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«Doctoral thesis for a doctoral degree at the Graduate School of Life Sciences, JuliusMaximilians-Universität Würzburg, Section Integrative Biology ...»

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The impact of logging and conversion to oil palm plantation on

Bornean stream-dependent frogs and their role as meso-predators

Die Auswirkungen von Abholzung des Regenwalds und Umwandlung zu

Ölpalmplantagen auf bachbewohnende Frösche und ihre Rolle als

Mesoprädatoren in Borneo

Doctoral thesis for a doctoral degree at the Graduate School of Life Sciences, JuliusMaximilians-Universität Würzburg, Section Integrative Biology

submitted by

Oliver Konopik


Lohr am Main

Würzburg 2014 Submitted on:……………………………………………………………………………

Members of the Promotionskomitee:

Chairperson: Prof. Dr. Thomas Dandekar Primary Supervisor : Prof. Dr. Ingolf Steffan-Dewenter Supervisor (Second): Prof. Dr. Hans-Joachim Poethke Supervisor (Third): Prof. Dr. Ulmar Grafe Date of Public Defence:………………………………………………………………… Date of Receipt of Certificates:………………………………………………………… Slow loris (Nycticebus menagensis), Sabah (Malyasia) 2014 “What escapes the eye … is a much more insidious kind of extinction: the extinction of ecological interactions” (Daniel H. Janzen) Table of content Table of content Affidavit

Curriculum Vitae

Publication list

Summary (English)

Zusammenfassung (German)

I. General introduction


II. Effects of logging and oil palm expansion on stream frog communities on Borneo, Southeast Asia

III. Logging and conversion to oil palm plantations alters trophic interactions and reduces multiple levels of anuran diversity

IV. From rainforest to oil palm plantations: shifts in predator population and prey communities, but resistant interactions

V. General discussion


Author contributions


Affidavit Affidavit I hereby confirm that my thesis entitled “The impact of logging and conversion to oil palm plantation on Bornean stream-dependent frogs and their role as meso-predators” is the result of my own work. I did not receive any help or support from commercial consultants. All sources and / or materials applied are listed and specified in the thesis.

Furthermore, I confirm that this thesis has not yet been submitted as part of another examination process neither in identical nor in similar form.

–  –  –

Hiermit erkläre ich an Eides statt, die Dissertation “The impact of logging and conversion to oil palm plantation on Bornean stream-dependent frogs and their role as meso-predators” eigenständig, d. h. insbesondere selbständig und ohne Hilfe eines kommerziellen Promotionsberaters, angefertigt und keine anderen als die von mir angegebenen Quellen und Hilfsmittel verwendet zu haben.

Ich erkläre außerdem, dass die Dissertation weder in gleicher noch in ähnlicher Form bereits in einem anderen Prüfungsverfahren vorgelegen hat.

–  –  –

DATE OF BIRTH December 18th, 1984 PLACE OF BIRTH Lohr am Main, Germany 2011-2014 PhD student at the Department of Animal Ecology and Tropical Biology, Julius-Maximilians-University of Würzburg 2005-2010 Studies in Biology, Julius-Maximilians-University of Würzburg

–  –  –

2005 Alternative service at the NABU-Centre Wollmatinger Reed, Konstanz 2004 Abitur, Franz-Ludwig-von-Erthal Gymnasium, Lohr am Main Publication list Konopik, O., Gray, C., Grafe, T. U., Steffan-Dewenter, I., Fayle, T. M. (2014) From rainforest to oil palm plantations: shifts in predator population and prey communities, but resistant interactions. Global Ecology and Conservation 2: 385-394.

http://dx.doi.org/10.1016/j.gecco.2014.10.011 Schwarz, C. J., Konopik, O. (2014) An annotated checklist of the praying mantises (Mantodea) of Borneo, including the results of the 2008 scientific expedition to Lanjak Entimau Wildlife Sanctuary, Sarawak. Zootaxa 3797: 130-168.

http://dx.doi.org/10.11646/zootaxa.3797.1.12 Konopik, O., Linsenmair, K.-E., Grafe, T. U. (2014) Road construction enables establishment of a novel predator category to resident anuran community: a case study from a primary lowland Bornean rain forest. Journal of Tropical Ecology 30: 13-22.

http://dx.doi.org/10.1017/S0266467413000795 Grafe, T. U., Goutte, S., Ahmadsah, H. H., Konopik, O., Scharman, M. & Bauer, U.

(2010) Updated check list of the amphibians of the Ulu Temburong National Park and Brunei Darussalam. Scientia Bruneiana 11: 53-59.

Konopik, O., Steffan-Dewenter, I., Grafe, T. U. (under review) Effects of logging and oil palm expansion on stream frog communities on Borneo, Southeast Asia. Biotropica.

Konopik, O., Ahmad Sah, H. H., Keller, A., Wong, A., Grafe, T. U., Steffan-Dewenter, I.

(submitted to Ecology) Logging and conversion to oil palm plantations alters trophic interactions and reduces multiples levels of anuran diversity.

–  –  –

Summary (English) I. Nowadays, tropical landscapes experience large-scale land use intensification and land conversion driven by increasing demand for resourses. Due to the continuously high demand for tropical timber and politically intended step increase in palm oil production, multiple rounds of logging and subsequent conversion to oil palm plantations became a regionally wide-spread land conversion pattern in Southeast Asia. Although many tree species and some animals are highly threatened by logging, a great number of species groups, such as birds or mammals, have been shown to persist in logged forests. Accordingly, many ecosystem services, such as dung removal, seed dispersal or the activity of scavengers, are functionally maintained in logged forests. In