, bird behavior, and interactions in the Brazilian Atlantic rainforest to parameterize the model and to explore the mechanisms behind the effects of forest successional stage on network properties. Network metrics derived from simulations were related to spatial structure, animal mobility, and behavior through multiple regression. We found out that network connectance, modularity, and evenness changed with successional stage, but nestedness showed little variation, in comparison to random expectations. In more advanced stages, plant individuals of the same species are likely to be more aggregated, and animal mobility variation tends to increase, which enhances network connectance and modularity. Our approach shows that forest successional stage affects mutualistic networks mainly through animal movement and the spatial configuration of plant species, but frugivore preference and morphological trait match also play a role in the structure of interaction networks.

Pedrosa, F.* & Galetti, M. Departamento de Ecologia, Universidade Estadual Paulista (UNESP), Brazil [email protected] Feral pigs as taxon substitutes: re-establishing lost ecosystem functions in defaunated landscapes. Recent invasion of feral pigs in two Brazilian hotspots of biodiversity challenges the assumption that it is an ecological disaster. While ecologists are witnessing the erosion of seed dispersal interactions worldwide due to disappearance of frugivores, there is an increasing recognition that alien species, in some cases, might restore species interactions. By evaluating stomach contents and faecal samples, we found that feral pigs consume fruits and disperse intact seeds, of a variety of native and alien species. We also evaluated two aspects of dispersal effectiveness: fruit removal and seed germination rate. Employing camera traps in front of large- seeded trees, we could estimate the contribution of feral pigs in removing fruits compared to native frugivores in three distinct sites. The action of feral pigs equals and exceeds the rate of seed removal by native large mammals in locations where the latter were historically extirpated, sites lacking both large native frugivores and feral pigs have undispersed fruits rotting below mother trees. We conducted seed germination tests by offering fruits to captive feral pigs and checking for seeds in their faeces. Defecated seeds germinated faster and at higher rates than those manually

43 pulped. Therefore, the results point to the role of feral pigs as effective seed dispersal agents. Indeed, they might be functioning as important replacements of lost ecological function in defaunated landscapes.

Pizo, M.A.1, da Silva, F.R.2 & Jacomassa, F.A.F.1 1Departamento de Zoologia, UNESP, Rio Claro, SP, Brazil, 2Programa de Pós-graduação em Biologia Vegetal, UNICAMP, SP, Brazil [email protected] Role of birds and bats in restored seed dispersal networks Seed dispersal is a key process in the restoration of ecosystem functionality, the main goal of ecological restoration. In tropical restored forests, bats and birds are the most important seed dispersers. We contrasted the roles of birds and bats in the seed dispersal networks formed in three restored forests with 15, 25 and 57 years since restoration, all planted with a high diversity of native tree species in the Brazilian Atlantic forest. From 2011 to 2013 we sampled feces collected from mist-netted birds and bats, and made 430 hours of direct observations on bird-dispersed fruiting plants to build seed dispersal networks for each study area. The networks were composed by 44 animals (9 bats and 36 birds) and 64 plants. Only 4 plants were used by both birds and bats. Bats dispersed more seeds than birds (26659 and 3035 respectively), however, birds showed more seed species richness (41 bird plants vs. 28 bat plants). Connectance was higher in bat- (0.23) than bird-networks (0.12). There was a higher network specialization in the former. The overall network was modular, also with a high specialization. High specialization is associated with a higher consumer diversity and more coexisting species. Bats dispersed pioneer tree species with small seeds (usually < 3 mm length), while birds dispersed both pioneer and later-successional species with larger seeds on average. Resource partitioning between the two disperser groups indicates that ecosystem function is being restored. The formation of well-defined subgroups or modules is important for the recovery of ecosystem functionality.

Prasad, S.1 & Westcott, D.A.2 1Jawaharlal Nehru University, New Delhi, India, 2CSIRO, Atherton, Australia [email protected] Developing a generic functional classification for seed dispersal A major limitation to improving predictions of plant movement in response to climate change has been realistic estimates of seed dispersal distances for plant communities. Mechanistic models are difficult to implement to derive generalizations for endozoochory unlike for anemochory. Functional classification of disperser services provides a useful tool to simplify characterization of endozoochory at community scales. Our aim was to develop a generic disperser functional classification for endozoochory, including communities with limited data and in heterogenous human-modified landscapes. We developed indices and a classification algorithm to group frugivores based on seed processing, quantity of seeds dispersed, seed dispersal distances, habitat specificity and range of fruits processed, and tested them on datasets from two communities with varying data availability. Our disperser effectiveness indices are focussed on the dispersal phase, and incorporate seed dispersal distances to classify disperser function. Our indices and functional classification offer flexibility to incorporate different data types that are used to quantify seed dispersal networks. We also provide tools to fine-tune the classification algorithm to meet the requirements of the research question. Our generic disperser functional algorithm and the associated indices provide flexible tools for various applications including comparison of dispersal services across disturbance gradients, and to derive estimates at the community level for seed dispersal kernels and quantity of seeds dispersed. As demonstrated by the applications with two tropical datasets, the generic disperser functional classification allows us to assess and compare the functional status of communities with respect to dispersal services.

Ramos, D.C.T.L.*1 & Silva, W.R.2 1Department of Ecology, São Paulo State University (UNESP), Rio Claro, Brazil, 2Department of Animal Biology, Campinas State University (UNICAMP), Campinas, Brazil [email protected] Vegetation functional and structural traits shaping seed rain underneath isolated trees Resources necessary for forest regeneration are usually depleted in disturbed sites, where seed dispersal and seedling establishment are also limited. Isolated trees play a fundamental role in these sites by concentrating seed rain in micro-sites favorable to native species establishment. This function depends on tree traits, which can be differentially important for attracting seed dispersers. Here we report on the effect of trees and the vegetation’s structure and function on the zoochorous seed rain promoted by frugivores in an abandoned pasture in Southeastern Brazil. We installed seed traps beneath 50 isolated trees of different species and dispersal syndromes. In one year, we collected a high seed density of, at least, 47 species underneath the isolated trees, and a scarce seed rain in the open field. The majority of the seed species was allochthonous in the study site, which points to the use of trees as stepping stones in the landscape by dispersers. Isolated trees’ canopy size and dispersal syndrome, vegetation cover, and number of surrounding zoochorous trees did not affect seed rain. Only tree height and number of surrounding zoochorous shrubs and herbs positively affected seed rain richness. In summary, perches as structural features are as important as fruit availability in intensifying seed rain. The relative importance of vegetation structural and functional characteristics on seed rain described in this study can potentially contribute to improve the management of disturbed sites and forest conservation.

Reynolds, C.* & Cumming, G.S. Percy FitzPatrick Institute of African Ornithology, DST/NRF Centre of Excellence, University of Cape Town, Rondebosch, Cape Town, South Africa [email protected] Bigger birds disperse better: seed traits matter less than vector for aquatic plant dispersal Seed dispersal is a fundamental ecological process and is strongly influenced by disperser and propagule characteristics. However, the relative importance of these characteristics for dispersal outcomes is unclear. We investigated differences in the dispersal of aquatic plants between Egyptian Goose (Alopochen aegyptiaca) and Red- billed Teal (Anas erythrorhyncha), two southern African duck species that differ markedly in size. The seeds of seven wetland plants with varying traits were fed to the ducks, and gut passage time and survival of ingested seed were determined. We examined the effect of disperser species and seed size and hardness on three dispersal characteristics and derived the corresponding dispersal kernels to explore differences in eventual distributions. The dispersal distance of a particular plant species was most strongly affected by disperser species. Egyptian Geese retained all seed species for longer than Red-billed Teal and the resultant dispersal kernels showed that seed dispersal by Egyptian Geese occurs to nearly twice the distance. Smaller seeds had significantly longer retention times and were also recovered in the highest numbers due to lower digestibility. There was no effect of disperser or seed traits on eventual germination. Our results suggest that seed traits may play a role in moderating dispersal distances within a particular vector, however, across vector species body size is more important in determining dispersal patterns. By exploiting different disperser species, wetland plants can be moved at different scales with little cost to germination. Seed traits and disperser ecology interact to influence the patterns of dispersal in aquatic systems and must be considered if we are to further develop our understanding of dispersal.

Rogers, H., Wandrag, E. & Dunham, A. Rice University, USA [email protected] Frugivore loss affects forest structure through an influence on treefall gap dynamics

Frugivores affect gap regeneration by facilitating colonization of fleshy-fruited species. We propose that frugivore loss will lead to reduced colonization by fleshy-fruited pioneer species in particular, reducing diversity, influencing canopy closure and ultimately affecting forest structure. The Mariana Islands offer a unique opportunity to examine the effect of frugivores on gap dynamics. Most tree species, including all pioneers, rely on frugivorous vertebrates to disperse their seeds, but an invasive snake was introduced to one island (Guam) and caused the functional extirpation of all native vertebrate frugivores. In contrast, the nearby islands of Saipan and Rota support similar forest to Guam, yet still have frugivores. We created experimental treefall gaps across all islands and monitored seedling regeneration for one year. We found a higher diversity of seedlings regenerating in gaps on islands with dispersers than on Guam. Gap regeneration on Saipan was faster than on Guam primarily because gaps on Saipan contained more than twice as many seedlings of pioneer species than those on Guam. At the current rate, our experimental gaps will close much sooner on Saipan than on Guam. This is consistent with the observation that there are two to four times more natural gaps on Guam than on islands with dispersers. We provide the first evidence that vertebrate seed dispersal can influence the physical structure of forests, with gap regeneration slowing in its absence. These results suggest that the on-going loss of biotic dispersers could result in forests that have lower diversity and more open forest structure.

Ruggera, R. A.*1,2,3, Blendinger, P.G.1,2 & Gomez, M.D.*1,3 1Consejo Nacional de Investigaciones Científicas y Técnicas, Argentina, 2Instituto de Ecología Regional, Universidad Nacional de Tucumán, Tucumán, Argentina, 3CIT – Jujuy, Cátedra de Biodiversidad y Desarrollo Sustentable, Facultad de Ciencias Agrarias, Universidad Nacional de Jujuy, Argentina [email protected] Linking structure and functioning in plant–frugivore interactions: importance of core frugivores and equivalence in their seed dispersal Multi-species mutualisms are thought to have a core of few species that disproportionally affect the remaining species in a network. Yet, the relative influence of core and peripheral species has not been properly tested, and the existence of geographical gradients in biodiversity has challenged the understanding on how reciprocal adaptations between frugivores and fruits could emerge. We propose that core species from different sites could be equivalent in the seed dispersal provided, which implies that cores could act as a unique selective force for fruit traits at a regional scale. We specifically ask: 1) Do core species affect partners more strongly than peripheral species? 2) Are core birds from different sites equivalent in the quantitative component (QC) of seed dispersal effectiveness provided? We monitored bird–fruit interactions in 10 sites of Yungas forest, NW Argentina. We recorded 3579 interactions with 69 fleshy-fruited plant species, 8 core bird species dispersed a larger seed number and seeds from more species than 46 peripheral bird species. Core birds had a higher QC than peripheral birds when consuming fruits of shrubs (Fobs=7.64, p=0.005), subcanopy (Fobs=19.1, p<0.001) and canopy (Fobs=4.5, p=0.02) trees, but not for epiphytes (Fobs=0.19, p=0.71) and Morus-like (Fobs=1.41, p=0.19) fruits. Through QC landscapes and simple regressions we found geographical replacements of core birds that had equivalent QCs. This study demonstrates that activity of a few core birds at a regional scale is extremely important for fruit removal of most plant species and that the ecological service provided by them is quantitatively equivalent.

Schleuning, M.1, Fründ, J.2 & García, D.3 1Senckenberg Biodiversity and Climate Research Centre, Frankfurt (Main), Germany, 2Dept. of Integrative Biology, Univ. of Guelph, Canada, 3Univ. of Oviedo, Depto. Biología de Organismos y Sistemas and Unidad Mixta de Investigación en Biodiversidad (CSIC-UO-PA), Spain [email protected] A trait-based concept to integrate structural and functional components of plant–frugivore interactions Network ecology has greatly advanced the understanding of the structural diversity of communities of interacting species. However, little advance has been made to understand the functional components of multi-species

46 interactions. We present a concept to develop the structural approaches of network theory into a trait-based framework for predicting ecosystem functions from biodiversity and mutualistic networks. An integration between structural and functional approaches in network studies requires the differentiation between three levels of biological organization that are determined by a sequence of processes. The levels correspond to: 1) the assembly of ecological communities by environmental filtering, 2) the formation of interaction networks by the matching of co- occurring species, and 3) the quantitative and qualitative components of mutualistic interactions providing the ecosystem function. We integrate these levels in a framework that categorizes relevant species traits by the respective organizational level. The identification of relevant plant and frugivore traits and the trait relationships across the organizational levels provide a generic template for testing the consequences of frugivore loss for seed- dispersal functions. For example, strong impacts on seed-dispersal functions are expected if large frugivores disappear first, interact preferentially with large-seeded plants and provide unique long-distance seed dispersal. A trait-based framework of the structural and functional components of plant–frugivore interactions will be useful to generalize the functional understanding of plant–frugivore interactions from local to regional spatial scales and to predict consequences of biodiversity loss for ecosystem functions by animals.

Solórzano-Filho, J.A. Faculty of Forestry, University of Toronto, Toronto, Canada [email protected] Big nut for a small mammal: The importance of Proechimys spp. (Echimyidae) in Brazil nut seed dispersal Neotropical rodents are important seed predators and dispersers of many tree species. Agoutis (genus Dasyprocta) have long been considered the exclusive seed disperser of the Brazil nut (Bertholletia excelsa), however some studies suggest that small rodents may also act as dispersers. I investigated this possible role of small rodents in the Southeastern Amazon. Experiments combining control and marked seeds of B. excelsa (n=1920) using fluorescent powder, inside and outside enclosures fitted with track plates were performed at six sites, with and without B. excelsa groves. These experiments allowed identification of the species of seed dispersers/predators as well as the fate and dispersed distance of the seeds. Densities of small mammals were determined by capture-mark-recapture techniques, and of the Brazil nuts by estimating the seed production of the individuals of B. excelsa based on their sizes. Proechimys spp. were the only small rodent species that removed seeds from the enclosures. Assuming that competition outside the enclosures had no effect on the amount of seeds removed by Proechimys spp., 34% of Brazil nuts outside the enclosures might be preyed upon or dispersed by them. These seeds were carried an average of 5.14 ± 3.73 m, and they took an average of 2.95 ± 1.45 days to be discovered, and no differences were observed between seeds placed inside or outside the enclosures. Of the marked seeds removed by Proechimys spp., 46% were effectively dispersed. My results show that Proechimys spp. can effectively disperse seeds and promote the establishment of new B. excelsa seedlings.

Sridhara, S.*1, Prasad, S.2, Westcott, D.3 & Edwards, W.1 1James Cook University, Australia, 2Jawaharlal Nehru University, India, 3CSIRO Ecosystem Science, Australia [email protected] / [email protected] Behaviour mediated by habitat heterogeneity and not fruiting trees better predicts seed deposition patterns by a tropical ruminant Current seed dispersal models, largely based on obligate avian frugivores, show strong links between spatial distribution of fruit plants and seed dispersal patterns. However, these models do not adequately incorporate behaviour of non-obligate frugivore dispersers that often track resources which may not overlap spatially with fruiting plants. Using chital (Cervidae, Axis axis) — a widely distributed deer species in India — as a model non- obligate frugivore disperser, we investigated whether its predator avoidance strategy, grazing strategy or simply the density of fruiting trees are robust predictors of seed dispersal patterns. In the foothills of the western Himalayas, India, we sampled 45 one-hectare plots to measure and model the relationship between resources such as fruiting

47 trees, availability of grass, cover from predators and quantity of seeds deposited. Results based on a GLM framework suggest that habitat type, which mediates the density of key resources such as grass and cover from predator, influences the quantity of seeds dispersed by chital. The presence of cover from predators and to a lesser extent the amount of grass within a plot best predicted the number of seeds found in a plot. Fruiting trees, however, had no influence on seed deposition. Our study underscores the need to incorporate greater level of behavioural details while understanding seed dispersal by non-obligate frugivores. These results also suggest that trophic interactions between disperser and their predators (mediated by cover) in addition to disperser behaviour alone (mediated by availability of grass) can influence seed dispersal patterns.

Terborgh, J.W.1, Zhu, K.2, Alvarez-Loayza, P.1 & Valverde, F.C.3 1Duke University Center for Tropical Conservation, USA, 2Carnegie Department of Global Ecology, Stanford University, USA, 3Botanical Research Institute of Texas, USA [email protected] Seed rain predicts diversity of a tropical floodplain forest in Perú For 8.5 years, we monitored the seed rain into a mature floodplain forest at the Cocha Cashu Biological Station in Peru using an array of 289 x 0.5 m2 traps. Here we address a perplexing issue: how seedfall at the scale of sapling establishment (ca 0.5 m2) can be reconciled with the overall diversity of a forest containing over 700 tree species. An area including all regular members of the tree community would be 6–8 orders of magnitude larger than the scale of establishment. We shall focus on species participating in the advance regeneration (AR), which collectively contribute 87% of the species in the community. However, >90% of the seed rain in this forest consists of seeds that are undispersed, or belong to lianas or small-seeded trees (e. g., Ficus) that do not participate in the AR. The rain of dispersed seeds of AR tree species is extremely scant (median, 4 per m2/yr). We will show that the diversity of AR saplings results from large spatial and temporal variability of seedfall. Synthetic sapling cohorts generated by randomly assigning success to seeds falling into each 0.5 m2 trap site capture much of the diversity of recruiting saplings. Weighting the probability of success by seed mass further enhances the reality of synthetic sapling cohorts. Density dependence must be extremely weak when only 4 seeds per m2/yr have the opportunity to compete. Thus, high species diversity does not arise through competition, but through extreme spatio-temporal heterogeneity in seedfall.

Thabethe, V.*, Wilson, A-L.*, Hart, L.A.* & Downs, C.T. DST-NRF Centre for Invasion Biology, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] Effect of invasive and indigenous avian species on seed germination of fleshy-fruited invasive alien plants in South Africa Avian frugivores play a key role in seed dispersal of many plant species, including invasive alien plants. This study assessed the effect of gut passage on the germination of selected invasive alien plant species in South Africa. Fruits of four fleshly-fruited invasive alien plant species: Solanum mauritianum, Cinnamomum camphora, Psidium guajava, and Morus alba, were fed to two species of indigenous turacos, Knysna (Tauraco corythaix) and Purple-crested (Gallirex porphyreolophus) Turacos, and invasive Rose-ringed Parakeets (Psittacula krameri). Seed retention time was determined as this may influence both seed dispersal and germination. Germination success of ingested seeds was compared to that of manually pulp-removed seeds, as well as to seeds in whole fruit. The germination success of seeds of all the invasive plant species increased significantly after ingestion by both turaco species compared to seeds from whole fruits. Germination success of manually pulp-removed seeds did not differ significantly from that of turaco-ingested seeds. In contrast, seed passage through the digestive tract of Rose-ringed Parakeets significantly reduced germination success and viability of ingested invasive plant species. Our results suggest that Knysna and Purple-crested Turacos are legitimate seed dispersers of fleshy-fruited invasive plants, while Rose-ringed Parakeets

48 are mainly seed predators. Although seed predation by Rose-ringed Parakeets negatively affects reproductive success of these plants, it is unlikely that this seed predator suppresses the population of these invasive alien plants in South Africa as the plants are already well established. Furthermore, they can facilitate dispersal by regurgitation and dropping uneaten fruits away from the parent plant.

Timóteo, S.1,2, Correia, M.1, Rodriguez, S.E.1 & Heleno, R.1 1CFE - Centre for Functional Ecology, Department of Life Sciences, University of Coimbra, Portugal, 2 Bristol Life Sciences Building, University of Bristol, UK [email protected] The refaunation of Gorongosa National Park and the reinstatement of the seed dispersal service Seed dispersal is vital for long-term vegetation dynamics and ecosystem stability. Large animals are disproportionally important dispersers for many large-fruited plant species and face higher extinction risk worldwide. We studied animal-seed dispersal in the Gorongosa National Park (GNP) in Mozambique. GNP supported some of the densest wildlife populations in Africa, until a 30-year long and devastating war pushed most large animals (e.g. lions, elephants, zebras, gnus) to the brink of extinction. Recently, legal protection and active restoration programs managed to recover many of those long-gone charismatic species into the GNP, particularly within the 6200 ha wildlife sanctuary where large animals density are closer to the pre-war densities.We used this unique natural experiment to evaluate the effect of defaunation and refaunation on the structure and functioning of seed dispersal interactions, building community-wide seed dispersal networks. Two habitats — mixed dry forest and miombo woodland, were sampled inside and outside the sanctuary by collecting animal dung along fixed transects, and performing focal observations of animals feeding. Retrieved seeds were identified by comparison to a reference collection, and interactions compiled into quantitative matrices. Several network and species-level descriptors of the interaction patterns were computed, including network specialization, niche overlap, interaction strength asymmetry, modularity and weighted nestedness. Results suggest that higher diversity of dispersers inside the sanctuary translates into more diverse and complex networks in both habitats. Overall, more seeds were dispersed inside the sanctuary and there was a higher redundancy of dispersers suggesting an effective restoration of the service provided to plants.

Tjelele, J.1,2, Ward, D.2 & Dziba, L.3 1Agricultural Research Council, Animal Production Institute, South Africa, 2School of Life Sciences, College of Agriculture, Engineering and Science, University of KwaZulu-Natal, South Africa, 3CSIR: Natural Resources and the Environment, South Africa [email protected] The effects of seed ingestion by livestock, dung fertilization, trampling, grass competition and fire on seedling establishment of two woody plant species The increasing rate of woody plant encroachment in grasslands or savannas remains a challenge to livestock farmers. The causes and control measures of woody plant encroachment are of common interest, especially where it negatively affects the objectives of an agricultural enterprise. The objectives of this study were to determine the effects of gut passage (goats, cattle), dung (nutrients), fire, grass competition and trampling on establishment of Acacia nilotica and Dichrostachys cinerea seedlings. Germination trials were subjected to 1) seed passage through the gut of cattle and goats and unpassed/untreated seeds (i.e. not ingested), 2) dung and control (no dung), 3) grass and control (mowed grass), 4) fire and control (no fire), and 5) trampling and control (no trampling). The interaction of animal species, grass and fire had an effect on seedling recruitment (P < 0.0052). Seeds retrieved from goats and planted with no grass and with fire (6.81% ± 0.33) had a significant effect on seedling recruitment than seeds retrieved from goats and planted with grass and no fire (2.98% ± 0.33). Significantly more D. cinerea and A. nilotica seeds germinated following seed ingestion by goats (3.59% ± 0.16) than cattle (1.93% ± 0.09) and control or untreated seeds (1.69% ± 0.11). Less dense grass cover, which resulted in reduced grass competition with tree

49 seedlings for light, space and water, and improved seed scarification due to gut passage were vital for emergence and recruitment of Acacia seedlings. These results will contribute considerably to the understanding of the recruitment phase of woody plant establishment.

Türke, M.1,2†, Fischer, C.3, Weisser, W.W.4 & Eisenhauer, N.1,2 1 German Centre for Integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, Deutscher Platz 5e, D-04103 Leipzig, Germany, 2 Institute for Biology, University of Leipzig, Johannisallee 21, D-04103 Leipzig, Germany, 3 Restauration Ecology, Department of Ecology and Ecosystem Management, Technische Universität München, Emil-Ramann- Strasse 6, D-85354 Freising, Germany, 4 Terrestrial Ecology Research Group, Department of Ecology and Ecosystem Management, Technische Universität München, Hans-Carl-von-Carlowitz-Platz 2, D-85354 Freising, Germany Earthworms act as seed dispersers in grassland and forests. Recent studies also reported gastropods to disperse vascular plants, ferns, mosses and lichens, mostly in forest ecosystems. In agricultural landscapes a number of weed species lack dispersal structures, and mechanisms of non-human dispersal are often unknown. Although a number of earthworm, snails, and slug species, many of which are invasive, can be found in agrienvironments, their role for the dispersal of weeds has been poorly explored. In temperate regions, we tested the feeding behavior of earthworms, invasive to North America, as well as of a European invasive horticultural pest slug being offered seeds of 12 arable weeds and three endangered herbs in the laboratory. Complementary video observations of seed depots were made in an organic rye field to identify seed consumers. In a second study, we tested invasive aquatic apple snails for their potential to disperse seeds of ten weed species growing in tropical irrigated rice fields. Removal of seeds was also assessed in rice terraces in the Philippines. Our results support the assumption that earthworms and gastropods facilitate weed movement in the agri-environment. Seeds of most weed species were readily swallowed by earthworms and by terrestrial and aquatic gastropods, and defecated seeds mostly germinated at similar rates as control seeds. We conclude that these invertebrates are overlooked dispersers of arable weeds. Weed dispersal by highly abundant invasive species might challenge traditional weed management. On the other hand, they might help re-establishing plant diversity in organic farming.

Kaplin, B.1, Carlo-Joglar, T.A.2, Acevedo, J.D.3 & Torres-Caba, J.*1 1Antioch University, New England, USA, 2Penn State University, USA, 3Texas A&M University, USA [email protected] The role of introduced rats (Rattus rattus) as seed dispersers and predators in La Olimpia Forest, Adjuntas, Puerto Rico Invasive species are known to cause significant threats to biodiversity, especially on island ecosystems. In La Olimpia Forest in Adjuntas, Puerto Rico, a secondary subtropical wet forest, the role of introduced black rats (Rattus rattus) in the dispersal of Sierra palm (Prestoea montana) seeds was studied. The Sierra palm, a native to the Greater and Lesser Antilles of the Caribbean islands, is important for maintaining soil stability on steep mountain slopes and its fruit, a valuable food resource for the endangered Puerto Rican parrot (Amazona vittata). Fruits of 18 Sierra palm individuals were collected, tagged and placed under their respective parent trees in depots of 25 seeds each. Approximately 75% of the seeds were removed from these seed depots. Automated camera traps photographed only black rats interacting with the fruits in the depots. The majority of the removed fruits had their pulp predated and seeds gnawed and damaged. However, several fruits were found intact or partially damaged. These results suggest that dispersal and predation are occurring as an interaction between black rats and the seeds of native Sierra palm in this forest. Through these interactions, rats are probably influencing the distribution of this native palm and the forest communities in Puerto Rico. By understanding such interactions we can focus on strategic management efforts for native plant species and the maintenance of ecological processes.

Trolliet, F.*1, Forget, P-.M.2, Huynen, M-C.1 & Hambuckers, A.1 1University of Liège, Belgium, 2Museum National d’Histoire Naturelle, Brunoy, France

50 [email protected] Effect of hunting pressure and forest fragmentation on seed dispersal of Staudtia kamerunensis (Myristicaceae) in the Western Congolian forest–savanna mosaic Numerous studies have investigated the impact of hunting and forest fragmentation on seed dispersal processes, but few of them in Africa. The African continent, however, holds the greatest abundance of large frugivores on Earth and is subject to increasing levels of hunting and forest fragmentation. Frugivory and seed dispersal in the pan- tropical family Myristicaceae has been well studied in the Neotropics, but remain barely known in Afro-tropical forests. Here we investigated how hunting, forest cover, and fruit availability influence the dispersal capacities of Staudtia kamerunensis in a mosaic composed of forest patches and savannas in D.R. Congo. We selected 34 fertile female S. kamerunensis trees distributed, at a landscape scale (300 km²), across 5 sites characterized by a gradient of hunting pressures and density of the study trees. The trees varied in size and surrounding forest cover (patch and corridors). We quantified the percentages of seed dispersal failure using litter trap and fruit remains during the entire 2013 fruiting season, and identified fruit eaters through focal observations. The most frequently observed frugivore was the White-thighed Hornbill (Bycanistes albotibialis). Our GLMM analyses show that increasing hunting pressure, decreasing abundance of B. albotibialis and decreasing forest cover near focal trees significantly increased dispersal failure. The limited forest cover of some patches, resulting from the highly fragmented feature of the landscape, and hunting pressure, are likely to impact negatively the abundance of B. albotibialis. Consequently, we suggest the dispersal system reached saturation because of a lack of effective seed dispersal away from parent trees.

Tsuji, Y. Primate Research Institute, Kyoto University [email protected] Inter-annual and inter-rank variation in characteristics of endozoochory by wild Japanese macaques (Macaca fuscata) Endozoochory is important to the dynamics and regeneration of forest ecosystems. Despite the universality of inter- annual variation in fruit production and existence of dominance hierarchy and related behavioural difference in group-living animals, few studies have addressed these effects on the effectiveness of endozoochory. I recently have evaluated the characteristics of endozoochorous dispersal by wild Japanese macaques (Macaca fuscata) in northern Japan. I collected faecal samples in the fall and quantified the proportion of faeces containing seeds, number of seeds per faecal sample, ratio of intact seeds, and seed diversity. The proportion of faeces containing seeds of any species did not show significant inter-annual variation, while species-level proportions did. The intact ratio of seeds varied significantly among ranks, and this relationship varied between years and dominant plant species. The number of seeds per faecal sample also varied among ranks, but did not between years. Finally, seed diversity did not show significant inter-annual and inter-rank variation, which was attributed to the longer duration of the macaque gastro-intestinal passage time and the mixing of seeds from different feeding bouts uniformly in the faeces. This study demonstrated that frequency and success of seed dispersal by macaques showed inter-annual and inter-rank variation, indicating low specificity across the seed–macaque network. The variation led by temporal and social factors in the quality of seed dispersal may provide evidence of high resilience in response to fluctuating environmental conditions in temperate forests.

Valido, A., Pérez-Méndez, N.1, Jordano, P.1, & García, C2 1Estación Biológica de Doñana (EBD-CSIC), Spain, 2Centro Investigação em Biodiversidade e Recursos Genéticos (CIBIO), Portugal [email protected] Downsized mutualisms: Demographic and genetic consequences of lizard body size reduction for an insular plant species Defaunation-driven body-size reduction of frugivores might entail the loss of ecological functions that drive recruitment and genetic patterns of plant populations. Here, we benefit from of a lizard extinction processes that

51 occurred in three Canarian islands, defining a gradient of body size: Gallotia stehlini (max snout–vent length: 280 mm, Gran Canaria), G. galloti (145 mm, Tenerife), and G. caesaris (111 mm, La Gomera). We studied the demographic and genetic consequences of lizard downsizing on Neochamaelea pulverulenta (Rutaceae), an endemic plant species that relies exclusively on lizards for seed dispersal. First, we detected that the frequency distributions of different age/size classes did not differ among islands. However, considering qualitative aspects of the dispersal outcomes, we found significant reductions in seedling recruitment outside the canopy, reduced effective recruitment rate, and seedling vigour in populations hosting small- to medium-sized lizard species. Second, we found a progressive truncation of the seed dispersal kernels mirroring the lizard body sizes, from a maximum seed dispersal distance that reached 94 m in Gran Canaria and only 4.5 m in La Gomera. Third, despite no evidence of genetic erosion at the local scales, the extent of the spatial genetic structure differed across the three scenarios. We detected an increased fine-scale spatial genetic structure and increased genetic clustering in populations hosting small- to medium-sized lizard species. Overall our results confirm, by using a combination of in-depth genetic and demographic methods, that defaunation-driven downsizing of seed dispersers has far-reaching implications on the regeneration, dispersal ability and genetic structure of plant species.

Wang, B. Xishuangbanna Tropical Botanical Garden, Chinese Academy of Sciences [email protected] Unequal seed dispersal by seed-size sensitive rodents: a case study in Chinese White Pine (Pinus armandii) Variation in individual seed production is high in many plant species. How does this variation affect seed-dispersing animal mutualists, and in turn the fitness of individual plants? In this study, we first surveyed the intra-specific variation in seed production in a Chinese White Pine population. For 134 target individual trees, there was very high variation in seed size, with mean seed mass ranging from 0.038 to 0.361 g among trees. This variation in seed size was correlated with neither tree size nor total seed set per tree. Furthermore, for 30 of the 134 trees that produced seeds two years later, there was a strong correlation in seed size between years, implying consistent differences among individuals. For a subset of 67 trees, we monitored the fate of 15,301 seeds: ignored by dispersers (8380), eaten (4381), or cached (2249). There was a large effect of seed size, with trees producing on average larger seeds having a much greater proportion cached. In contrast, the proportion of seeds cached was independent of total number of seeds produced. The probability that seeds are cached is an important proximate component of fitness, because cached seeds (as opposed to ignored or eaten seeds) are most likely to give rise to successful seedlings and adults. Even though seed size has a large potential to affect the fitness of individuals, small seeds are nonetheless important as food for rodent dispersers. Therefore, the fitness of large-seeded trees could be enhanced by small- seeded neighbors that help maintain populations of mutualistic dispersers.

White, J.D.M., Midgley, J.J. & Bronner, G.N. Department of Biological Sciences, University of Cape Town, South Africa [email protected] First use of camera traps to study scatter hoarding in Cape fynbos We expand on the current research of scatter hoarding in the Cape fynbos by investigating small mammal behaviour using remotely activated camera traps. All previous research in the Cape has used indirect circumstantial field and controlled arena trial evidence to corroborate likely scatter-hoarding small mammals. We provide the first direct video evidence of seed predation and scatter hoarding behaviour of several Cape small mammals. We spread the known geographic range of scatter hoarding within Cape fynbos by investigating novel nut-fruited plant taxa (the diverse but poorly studied genus Cliffortia and new Leucadendron spp.), which are predicted to be scatter hoarded. Hairy-footed Gerbil (Gerbillurus paeba) and Cape Spiny Mouse (Acomys subspinosus) are both shown to scatter hoard L. loranthifolium and L. sessile nuts, while G. paeba also scatter hoards L. pubescens nuts. The Striped Field Mouse (Rhabdomys pumilio) and Namaqua Rock Mouse (Aethomys namaquensis) are shown to have high predation

52 rates on various indigenous nuts (including C. cuneata and C. phillipsii nuts at Helderberg Nature Reserve), but never to bury one. General responses to seeds (distances removed, predation rates, burial rates) are summarised for each recorded small mammal species.

Wilson, A-L.*, Shuttleworth, A. & Downs, C.T. DST-NRF Centre for Invasion Biology, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] Assimilation efficiency of the Red-winged Starling Onychognathus morio fed Strelitzia nicolai arils Most fruits consumed by avian frugivores comprise a fleshy pulp. In contrast, the fruits of the Strelitziaceae family of flowering plants consist of a hardened seed with a woolly aril at its tip. Although frequently consumed by birds, the digestibility of these woolly arils remains unknown. Consequently, we fed whole Strelitzia nicolai seeds with arils to captive Red-winged Starlings (Onychognathus morio) to determine daily food and energy intake, digestive transit time and assimilation efficiency. Strelitzia nicolai arils were found to contain a remarkably high lipid content (66.78 %), with a correspondingly low water (7.61 %) and protein (5.79 %) content. Birds generally ate only the aril of S. nicolai seeds, but if they swallowed the seeds whole, then they either defecated or regurgitated the seeds without the aril. Red-winged Starlings were found to have fast digestive transit times (approx. 24 min) and very high apparent assimilation efficiencies (approx. 96 %) of the aril of S. nicolai seeds. The results obtained in this study indicate that the woolly pulp of S. nicolai is clearly a good source of energy for Red-winged Starlings and consequently may explain why they willingly feed on this seemingly unappealing food source.

Wotton, D.M.1,2, McAlpine, K.G.2 1Moa’s Ark Research, 2Department of Conservation, University of Canterbury [email protected] Frugivory and seed dispersal of invasive plants in New Zealand Frugivores play a key role in spreading non-native invasive plants, via seed ingestion and subsequent dispersal. We reviewed seed dispersal of fleshy-fruited invasive plants in New Zealand, focusing on plant and frugivore traits that influence spread. We found almost a third (32.9%) of 295 invasive species have fleshy fruits adapted for vertebrate dispersal. Fruiting phenology differs between invasive and native plants, with many invasive species fruiting from late autumn to early spring (May to September) when native fruits are scarce. Invasive plant fruiting duration does not differ from natives. The main dispersers are four non-native birds (Turdus merula, Sturnus vulgaris, T. philomelos and Acridotheres tristis), one native bird (Zosterops lateralis), three endemic birds (Hemiphaga novaeseelandiae, Prosthemadera novaeseelandiae and Anthornis melanura) and two non-native mammals (Trichosurus vulpecula and Sus scrofa). All 10 species eat significant quantities of fruit annually or seasonally and are widespread in New Zealand. The bird dispersers rarely damage seeds, while the mammals destroy some seeds. T. merula, Z. lateralis and S. vulgaris are the most important dispersers, occupying the largest range and dispersing the greatest number of species. Sturnus vulgaris also disperses seeds long distances, including to island reserves several kilometres offshore. The large-bodied H. novaeseelandiae (New Zealand pigeon) is the key disperser for three invasive plants with large, single-seeded fruits. Seed dispersal by frugivores is a significant means of spread for invasive plants in New Zealand. However, surprisingly little research has been conducted and its management poses a challenge.

Zungu, M.M.* & Downs, C.T. DST-NRF Centre for Invasion Biology, School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa [email protected] Effects of tannins on fruit selection in three southern African frugivorous birds Tannins are common secondary compounds in plant material and are known for their ability to bind to protein which reduces nitrogen availability in the diet. In fruits, these compounds are responsible for their astringency which is

53 thought to result in reduced food intake. In this study, the repellent effects of tannins were examined in three species of frugivorous birds: Red-winged Starlings Onychognathus morio, Speckled Mousebirds Colius striatus and Cape White-eyes Zosterops virens. Birds were fed artificial fruit diets containing varying levels of tannins in paired choice tests with the amount of food eaten by birds used to determine preference. Red-winged Starlings preferred the control diet, were indifferent to the medium tannin diet and were deterred by the high tannin diet whereas Speckled Mousebirds and Cape White-eyes were not deterred at all concentrations. The discrepancy in the results was attributed to differences in taste sensitivity, tolerance levels and detoxification mechanisms of secondary compounds between species. Because fruit selection and ultimately fruit removal rates affect plant community composition, the disparity in the results suggests that frugivorous birds do not contribute equally to plant community dynamics. However, plant secondary compounds in fruits are diverse and their effects are similarly diverse and there is potential that different groups of secondary compounds generate disparate effects. Similar studies on other types of secondary compounds may thus contribute towards a broader understanding of the role of secondary compounds in mediating fruit–frugivore interactions.

Zwolak, R., Bogdziewicz, M.* & Wróbel, A. Adam Mickiewicz University, Faculty of Biology, Department of Systematic Zoology, Poland [email protected] Benefits of masting for the European beech: timing of granivore satiation and benefits of seed caching support the predator dispersal hypothesis The predator dispersal and predator satiation hypotheses provide alternative explanations for masting. Both assume satiation by seed-eating vertebrates. They differ in when satiation occurs: before or after seed removal and caching by granivores (predator satiation and predator dispersal, respectively). This difference is often overlooked, but it is demographically important because cached seeds are dispersed and often have a microsite advantage over non- dispersed seeds. We conducted rodent exclosure experiments in mast and non-mast years to test contrasting predictions of the predator dispersal and predator satiation hypotheses in our study system: yellow-necked mice (Apodemus flavicollis) interacting with European beech (Fagus sylvatica). Specifically, we tested whether 1) removal of seeds from forest floor is similar during mast and non-mast years (i.e., lack of satiation before seed removal), 2) masting decreases removal of cached seeds (i.e., satiation after seed removal), and 3) seed caching increases probability of seedling emergence relative to non-cached seeds. We found 1) masting did not result in satiation of seed removal, 2) masting decreased removal of cached seeds, and 3) seed caching dramatically increased the probability of seedling emergence relative to non-cached seeds. European beech benefits from masting by scatter- hoarder satiation that occurs only after seeds are removed and cached in suitable microsites. These findings do not exclude other evolutionary advantages of beech masting. However, they indicate that fitness benefits of masting range beyond the most commonly considered advantages of predator satiation and increased pollination efficiency.

FSD POSTER ABSTRACTS

Burns, K.C. Victoria University of Wellington [email protected] What structures fruit–frugivore webs: traits or chance? The simplest and arguably the most ubiquitous pattern in seed dispersal mutualisms is size coupling: large frugivores tend to consume larger fruits and small frugivores tend to consume smaller fruits. Despite the simplicity of this pattern, the potential mechanisms responsible for fruit–frugivore size coupling are mechanistically divergent and poorly resolved. Size coupling could arise deterministically, if large frugivores actively seek out larger fruits to maximize their foraging efficiency. Alternatively, size coupling could also arise passively, if frugivores forage randomly, but are able to consume only those fruit species that are smaller than their gape width. I observed birds forage for fruits in a New Zealand forest reserve at approximately five-day intervals for six years to test for fruit– frugivore size coupling. I then derived a suite of network analyses to establish whether fruit–frugivore size coupling was best explained by active or passive foraging by frugivores. Results showed a strikingly strong pattern in size coupling, the average size of fruits consumed by each frugivore species increased with their maximum gape width. Simulation analyses revealed that over 70% of variation in interaction frequencies in the observed fruit–frugivore web could be explained by a size-constrained, passive, foraging model. Foraging models in which birds foraged actively for different-sized fruits to improve their foraging efficiency performed more poorly. Results were therefore consistent with the hypothesis that apparently nonrandom patterns in seed dispersal mutualisms can sometimes arise from simple stochastic processes.

Cestari, C.1, Pizo, M.A.1 & Loiselle, B.2 Universidade Estadual Paulista, Brasil1, University of Florida, USA2 [email protected] Foraging costs of a fruit-eating lekking bird: Integrating sexual selection and optimal foraging theories Male reproductive success in lek mating systems is correlated with territory attendance, display activity, and aggression rate. Likely consequences of these behaviors by top-ranking males are: (1) less time available for foraging, and (2) increased energetic demands if displays are ‘costly’ in terms of energy investment. Here we examine how display effort of males potentially affects fruit foraging decisions using an experimental approach where fruits are offered on display courts. We hypothesized that male rank, as indicated by territory attendance and display rate, would be related to consumption of fruits on the courts. To test this hypothesis we used Manacus manacus (Aves: Pipridae) as a model to represent lek systems. We placed branches with 100 Myrsine ferruginea fruits next to 23 display courts in five lek areas of M. manacus in southeastern Brazil. The differential consumption of M. ferruginea fruits and display rate (“snap-jump” bouts) by males were recorded from the arrival of the first male in the lek area to 8:30 and 9:00 h using mini-cameras set up at 2 m from the males’ courts. We found a positive correlation between the number of consumed fruits and display rate by males, even after correcting for time spent away from display courts. This result supports our prediction that males that display more consume more fruits, likely to meet larger energetic needs for reproductive success, while minimizing time spent away from their courts. Our next steps will focus on the how the differential rank of males affects fruit choice in terms of energetic rewards.

Cortinoz, J.1, Traveset, A.2 & Siva, W.3 1Graduate Program in Ecology, University of Campinas, Brazil, 2Institut Mediterrani d'Estudis Avançats (CSIC-UIB), Spain, 3Dept. Animal Biology, University of Campinas,IB/UNICAMP, Cx Postal 6109 Campinas- SP, Brazil. [email protected]

Fleshy fruit use by pulp and seed- eating insects in a tropical island: a network approach to evaluate community patterns Patterns of insect frugivory at community level are little explored. Even though fruit occurrence can be highly unpredictable and ephemeral, many insect species develop inside fruits, feeding on pulp or preying upon seeds. There is scarce information about host selectivity by fruit-feeding insects, whether it varies between guilds (pulp vs. seed-eating insects) and on the biotic and abiotic factors that shape these interactions. The objective of this study was to evaluate patterns of fleshy fruit use by insects at a community level and determine how fruit traits and phenology mediate these interactions. We studied interactions between insects and fruits from 12 Myrtaceae species in an Atlantic forest Island in southeast Brazil for two years. We used a network approach assessing metrics that describe structure both at the network and species level. We tested which fruit traits influence host selectivity and compared resource use between insect guilds. We found high connectance, low nestedness and moderate specialization in the interactions. Although fruit availability changed between years, few parameters of the network were affected. Fruit and seed size are important factors determining the number and specificity of plant interactions with animals. Selectivity differs between insect guilds, showing that pulp-eating insects tend to be generalized, while seed predators are more specialized. This could indicate a higher physiological and phenological dependence of seed predators on their hosts than pulp-eating insects. Insect frugivory should be further explored as a good model for consumer-resource studies and coevolution between herbivores and plants.

Dracxler, C.M.*1, Confais, A.2 & Forget, P-.M.1 1Muséum National d’Histoire Naturelle, Département Ecologie et Gestion de la Biodiversité, France, 2Université de Tours, France [email protected] Effects of anthropogenic activities on vertebrate and invertebrate seed predation Seed predators play an important role in shaping plant populations. By consuming seeds, predators directly affect seed survival, influencing subsequent phases of the reproductive cycle. The final outcome of seed predation is influenced by several aspects of both sides of the interaction, such as predator type, seed size and fruit timing. In disturbed forests, however, responses by vertebrate and invertebrate seed predators to human activities remain unclear. We initiated a meta-analysis to disentangle the effects of anthropogenic activities — hunting, forest fragmentation and logging — on both pre- and post-dispersal seed predation by vertebrates and invertebrates. We listed 80 studies published from 1983 to August 2014 providing information on seed predation in control and disturbed forests. Preliminary results show that seed predation is affected by disturbance type, seed predator, and by the interaction between these factors. While invertebrate seed predation seems to be positively affected by hunting and fragmentation, vertebrate seed predation rates decreased in hunted forests. Considering the vulnerability of vertebrates in hunted forests and their contribution to seed dispersal, disturbed forests may present a reduction of seed removal and consumption events, favoring seed accumulation beneath parent plants. This pattern, in turn, can affect seed predation by invertebrates, which are more likely to be host-specific and to act in a seed density dependent manner. Our findings reveal consistent effects of hunting and forest fragmentation on vertebrate and invertebrate seed predation and indicate study gaps that may contribute to the understanding of seed predation in a multi-trophic interactions perspective.

Ehlers Smith, D.A.*, Ehlers Smith, Y.C. & Downs, C.T. DST-NRF Centre for Invasion Biology, School of Life Sciences, Pietermaritzburg, University of KwaZulu-Natal, South Africa [email protected] Primates as seed predators — a proposed investigation into the impact of primate granivory in tropical forests Up to 75% of tropical tree species produce fruits adapted for animal dispersal, and it is estimated that animals move up to 95% of seeds produced. Primates account for 25–40% of frugivore biomass in tropical forests and are ideally suited to seed dispersal, being highly mobile and adept at manipulating a wide variety of fruits. However, some primates, such as the family Colobinae, whose digestive morphology permits consumption of unripe seeds, may in

56 fact act as seed predators, but their effect on tropical forest production and ecology is poorly understood. We aim to quantify the impact of seed predation by primates in Sabangau Forest, Central Kalimantan, Indonesia, using a granivorous primate species, the red langur (Presbytis rubicunda), 76% of whose mean annual diet comprises unripe seeds. We intend to test if the species is a true seed predator by processing faeces for intact seeds, which will be planted to assess germination viability. We will quantify the impact of granivory on forest ecology by comparing density and fruit phenology of the most important fruit species in the diet with individual ingestion rates and known population density, home-range size and day-range length of red langurs. As red langurs eat unripe seeds, we also aim to quantify their impact of competitive exclusion on sympatric, frugivorous primates by calculating the degree of overlap with the known diet of Bornean orang-utans (Pongo pygmaeus) and Bornean southern gibbons (Hylobates albibarbis). This study has the potential to improve our knowledge of tropical forest ecology and inter-species interactions.

Escribano-Avila, G. Institute of Ecology, Technical Particular University of Loja, Ecuador [email protected] The relevance of size on potential plant–frugivore interactions of the Tumbesian dry forest: is bigger always better? Tropical dry forests are among the most diverse and threaten ecosystems. Unfortunately, we lack basic knowledge on regeneration dynamics in which seed dispersal plays an instrumental role. The main dispersal syndrome in tropical dry forests is endozoochory. Yet, the plant–frugivore interactions that shape the seed dispersal network are still unknown. This information, despite being urgently needed to promote ecosystem recovery and conservation of tropical dry forests, is very costly to obtain in terms of time and money. Here we provide information on the potential dispersal network of the tropical dry forest using bibliographic and easily-collected field data. Large-sized frugivores are able to disperse a wider array of tree species and seeds to further distances than smaller ones. Thereby it is expected that large-sized frugivores play a key role in the dispersal of tropical dry forest species. However large-sized frugivores are usually the most threatened in the community, and it is therefore necessary to account for their conservation status and threats. Overall results showed that “size” of both, fruits and frugivores, shaped the potential dispersal network. Larger-sized frugivores acted as network-generalists as they dispersed most plant species, thereby playing a key role as network connectors. The largest-sized frugivores were the main and virtually the only dispersers of plants bearing the biggest fruits (i.e. network-specialists), thereby being essential in the maintenance of seed dispersal for big-fruited species. However, large-sized frugivores are threatened by habitat loss, fragmentation and hunting, and it is therefore necessary to establish urgent conservation measures to promote regeneration dynamics of the Tumbesian dry forest in Ecuador.

Falcón, W.*1, García, C.2, Ozgul, A.1 & Hansen, D.1 1Institute of Evolutionary Biology and Environmental Studies, University of Zürich, Switzerland, 2Centre for Research of Biodiversity and Genetic Resources (CIBIO), University of Porto, Portugal [email protected] The effects of animal-mediated intra and inter-island seed dispersal on the population genetics of Solanum aldabrense on Aldabra Atoll Dispersal of seeds mediated by animals has a profound effect on the population structure of plants. The Aldabra endemic Solanum aldabrense is highly dependent on frugivores for seed dispersal, and there are many available genetic resources for the genus, thus making it an ideal candidate to assess the outcomes of endozoochory. We aim to do this by coupling genetic and demographic sampling techniques, vegetation types, and movement patterns of frugivores. We will focus fine-scale demographic and genetic sampling on Picard and Polymnie to assess intra-island dispersal. We will identify life-history stages, and use seedlings to evaluate reproductive plants to seedling dispersal events. Standard genetic techniques and analyses will be employed to elucidate the effects of endozoochory on a)

57 dispersal distances, b) the effects of habitat and plant neighbourhood on demographic parameters, and c) the contributions to seedling establishment by plants in different life-history stages. We will also investigate the potential effects of tortoise-mediated dispersal (tortoises are absent on Polymnie). Moreover, to assess the effects of animal-mediated inter-island dispersal on gene flow, we will collect genetic samples on the other three main islands and islets. We hypothesise that tortoises will provide intra-island, long-distance dispersal services, while birds will provide short-distanced and clumped dispersal of genetically similar individuals, and rarely, inter-island seed dispersal. Parent-to-seedling dispersal distance will be limited by habitat availability and neighbourhood composition, while plants with higher reproductive output will be more represented amongst sampled seedlings. Finally, we expect to find low levels of inter-island gene flow due to dispersal by birds.

Jara-Guerrero, A.1, Espinosa, C.I.1, De la Cruz, M.2,3, Méndez, M.3 1Departament of Natural Sciences, Technical Particular University of Loja, Ecuador, 2Departament of Plant Biology, Politecnica University of Madrid, Spain, 3Biodivesity and Conservation Area, Rey Juan Carlos University, Spain [email protected] Legitimacy of deer as seed dispersers of woody species in a tropical dry forest This work analyzes the role of deer on the regeneration of woody species in a tropical dry forest. This study was conducted in the Arenillas Ecological Reserve, southwestern Ecuador. During 23 months, all deer faecal pellets observed along a 3.4 km path were collected once monthly. All seeds present in the pellets were recorded. The germination per cent of seeds that passed through deer guts was compared with control seeds taken from the plants as reference for the legitimacy of deer as seed dispersers. In total, 386 pellets were collected and 65.3% had seeds. Seeds of 11 woody species were identified in the pellets. Seeds of Chloroleucon mangense, Caesalpinia glabrata, Senna mollissima and Piptadenia flava were the most abundant in the pellets. These species have legume fruits and are considered autochorous species. We also found species with berry fruits (Cactaceae, Randia aurantiaca and Jacquinia sprucei). The seeds of only eight species germinated from pellets, therefore, the legitimacy of deer as seed dispersers was analyzed only for four species (C. mangense, C. glabrata, L. trichodes and S. mollissima) for which we had a control. Deer were legitimate dispersers at least for two species, C. mangense and S. mollissima. The percentage of germination for the other two species, C. glabrata and L. trichodes, was significantly lower in seeds from pellets than the control. However, decrease in germination could be compensated for by longer distance dispersal.

Kitamura, S.1, Chodapisitkul, S.2, Sooktawee, A.2 & Poonswad, P.3 1Ishikawa Prefectural University, Japan, 2Dusit Zoo, Thailand, 3Mahidol University, Thailand [email protected] Seed retention times for wild fruits consumed by captive hornbills (Buceros bicornis and Rhyticeros undulatus) in Thailand Frugivorous vertebrates provide important ecosystem services by transporting seeds of fleshy fruited plants. The time it takes seeds to pass through the gut of vertebrates is an important aspect of endozoochorous seed dispersal because it influences seed dispersal distance. Theoretically, the longer the retention time, the more likely a seed is to be widely dispersed. Regurgitation times for four fruit species (Dimocarpus longan, Bhesa robusta, Horsfieldia amygdalina, and Aglaia spectabilis) were determined using 10 captive hornbills (7 Great Hornbills and 3 Wreathed Hornbills) in Dusit Zoo, Thailand, from September 2010 to November 2012. Ripe fruits were fed to these hornbills and the length of time from consumption to regurgitation was noted for each fruit species. Most seeds were regurgitated within an hour but seed retention times varied significantly among fruit species. Median retention times were 28 min for A. spectabilis, 50 min for B. robusta, 47 min for H. amygdalina and 64 min for D. longan in Great Hornbills and 47 min for A. spectabilis, 45 min for B. robusta, 42 min for H. amygdalina and 78 min for D. longan in Wreathed Hornbills. The dispersal distance of a seed is a function of its retention time within its disperser and the disperser's range and rate of travel. Wide ranging daily hornbill movement patterns and the reasonably long

58 seed retention times presented here, suggest that hornbills may frequently disperse seeds some distance from their sources.

Kuhlmann, M.*1 & Ribeiro, J.F.2 1University of Brasilia, 2Embrapa Cerrados [email protected] Phylogenetic considerations on the frugivory mutualism in the Cerrado biome, Brazil The Cerrado includes forest, woodland and herbaceous environments and is one of the richest biome of the planet. It is a hotspot, with over 12,000 species of vascular plants (40% endemic) and more than 2,500 species of vertebrates. It is estimated that about 4,000 species of Cerrado plants depend on animals for seed dispersal and that 50% of the bird and mammal species consume fruits. This study aimed at evaluating different animal dispersal mechanisms for angiosperm families within the Cerrado biome, and to determine which lineages exhibit greater restriction in the plant–frugivore mutualism. To address these questions, we evaluated about 1,000 animal- dispersed fruit species belonging to 300 genera and 100 families present in the biome. Four categories of the frugivory mutualism were considered: bird fruits, bat fruits, non-flying-mammal fruits and ant fruits. Families with more than one dispersal agent were considered as “mixed”. Among the taxa analyzed, 50% of species and 50% of genera had characteristics related to bird frugivory, such as small and colorful diaspores, and over 60% of the families were restricted to only one kind of frugivore. Within these specialized families, 80% were exclusively bird fruits. The percentage of families with restricted mutualism in the five major lineages of angiosperms was 78% in basal angiosperms, 67% in monocots, 75% in basal eudicots, 57% in rosids and 61% in asterids. Therefore, phylogenetically, it seems that restrictive frugivory mutualism is more common than “mixed” mutualism among Cerrado families, and that plant–bird relationships must have had a prominent role in the evolution of fruit types in this biome.

Martínez-López, V. *1,2, Zapata, V.*1, Smith-Ramírez, C. 3,4 & Robledano-Aymerich, F.1 1Department of Ecology and Hydrology, University of Murcia, Spain, 2Department of Zoology and Anthropology, University of Murcia, Spain, 3Institute of Ecology and Biodiversity (IEB), Chile, 4Faculty of Forestal Science, Austral University of Chile, Chile [email protected] Dispersal of fleshy-fruited trees and shrubs by birds in Mediterranean-type regions: a review. Seed dispersal has been studied worldwide. Consequently, there is much information available about different topics (e.g. long distance dispersal, effects of fragmentation on seed dispersal patterns, seed dispersal networks, etc). However, fewer studies address these topics together in order to offer an integrative picture about where we are, and which are the main questions to be resolved. In this context, we have reviewed research on the dispersal of fleshy- fruited trees and shrubs, focusing our search on a single bioclimatic region (Mediterranean-type) and on a single disperser assemblage (birds). Our aims were: 1) to define and to compare seed dispersal patterns among Mediterranean areas and 2) to identify the main gaps in our study topic in the studied areas. The results to date show huge differences in the number of papers published among the different areas of the Mediterranean biome. There is much more knowledge about the Mediterranean Basin than from any of the remaining regions. Furthermore, some environments monopolize the research effort on dispersal, such as natural areas or fragmented landscapes, while others are poorly studied such as agroforestry areas or semiarid environments. In conclusion, we propose a series of topics and areas where research effort should increase, preferably through transnational cooperative studies, with the dual aim of increasing the local knowledge base and enabling future comparative studies and meta-analyses.

Mokotjomela, T.M.*1, Downs C.T.2, Knight, J.1 & Esler, K.3 1University of the Witwatersrand, South Africa, 2School of Life Sciences, University of KwaZulu-Natal, Pietermaritzburg, South Africa, Department of Conservation Ecology and Entomology & Centre for Invasion Biology, Stellenbosch University, Private Bag X1, Matieland 7602, South Africa [email protected] Dispersal agents for conservation of fleshy-fruited plants in South Africa Knowledge of keystone avian seed dispersal agents and their effectiveness in seed dispersal is critical for increasing adaptation of native fleshy-fruited plant species to habitat fragmentation and climate change by increasing their range. Seed dispersal effectiveness (SDE) is determined as a product of quantity (numbers of seeds dispersed) and quality (probability that a dispersed seed becomes a new adult plant) components. However, there is limited understanding of avian seed dispersal effectiveness in South Africa despite there being about 2000 plant species threatened by inherently rare avian seed dispersal services, competition with predominant alien flora, and habitat fragmentation. Previous studies have investigated the quality or quantity aspects of SDE separately, but no study has examined the SDE as a composite function of quality and quantity aspects in South Africa. By collating information on quality and quantity aspects of SDE from the published local literature, we derived the relative seed dispersal effectiveness of different bird species for various plant species. Regression analysis was applied to extrapolate for other plant species with similar seed/fruit characteristics, also considering potential changes in plant species’ spatial overlap with geographic distributions of important bird species. This approach allows identification of keystone avian dispersal agents at broader spatial scales, and thus may guide efforts for conservation of their dispersal services for native flora. With reference to threats to avian seed dispersal processes in South Africa (e.g. competition with alien plants and climate change), the conservation implications of avian seed dispersal effectiveness for the native flora are discussed.

Moran, C.1, 2, 3 & Catterall, C.1 1Griffith School of Environment, Griffith University, Brisbane, Australia, 2College of Marine and Environmental Sciences, James Cook University, Cairns, Australia, 3CSIRO Tropical Forest Research Centre, Atherton, Australia [email protected] Seed dispersing birds respond to local rainforest cover: Consequences for seed fate Most Australian rainforest plants are dispersed by fruit-eating birds. In partly deforested landscapes, seed dispersal within and between forest patches is influenced by these birds’ use of forest fragments, and by their patterns of fruit consumption. If the dispersers of a plant species decline or disappear, this will set an ultimate limit to the fate of its seeds – reduced dispersal. Therefore an important question for the conservation of rainforest plant communities is how to sustain or recover seed disperser abundances in fragmented forests. We assessed the effects of fragment size and surrounding forest cover on communities of seed disperser birds in an extensively-cleared Australian rainforest landscape, where different bird species vary in both their sensitivities to fragmentation and their roles as seed dispersers. In surveys of single one-hectare plots within 25 rainforest fragments, we recorded 20 seed disperser species. We used regression modelling to test how well particular seed disperser variables (species abundances, richness and abundance of functional groups) could be predicted by fragment size and six measures of surrounding forest cover (within 200m, 1000m and 5000m radii, for cover of rainforest and of all forest types). Model comparisons showed that the amount of rainforest cover within 200m was the best predictor of the abundances of fragmentation-sensitive disperser species, and of birds which disperse most plant species (including plants which have few dispersers). We conclude that a high proportion of local rainforest cover will help maintain seed disperser assemblages and seed dispersal in forest fragments and during forest restoration.

Muñoz, M.C.*1,2, Schaefer, H.M.3, Böhning-Gaese, K.1,2 & Schleuning M.1 1Biodiversity and Climate Research Centre (BiK-F) & Senckenberg Gesellschaft für Naturforschung, Frankfurt am Main, Germany, 2Department of Biological Sciences, Goethe Universität, Frankfurt am Main, Germany, 3Department of Evolutionary Biology and Animal Ecology, Faculty of Biology, University of Freiburg, Freiburg, Germany [email protected]

Linking mutualistic plant-frugivore networks and morphological traits to predict seedling establishment in the tropical Andes Frugivorous animals and fleshy-fruited plants form mutualistic networks of seed-dispersal interactions, one of the main ecological processes to maintain forest regeneration. Species traits affect mutualistic interactions, but little is known about the traits that influence seed-dispersal interactions and plant recruitment. The aim of our study was to identify functional traits of interacting species that promote seedling establishment. Along an elevational gradient (1800 to 2700 m) in the Colombian Andes, we carried out fieldwork in ten plots (100 x 20 m2). During 1200 hours of observation, we recorded fruit consumption on 21 fleshy-fruited plant species by 54 frugivorous bird species. We measured seed mass and counted the number of seeds of all consumed species, and recorded 11 morphological birds traits related to frugivory. In each plot, we set up 60 subplots (1 m2) to record seedling establishment. First, we tested which bird and plant traits influenced the frequency of plant–frugivore interactions. Second, we quantified the contribution of each bird species to seedling establishment and tested which traits were related to the seedling numbers in the study plots. We found that body mass and seed mass were the most important traits that determined fruit removal and the number of seedlings in the forest. Our preliminary results show that large birds promote seedling establishment, and plant species with large seeds have high chances of establishment. Thus, our findings suggest that body mass and seed mass are important functional traits that strongly impact forest regeneration in tropical mountain forests.

Naoe, S.1, Masaki, T.1 & Sakai, S.2 1Forestry and Forest Products Research Institute, Japan, 2Kyoto University, Japan [email protected] Temporal variation in community-level fruit abundance as a determining factor of seed dispersal of woody species that share frugivores In animal-dispersed plants, seed dispersal patterns of the same species in the same habitat can differ greatly temporally. However, very few studies have investigated the factors of temporal variation in seed dispersal. Our aim was to evaluate the effects of community-level and conspecific fruit abundance and frugivore phenology on seed dispersal by birds in a Japanese temperate forest. To this aim, we targeted six bird-dispersed woody species which share similar fruit characteristics but have different fruiting seasons, and investigated their fruit removal rate and seed dispersal distance for three years. The results indicated that annual and seasonal variation in community-level fruit abundance determined both fruit removal and the seed dispersal distance across species, but the effect differed with bird seasonality. Abundant fruit satiate fruit removal only in fruit-feeding seasons but not in insect-feeding seasons. A scarcity of fruits increased the dispersal distance, but only during the migratory season. This difference was probably due to the different foraging behaviors of passing migrants and territorial birds (i.e., permanent residents and summer birds). Our results illustrated that temporal variation in community-level fruit abundance could be a determining factor for seed dispersal in a temperate forest. The effects of temporal variation in community-level fruit abundance on seed dispersal patterns should be investigated in other ecosystems, particularly in other temperate regions, in order to understand their importance and associations with frugivore phenology.

Pérez-Méndez, N.*1, Jordano, P.1, García, C.2 & Valido, A.1 1Estación Biológica de Doñana (EBD-CSIC), Spain, 2Centro Investigação em Biodiversidade e Recursos Genéticos (CIBIO), Portugal [email protected] Defaunation-driven downsizing of frugivorous lizards determines genetic diversity and connectivity of Neochamaelea pulverulenta (Rutaceae) populations in the Canary Islands The distribution of genetic variation within species results from both historical and on-going ecological processes. How anthropogenic pressures affect this pattern remains a key goal of conservation biology. For instance, current extinction rates and the resulting downsizing of large-bodied frugivores may severely collapse the distribution of

61 genetic variation of plant species they feed on at the large scale, since large frugivores provide long-distance dispersal services. Thus, seed dispersal limitation is expected to hamper genetic connectivity causing genetic erosion within populations and genetic differentiation among populations. To test these hypotheses, we analysed the outcomes of the mutualistic interactions among Gallotia lizards (Lacertidae) and Neochamaelea pulverulenta (Rutaceae), a plant species relying on these frugivores for seed dispersal. The study was carried out on three islands with lizards differing in their body sizes: G. stehlini (max. snout to vent length: 280 mm, Gran Canaria), G. galloti (145 mm, Tenerife), and G. caesaris (111 mm, La Gomera). By applying population graph theory, we built a genetic network for each island based on the allelic frequencies of n= 80 populations. We tested the role of the defaunation process in determining population “hubs” that could act as a source of propagules, associated with the largest lizard species (Gran Canaria), and peripheral populations interacting with the less-effective, small-sized lizards. We tested for connectivity bottlenecks, whenever populations show differentiation well above the expectation based on isolation by distance, influenced by defaunation-driven downsizing. Our results suggested that defaunation-driven downsizing poses long-lasting genetic consequences for endozoochorous plant species.

Ramos, D.C.T.L.*1, Pizo, M.A.2 & Ribeiro, M.C.¹ 1Department of Ecology, São Paulo State University (UNESP), Rio Claro, Brazil, 2Department of Zoology, São Paulo State University (UNESP), Rio Claro, Brazil [email protected] Movement behavior of seed disperser birds in a gradient of forest loss Movements of bird dispersers connecting fragmented landscapes promote genetic diversity, colonization of new areas, and persistence of plant populations. However, little is known about how disperser behavior responds to different degrees of human changes on landscape structure and its consequences for quality and conservation of seed dispersal. In an on-going study, we aim to understand how movement behavior and use of different types of perches of four abundant bird seed dispersers change in a gradient of forest loss. We have completed 162 hours of direct observation of bird movements in six 600 x 600 m landscapes with different percentages of forest cover and cattle grazing pastures in the surroundings of a large Atlantic Forest corridor of Southeastern Brazil. Patagioenas picazuro and Dacnis cayana perch for longer periods and fly larger distances than Turdus sp. and Tangara sayaca, and may perform distinct functions. Birds perch longer and fly larger distances in low forest cover landscapes. The probability of perching in structurally simpler landscape features, like remnant trees and live fences, tended to increase with declining forest cover. Based on our results, as a consequence of changes in bird behavior, low forest cover landscapes may have farther seed dispersal and their functional connectivity may have an increased dependence on the distribution of stepping stones. We suggest that those differences should be considered in improving management practices for vegetation recovery and conservation in anthropic landscapes.

Schupp, E.W.1, Bennion, L.D.1, Gómez, J.M.2 & Jordano, P.3 1Utah State University, USA, 2Estación Experimental de Zonas Aridas, 3Estación Biológica de Doñana [email protected] Preliminary observations of dispersal and recruitment dynamics of Prunus fasciculata, a dry‐ fruited native almond of the SW United States This poster presents preliminary observations on dispersal and recruitment of a widespread, patchily distributed native almond of the southern Great Basin and the Mojave Desert, USA. This dioecious species grows in clumps along intermittent washes (arroyos, wadis). Goals at this phase of investigation are to: quantify interactions of rodent species with seeds, evaluate seed traits and germination, and investigate the origin of shrub clumps. Observations with game cameras suggest that the diurnal rodent Ammospermophilus leucurus is the most frequent interactor with seeds. Squirrels climb and harvest seeds directly from the shrub. Many are consumed in place but caches are also made. Seeds, which are not produced every year, are 10–12 mm long and weigh about 0.9 g. Unlike

62 commercial almonds, the exocarp remains intact enclosing the seed. About 72% of seeds collected from shrubs are filled and appear healthy. Following cold–moist stratification, 39% germinated. However, exocarps inhibit germination, for seeds with the exocarp intact, germination was 30%. It is not yet known whether rodents husk seeds before caching them. To make an initial assessment of the origin of clumps, leaf tissue from 4–13 individual branches was sampled from four clumps. Samples are being sequenced to determine whether clumps are single genetic individuals or represent multiple genetic individuals. Results will inform more detailed studies in the future. Although preliminary, results suggest this is a fascinating species for investigating the relative roles of biotic interactions (e.g. dispersal, facilitation) and abiotic conditions (e.g. microtopographic position) driving patterns of recruitment of a desert shrub.

Shi, L., Lin, Y. & Wang, Z. College of Animal Science, Xinjiang Agricultural University, China [email protected] Is the Turpan Wonder Gecko Teratoscincus roborowskii (Squamata, Gekkonidae) a legitimate seed disperser of a desert shrub Capparis spinosa (Capparaceae) In this study we determined the effect of seed passage through Teratoscincus roborowskii digestive tracts on germination of the fleshy-fruited desert shrub species Capparis spinosa (Capparaceae), and evaluated the effect of this passage on seed coat characteristics. In addition, we assessed the spatial patterns of faecal deposition by lizards onto various microhabitats available in the desert environments of the Turpan Basin, Western China. An assessment of seeds following passage through the gut of lizards indicated that seeds remained intact and viable and that the proportion of viable seeds from lizard faeces was not significantly different from that of seeds collected directly from fruits. The germination rate and the final percentage of lizard-ingested C. spinosa seeds were greater than control seeds. Comparing scanning electric photographs and seed coat histological sections of control and passage seeds with different retention times, we suggest that seed coats from lizard-ingested seeds were probably thinning and softening by abrasion inside the lizards’ digestive tract. Twenty-seven percent of the lizard’s faeces were deposited on bare soil near dead wood commonly inhabited by lizards. However, this microhabitat represents only 3% of the available ground cover at the study site. By enhancing seed germination and depositing intact and viable seeds onto potential safe sites for recruitment, the lizard Teratoscincus roborowskii is acting, at least qualitatively, as an effective disperser of Capparis spinosa.

Smith-Ramirez, C.1, Mora, J.P.2 & Zapata, V.M.*3 1Ecology and Biodiversity Institute (IEB-Chile), 2Science Faculty, Austral University of Chile, 3Department of Ecology and Hidrology, University of Murcia, Spain [email protected] Wind as a legitimate seed dispersal agent of endozoochorus fruits on Robinson Crusoe Island, Chile We ask whether fleshy fruits could be dispersed by wind in areas where it is strong and frequent, how frequently wind dispersal occurs compared to bird dispersal and whether wind could be a legitimate seed dispersal agent. Between January 2011 and June 2010 (17 months), 340 to 282 seed traps were placed in a forest stand on Crusoe Island. The traps were placed in four environmental types: invasive scrubland, forest, forest border with canopy gaps, and forest gaps. We considered that fruits fallen in traps within forest gaps are transported by wind since there is no canopy above. We assume that fruits of native species fallen in scrublands, and the exotic fruits fallen in forests are moved by wind. We found that 79.9% of the fruits fallen into traps in forest gaps were fleshy fruits the others were dry fruits. In total Turdus (the only dispersing bird on Crusoe Island) moved 73.5 % of the seeds gathered in the traps, the other seeds remaining in the fruits were moved by wind. We found differences in the total seeds, whether endemic or exotic, and species numbers transported by wind or birds. The birds dispersed seeds mostly from heavy fruits. We sowed seeds (defecated by Turdus and dispersed by wind) of three species. We found that wind was a

63 legitimate seed dispersal agent of two of them. We conclude that in windy places, wind could be a legitimate dispersal agent of some fleshy fruit seeds, and as important as birds.

Strasberg, D.1, Flores, O.1 & Thébaud, C.2 1UMR PVBMT, Université de La Réunion, France, 2Laboratoire EDB, Université Toulouse III Paul Sabatier, France [email protected] Lack of recovery of native fleshy fruited species following large-scale disturbance on Reunion (Mascarene archipelago) On the island of Réunion (Mascarene archipelago), a lack of recovery of native fleshy fruited plant community was observed on a chronosequence produced by lava flows of the active Piton de la Fournaise volcano. We compared fruit characteristics, species composition and abundance of plant communities growing on different aged lava flows (8 to about 500 yrs old). Results showed that wind-dispersed plants dominate the young lava flows, less than 70 yrs old whereas 85% of woody plants are fleshy-fruited species in old-growth forests on about 500 yrs old lava flows. Native woody plants with medium and large fleshy fruits (length > 20 mm) are absent from lava flows younger than 200 yrs. Data confirm that historical extinction of native frugivores in the Mascarenes limits dispersal of most native trees. On sites previously surveyed in the 1970s and the 1990s, species composition and abundance measurements showed that lava flows initially dominated by native wind dispersed species are rapidly colonized by exotic fleshy- fruited species. The dispersal of the latter species is facilitated by the introduced red-whiskered bulbul Pycnonotus jocosus that invaded the island in the 1990s. We conclude that the drastic changes in frugivores community composition alter succession processes and preclude the recovery of native plant communities after large-scale disturbances.

Takahashi, K.1 & Takahashi, K.2 1Nagano University, Japan, 2Shinshu University, Japan [email protected] Spatial distribution and size of small canopy gaps created by the Japanese Black Bear The Japanese Black Bear, a large-bodied frugivore, was believed to frequently create small gaps in the tree crown during fruit foraging. However, there were no authenticated reports of black bear-created canopy gaps. To characterize physical canopy disturbance by black bears, we examined a number of parameters, including the species of trees in which canopy gaps were created, gap size, the horizontal and vertical distribution of gaps, and the size of branches broken to create gaps. The size of black bear-created canopy gaps was estimated using data from branches that had been broken and dropped on the ground. The disturbance regime was characterized by a highly biased distribution of small canopy gaps on ridges, a large total overall gap area, a wide range in gap height relative to canopy height, and diversity in gap size. Surprisingly, the annual rate of bear-created canopy gap formation reached 141.3 m2 ha–1 yr–1 on ridges, which were hot spots in terms of black bear activity. This rate was approximately 6.6 times that of tree-fall gap formation on ridges at this study site. Furthermore, this rate was approximately two to three times that of common tree-fall gap formation in Japanese forests, as reported in other studies. Our findings suggest that the ecological interaction between black bears and fruit-bearing trees may create a unique light regime, distinct from that created by tree falls, which increases the availability of light resources to plants below the canopy.

Tsuji, Y. Primate Research Institute, Kyoto University [email protected] Estimate of the seed shadow created by wild Japanese macaques as seed dispersers in Japanese cool-temperate forest I evaluated the inter-yearly difference in distances of seed dispersal by a group of wild Japanese macaques (Macaca fuscata), a species of cercopithecine in the temperate forest in Japan, in order to address effect of inter-annual

64 variation in fruiting of staple foods on the dispersal distance, one of the characteristics of endozoochory. Since we could conduct continuous observation from dawn to dusk, and gut passage time of the macaques was known, dispersal distance of seeds could be obtained by recording the locations of feeding and ranging patterns of the target troop, with the straight distance between the two locations being the dispersal distance. The mean (± SD) dispersal distances of 11 sap fruit species by the macaques were 487 ± 264 m (range: 8–1246 m) and inter- yearly/seasonal variation was not found. The present study provides the effect of inter-annual difference in fruiting, and consequent effect on macaque ranging behaviour on the dispersal distance of seeds by the macaques, an important aspect of endozoochorous seed dispersal in the temperate region. The independent bout method that we employed could easily provide essential information contributing to estimation of seed dispersal patterns caused by the subject animals. Since there are many study sites at which researchers can observe habituated animals, this method should be applied in other areas to study seed dispersal by the primates.

Vogeler, A.V.1, Bohning-Gaese, K.2,4, Vollstaedt, M.2 & Tschapka, M.1,3 1University of Ulm, Institue of Evolutionary Ecology and Conservation Genomics, Germany, 2Biodiversity and Climate Research Centre (BiK-F) and Senckenberg Gesellschaft für Naturforschung, Germany, 3Smithsonian Tropical Research Institute, Balboa, Panama, 4Johann Wolfgang Goethe University of Frankfurt, Department of Biological Sciences, Germany [email protected] Habitat use of pteropodid bats and their importance as long-distance seed dispersers on the slopes of Mt. Kilimanjaro, Tanzania Although frugivorous bats have been frequently identified as important pollen and seed dispersers, little information is available on the role of the fruit bats as seed dispersers in Africa, including on how land-use changes may affect this ecosystem service. Here, we examined seed contents in fecal samples of six pteropodid bat species (Eidolon helvum, Epomophorus minimus, Epomophorus wahlbergi, Rousettus aegyptiacus, Rousettus lanosus and Lissonycteris angolensis) captured in five different habitat types (maize, savanna, home-garden, coffee plantation, lower mountain forest) on Mt. Kilimanjaro, Tanzania. Additionally, we used an experimental setting in which fruit bats were kept in a flight cage to investigate gut passage and retention time of seeds. Animals were fed and subsequently filmed as well as directly observed to assess transit times of seeds over a 36-hour period. Preliminary analyses show distinct differences among fruit bat communities in the different habitat types, which indicates that human land use modifies community composition of these keystone species. In most fecal samples we found Ficus spp. (Moraceae) seeds. Maximum retention of viable Ficus seeds in E. wahlbergi exceeded 12 hours. Our study stresses the importance of fruit bats as seed dispersers on Mt. Kilimanjaro, in particular for Moraceae. Their high degree of mobility in combination with seed retention times of more than 12 hours suggests a high potential for long distance seed dispersal. They also link different habitat types. Therefore the changes in community composition caused by habitat degradation may have negative consequences for seed dispersal by fruit bats.

Vollstädt, M.1, Schleuning, M.1, Tschapka, M.2,3, Vogeler, A.2 & Böhning-Gaese, K.1,2 1Biodiversity and Climate Research Centre (BiK-F) and Senckenberg Gesellschaft für Naturforschung, Germany, 2University of Ulm, Institute of Evolutionary Ecology and Conservation Genomics, Germany, 3Smithsonian Tropical Research Institute, Balboa, Panamá, 4Johann Wolfgang Goethe University of Frankfurt, Department of Biological Sciences, Germany [email protected] The modularity of a seed-dispersal network on the slopes of Mt. Kilimanjaro, Tanzania Climate change and the increase in land-use intensities threaten the functioning of ecosystems worldwide. It is not fully understood to what extent these anthropogenic changes influence the dynamics of mutualistic interaction networks. There are only few studies investigating the effects of climate and land-use on interaction networks. Our study is the first to examine the modular structure of a seed-dispersal network both along an elevational and a land-

65 use-gradient. In our on-going study, we investigate an interaction network of frugivorous birds and fruiting plant species on Mt. Kilimanjaro, Tanzania. We investigate plant–frugivore interactions in 12 different habitat types (ranging from pristine to highly disturbed habitats) and at 6 elevational belts (800–4200 m asl). Here, we analyze the modularity of this large seed-dispersal network, modularity is defined by the presence of modules comprising highly interlinked subsets of species that are separated from the rest of the network. Our preliminary results show that the seed-dispersal network on Mt. Kilimanjaro is organized into different modules. More specifically, our results show that these distinct modules correspond to specific habitat types, comprising modules of species occurring in specific forest types and in habitats outside the forest belt. The modular structure of the seed-dispersal network suggests that the different habitat types on Mt. Kilimanjaro represent spatially distinct sub-networks which are only weakly interconnected by frugivorous bird species moving between the respective habitat types.

Wróbel, A.* & Zwolak, R. Adam Mickiewicz University, Faculty of Biology, Department of Systematic Zoology, Poland [email protected] A comparison of acorn dispersal by different rodent species: does size of the disperser matter? Most zoochoric plants rely on multiple species of animals for seed dispersal, these dispersers often vary in their influence on plant recruitment. We used selective rodent exclosures to determine the impact of different groups of rodents on the fate of red oak (Quercus rubra) acorns. The rodents were divided into 1) small-sized (Peromyscus sp. and Myodes gapperi), 2) the previous group plus medium-sized (Tamias striatus andGlaucomys volans), and 3) the two previous groups plus large-sized rodents (Sciurus carolinensis). Acorn removal rates were significantly lower for the small-sized group relative to the medium- and large-sized groups. When only small-sized species had access, a significantly higher proportion of removed acorns was never found (48% vs. 15% for medium-sized and 38% for large-sized), probably reflecting frequent larder-hoarding. In comparison to other groups, the small-sized group cached much fewer (0.06% vs. 48% for medium-sized and 40% for large-sized) and partially ate more removed acorns (91% vs. 41% for both medium-sized and large-sized) than the other groups. The average acorn dispersal distance was the longest for the medium-sized treatment (2.5 m ± 0.16 SE), intermediate for the large-sized treatment (2.0 m ± 0.16 SE), and the shortest for the small-sized treatment (1.8 m ± 0.15 SE). Considerably lower proportion of cached acorns and shorter distances of dispersal indicate that small-sized rodents provide low quality acorn dispersal relative to medium and large-sized rodents. This finding provides strong support for the notion that scatter-hoarders are not equal in their effects on plant reproduction.

Zaniratto, C.P.1, Costa, J.C.2 & Silva, W.R.3 1Graduate Program in Ecology, University of Campinas, Brazil, 2Costa & Faria Agro-Environmental Consultancy, Piracicaba, 3Brazil, Dept. Animal Biology, University of Campinas, Brazil [email protected] Dense understory and frugivorous bird species increase seed rain in a restoration physiognomic mosaic of Atlantic Forest in Southeast Brazil. Zoochory plays an important role in the dynamics of plant communities, allowing the colonization of new areas and ensuring the maintenance of tropical diversity. Compared to terrestrial frugivores, birds are more efficient seed dispersers due to their high mobility and capacity of dispersing most of the consumed seeds without damage. These characteristics make birds important agents in forest restoration, as they can improve seed deposition by transporting seeds from well-preserved places to degraded ones. We evaluated how the presence of frugivorous birds and vegetation structure locally influence the seed rain in a restoration physiognomic mosaic of Atlantic Forest in São Paulo state, southeast Brazil. The seed rain was collected year-round by seed traps set in 20 sampling points randomly distributed in the study area. At each point we recorded birds in 30 m fixed-radius point counts, as well as the richness of ornithochorous plants, canopy opening and three ranges of understory density. The effect of these variables on the seed rain was evaluated by model selection analysis. Seed richness was positively related to bird

66 richness and understory density. These results suggest that the ornithochorous seed rain can be influenced, at the local scale, both by the presence of frugivorous birds and the structural complexity of the vegetation. Thus, recovery of the understory is an essential step in the restoration process, leading to a richer bird assembly and, consequently, to a more diverse plant assembly.

Zapata, V.*1, Martínez-López, V.1,2 & Robledano-Aymerich, F.1 1Department of Ecology and Hydrology, University of Murcia, Spain, 2Department of Zoology and Anthropology, University of Murcia, Spain [email protected] Assessment of bird dispersal potential for the passive restoration of Mediterranean semiarid old fields from forest source patches Dispersal by bird frugivores has been scarcely studied in semiarid Mediterranean areas, and hence few evidences exist of passive restoration based on this ecosystem service. As part of a study about old field recovery in Southeastern Spain, we assessed the dispersal potential of fleshy-fruited shrubs from forest remnants to nearby disturbed habitats (old fields, dwarf shrublands, dry pasture). Different approaches, scales and methodologies (e.g. bird netting and censuses, estimations and observations of seed removal, genetic studies) have been used. The main aims were 1) to assess bird-mediated seed dispersal flow within and around forest source patches, and to identify the factors limiting seed export, 2) to evaluate the performance of seed dispersal towards old fields and its relationship with bird assemblages, through multispecies indices based on their disperser character (legitimate vs illegitimate), and 3) to propose guidelines for the passive or assisted recovery of disturbed areas, based on bird dispersing activity. Both internal regeneration and old- field colonization by shrubs were favored by availability of perches (dead or decaying trees) and facilitative effects. The main limiting factors were lack of seed sources, the scarcity of favorable regeneration microsites, competition by earlier colonizers, weather, and probably herbivory. Furthermore, forest areas can display a limited seed export, which can also be negatively affected by nearby urban gardens acting as sinks. Though difficult, in semiarid areas passive regeneration can benefit from bird dispersal services, provided that dispersal filters are removed or mitigated, and that critical elements (e.g. natural or artificial perches, microsites, water) are wisely managed.

Acknowledgements

The FSD 2015 Local Organising Committee are most grateful for the financial support received from the following organisations:

 University of KwaZulu-Natal

 Oppenheimer & Sons

 Centre for Invasion Biology (CIB) DST-NRF Centre of Excellence for Invasion Biology, University of Stellenbosch

 National Research Foundation

Summary list of delegates

Surname Name Affiliation E-Mail

Acosta Rojas Diana Universidad de los Andes [email protected] Carolina

Albert- Aurélie Muséum National d’Histoire Naturelle-UMR 7179 CNRS-MNHN [email protected] Daviaud Ally Ebrahim University of KwaZulu-Natal, Pietermaritzburg [email protected]

Baltzinger Christopher Irstea [email protected]

Bello Carolina Universidade Estadual Paulista UNESP [email protected] Bender Irene German Centre for integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig [email protected]

Blendinger Pedro CONICET & Instituto de Ecología Regional, Universidad Nacional de Tucumán [email protected]

Boettinger Janis Utah State University [email protected] Bogdziewics Michal Department of Systematic Zoology, Adam Mickiewicz University in Poznań michalboqdziewicz@qmail,com Böhning- Katrin Senckenberg Biodiversity and Climate Research Centre [email protected] Gaese Borges Renee Centre for Ecological Sciences, Indian Institute of Science [email protected]

Buzenac Jacqueline Muséum National d’Histoire Naturelle-UMR 7179 CNRS-MNHN [email protected] Campos Claudia Instituto Argentino de Investigaciones de las Zonas Áridas (IADIZA) [email protected] Cestari Cesar UNESP [email protected] Chapman Hazel University of Canterbury [email protected] Chen Si-Chong University of New South Wales [email protected] Chibesa Moses University of KwaZulu-Natal, Pietermaritzburg [email protected] Christianini Alexander Universidade Federal de Sao Carlos [email protected]

Clausi Bernardo Myrtlau, Brazil [email protected] Culot Laurence Universidade Estadual Paulista (UNESP) [email protected] Davenport Lisa Duke University Center for Tropical Conservation [email protected]

Downs Colleen University of KwaZulu-Natal, Pietermaritzburg [email protected] Drabik- Martyn University of KwaZulu-Natal, Pietermaritzburg [email protected] Hamshare Effiom Edu Lund University/Cross River State Forestry [email protected]

Ehlers Smith Yvette University of KwaZulu-Natal, Pietermaritzburg [email protected] Ehlers Smith David University of KwaZulu-Natal, Pietermaritzburg [email protected]

Escribano- Gema Universidad Técnica Particular de Loja [email protected] Avila Falcón Wilfredo Institute of Evolutionary Biology and Environmental Studies,University of Zürich [email protected] Farwig Nina University of Marburg, Faculty of Biology - Conservation Ecology [email protected] Fedriani Jose Maria Instituto Superior de Agronomia, Universidade de Lisboa [email protected] Laffitte Fonturbel Francisco Universidad de Chile [email protected] Forget Pierre- Muséum National d’Histoire Naturelle [email protected] Michel

Fricke Evan University of Washington [email protected] Galetti Mauro Departamento de Ecologia, Universidade Estadual Paulista [email protected] García Daniel University of Oviedo [email protected] / [email protected] Goldson S.

Gomez Daniella CIT – Jujuy, Cátedra de Biodiversidad y Desarrollo Sustentable (Univ. Nac. de [email protected] Jujuy) Goncalves Cristina UNESP [email protected] González-Varo Juan Estación Biológica de Doñana [email protected]

Gordon Katherine University of Cape Town [email protected]

Govender Keeveshnee University of KwaZulu-Natal, Pietermaritzburg [email protected] Green Ronda Environmental Futures Research Institute, Griffith University [email protected] Hansen Dennis Institute of Evolutionary Biology and Environmental Studies, University of Zürich [email protected] Hart Lorinda University of KwaZulu-Natal, Pietermaritzburg [email protected]

Haugaasen Torbjǿrn Norwegian University of Life Sciences [email protected]

Haurez Barbara Laboratory of Tropical Forestry University of Liège, Passage des Déportés, 2, [email protected] 5030 Gembloux Hui Cang Stellenbosch University [email protected]

Jansen Patrick Wageningen University & Smithsonian Tropical Research In [email protected]

Johnson Steven University of KwaZulu-Natal, Pietermaritzburg [email protected]

Jones Landon University of Louisiana Lafayette [email protected]

Jordaan Maryna University of KwaZulu-Natal, Pietermaritzburg [email protected]

Kaplin Beth Antioch University New England and University of Rwanda [email protected] Kelly Dave Biological Sciences, University of Canterbury, [email protected]

Kitamura Shumpei Ishikawa Prefectural University [email protected] Kromann- Anders MEME (Management and Ecology of Malaysian Elephants) [email protected] Clausen

Kuhlmann Marcelo University of Brasilia [email protected] Kwit Charles University of Tennessee [email protected] Loayza-Freire Andrea Universidad de La Serena, Instituto de Ecología y Biodiversidad [email protected] Marques- Caroline Muséum National d’Histoire Naturelle [email protected] Dracxler Martínez Vincente University of Murcia [email protected] López

Martins Milene Universidade Federal de Sao Carlos [email protected] McConkey Kim NIAS [email protected]

MacFadyen Duncan E. Oppenheimer & Son [email protected] Midgley Jeremy University of Cape Town [email protected] Mokotjomela Thabiso University of the Witwatersrand [email protected] Moran Catherine Griffith University & James Cook University & CSIRO, Australia [email protected] Mueller Thomas Senckenberg Biodiversity and Climate Research Centre & Goethe University [email protected] Frankfurt Munoz-Neyra Marcia Biodiversity and Climate Research Centre (BiK-F) [email protected]

Nathan Ran The Hebrew University of Jerusalem [email protected]

Neuschulz Erika Lena Senckenberg Biodiversity and Climate Research Centre Frankfurt [email protected] Niebuhr Bernardo Departamento de Ecologia, Universidade Estadual Paulista [email protected] Santos Olsson Ola Lund University [email protected]

Pedrosa Felipe Departamento de Ecologia, Universidade Estadual Paulista (UNESP) [email protected] Pérez-Méndez Nestor Estación Biológica de Doñana (EBD-CSIC) [email protected] Pfeiffer Morgan University of KwaZulu-Natal, Pietermaritzburg [email protected] Pillay Kerushka University of KwaZulu-Natal, Pietermaritzburg [email protected]

Pizo Marco Departamento de Zoologia, UNESP [email protected]

Prasad Soumya Jawaharlal Nehru University [email protected] Ramos Danielle Sao Paulo State University [email protected] Rane Ulhas Centre for Ecological Sciences, Indian Institute of Science [email protected]

Reynolds Chevonne University of Cape Town, Percy Fitzpatrick Institute of African Ornithology [email protected] Richardson Dave Centre for Invasion Biology, University of Stellenbosch [email protected]

Riddhika Kalle University of KwaZulu-Natal, Pietermaritzburg [email protected]

Rogers Haldre Rice University [email protected] Ruggera Román CIT – Jujuy, Cátedra de Biodiversidad y Desarrollo Sustentable (Univ. Nac. de [email protected] Jujuy) Schleuning Matthias Senckenberg Biodiversity and Climate Research Centre [email protected]

Schupp Eugene Utah State University [email protected] Shi Lei Xinjaing Agricultural University [email protected]

Shuttleworth Amy-Leigh University of KwaZulu-Natal, Pietermaritzburg [email protected] Silva Wesley University of Campinas [email protected] Singh Preshnee University of KwaZulu-Natal, Pietermaritzburg [email protected] Smith Cecilia Universidad Austral de Chile [email protected]

Solorzano- Jorge A. Faculty of Forestry, University of Toronto [email protected] Filho

Sridhara Sachin James Cook University [email protected]

Strasberg Dominique Reunion University [email protected]

Summers Mark University of KwaZulu-Natal, Pietermaritzburg [email protected] Takahashi Kazuaki 658-1 Shimonogo, Ueda, Nagano [email protected] Terborgh John Center for Tropical Conservation, Duke University [email protected] Thabethe Vuyisile University of KwaZulu-Natal, Pietermaritzburg [email protected] Tharmalingha Ramesh University of KwaZulu-Natal, Pietermaritzburg [email protected] m Timoteo Sergio Centre of Functional Ecology, Department of Life Sciences, University of [email protected] Coimbra Tjelele Julius Agricultural Research Council, South Africa [email protected]

Torres Caba Jahdiel Centre for Tropical Ecology and Conservation, Antioch University, New England [email protected] Traveset Anna IMEDEA, Spanish Research Council [email protected] Trolliet Franck University of Liege [email protected] Tsuji Yamato Primate Research Institute, Kyoto University [email protected]

Turke Manfred German Centre for integrative Biodiversity Research (iDiv) Halle-Jena-Leipzig, [email protected] Valido Alfredo Estación Biológica de Doñana (EBD-CSIC) [email protected]

Vogeler Anna University of Ulm, Institute of Evolutionary and Conervation Genomics [email protected] Vollstädt Maximilian Biodiversity and Climate Research Centre (BiK-F) and Senckenberg Gesellschaft [email protected] für Naturforschung Wang Bo Xishuangbanna Tropical Botanical Garden, Chinese Academy Sciences [email protected]

Westcott David CSIRO [email protected] White Joseph Department of Biological Sciences, University of Cape Town [email protected] Wotton Debra Moa’s Ark Research and University of Canterbury [email protected] Wróbel Aleksandra Uniwersytet im. Adama Mickiewicza [email protected]

Zapata Pérez Víctor University of Murcia Spain [email protected] Manuel Zungu Manqoba University of KwaZulu-Natal, Pietermaritzburg [email protected] Zwolak Rafal Department of Systematic Zoology, Adam Mickiewicz University in Poznań [email protected]

How to attend a scientific conference to your best advantage

John Cooper and Les G Underhill Animal Demography Unit, Department of Zoology University of Cape Town, Rondebosch, 7701 South Africa

The golden rule of conference attendance is simple. Escape the bubble of your cozy group of colleagues. Keep a degree of space between yourself and the people you can interact with at home when you are not at the conference, so as to allow time to meet new people, have useful conversations with them and hopefully establish long-lasting professional relationships. However, do touch sides with your own people, if only for de-briefing, at least once a day. In this way you will be able to act as a colleague should, introducing your new acquaintances and friends (and meeting theirs), and mooting joint research programmes and the like.

If you do form a group from among your colleagues, for example to have lunch at the conference venue, try every time to invite one or two of your new acquaintances to join you. There are always conference attendees without home colleagues who will welcome such invitations. Much good business comes out of the informal conversations that take place at such times: even sitting beside the right person on the bus can pay dividends!

Although you may think the most important thing at a conference is attending as many talks and poster sessions as feasible, and making your own presentation, the really most important business of all is transacted outside the formal sessions. So do not feel you are letting your sponsor down by missing a talk or two to hold or to continue a conversation which you think will be important to you, either by gaining specific knowledge you think you will need, or by establishing a good contact who may help with your future career, or next piece of planned research. Important things happen “in the margins” of the conference. This is where you may find yourself a PhD supervisor, a postdoctoral position, where you find new research partners, where you seek out invitations for visits to research institutes, where you find yourself a sabbatical position, where you find yourself a job, where you make yourself available for election to the executive committee, where you offer to organize the next conference, where you find PhD students to supervise and postdocs to mentor, where you find staff, where you find examiners for PhD theses, etc. If you don’t come home from the conference both exhausted but also fulfilled, and with a notepad full of new contacts and ideas, something probably went wrong somewhere. However, missing sessions for activities such as bird-watching and shopping is unethical, unless you are paying your own way completely out of your own pocket. Although you are may not hear an adverse comment, your absences do not go unnoticed.

If there are parallel sessions, choose your session carefully. It often is a disaster to move between parallel sessions, because it is quite rare for the conference timekeeping to be good enough to make this reliable. If there are no talks to go to that particularly interest you, go and learn about something new. Look at the posters carefully. If there is one that interests you, search out the owner and discuss it with her/him.

When you get your conference pack, look carefully through the name and address list of people attending. If you see that people whom you want to meet are attending, make a list of them and search them out. You can do this in at least two ways: look at the name badges, or ask people if they can point out ABC to you. If you have read one of their papers, you can say: “Hi, I’m DEF, and I found your paper in HIJ on XYZ very interesting and helpful.” That little bit of flattery ought to get you into conversation. If these high fliers are abrupt with you, it is their problem, not yours. You really need to work hard, right from the start of the conference, to meet all the people you want to talk to. Ask your colleagues, and especially your supervisor, to make introductions for you. If you tend to be on the shy side, this is a good way to get to meet new people, especially the more senior ones. A good supervisor or senior

72 colleague will make the introduction, start the conversation going with a statement on what interesting work you have been doing, ask you a question she/he already knows you will be able to answer well, perhaps make an interjection or two, then excuse him/herself, leaving you well introduced and feeling confident to continue the conversation on your own.

If you have a laptop with you, transfer the notes on paper into a file of things to do once you get home, organized in some more rational way than the sequential paper notes: ideas; emails to send off as soon as you get home (many of these ought to be thank you emails, to anyone who was helpful in some way – ideas, discussions, took you out for a meal, future invitations, gave you a reprint, etc). Small courtesies pay long-term dividends.

If you are a MSc/PhD student and your supervisor or other sponsor has paid for your conference costs, what does he or she expect of you? Primarily, you should do your presentation to the best of your ability:

1. Meet your session chair in advance, preferably the previous day. Ask for an introduction if you do not know her/him. Ensure the chair knows the facts about you and your work which you would like to be made public. Hand in your presentation as requested – listen carefully for the instructions about this. Check that it projects properly yourself, do NOT leave this to anyone else. Be ready for a problem with the audio-visual equipment and do not panic if your illustrations do not immediately come up on the screen, just solve the problem, or ask for help, and then proceed.

2. Keep to time. Practice if you feel you need to ensure this. Present your talk to your colleagues at home first and listen to their criticisms and act on them.

3. Speak slowly and distinctly and avoid slang, jokes and irrelevant “wake-up” pictures at all costs. In general, most members in your audience will not have English as their home language. Be respectful of them. If you use a term specific to your discipline, explain parenthetically. For example: “I measured the width of the parietal bone, a bone at the front of the skull…”. Explain abbreviations the first time they are used. Stick to simple English words, and stick to one term for each concept: no need to use “elegant variation”. An example: “tail feather” is much better than ”rectrix”. Don’t use slang. It is “kilometres”, not “kims” or “kays”; such loose talking is unprofessional and disrespectful to the audience.

4. If you need to use the terms for the seasons, you need to make it clear which hemisphere you are in – this is particularly important if you are doing a “southern hemisphere” talk and most of your audience is from the northern hemisphere. You need to make it clear then that you are talking about the southern summer, because when you say “summer”, northern hemispherists think “June, July and August”. Technically the northern seasons are boreal and the southern seasons are austral, so you can take about the austral summer, but even this is not widely understood in the northern hemisphere. Best of all, try to paraphrase round these confusions.

5. Check your voice volume with a colleague in the empty lecture room in advance. She/he is in the back row, can you be heard clearly? If you know you have a tendency to speak softly, or mumble, this is critically important.

6. It does not matter if you are nervous, everyone in the audience will be nervous when they get up to present, irrespective of their seniority and experience. If you feel extremely nervous at the start, take a drink of water to avoid a dry throat and tell the audience: “This is my first talk to an international audience and I’m a bit nervous, but I am excited about my work and hope you will be too”. Have your starting sentence ready in your head, word perfect. It is best to avoid reading from a prepared text. If you feel you really have to do this, then from time to time, lift up

73 your head, look directly at the audience and make a few statements that are clearly not in your notes. This will help establish rapport. Far better is to have a few words to prompt you on the laptop adjacent the projected illustration.

7. Start off with your second-best story and picture, and end on your best. In between be succinct and to the point. Avoid “trailing off” your sentences. At all times never end your talk with something like “…and that’s all I have to say”, end positively with thanking the audience for their attendance. When you start, if you feel the session chair has not introduced you properly by all means say who you are, but briefly. At the end avoid long-winded thanks and acknowledgements, they should rather be on your closing picture. Just say I owe a big thanks to my supervisor/colleague Professor John Bloggs and to the Myrtle Aspidastra Bursary Fund.”

8. Nowadays we have our illustrations on a lap top in front of us. Use this as your prompt. There is no reason, at any time, to turn and face the screen and talk half-turned from, or worse, with your back to the audience. Never walk up to screen and make it move by touching it, neither try to get you hand to cast a shadow on the salient figure. If you need to use a pointer, use a mouse. Laser pointers tend to exaggerate your nervously shaking limb and annoy the audience. Well-designed illustrations should not need a pointer, rather build the picture in steps to make your points in a logical order. Make sure from your speaking position that all will be able to see your pictures, and check that the writing is legible from the back row. Don’t overload your pictures with detail so you have to say “Please ignore all the columns except the seventh”. This suggests to the audience you have borrowed a picture from a previous talk at best, and, at worse, that you are not very good..

9. Know exactly how you are going to end.

Conferences in 2016 you may be interested in

 BOU Urban Birds conference, Leicester, UK , 5 – 7th April 2016

 17th International Bat Research Conference, Durban South Africa, 31st July- 5th August 2016

 Pan African Ornithological Congress 14, Dakar, Senegal, 17-23rd October 2016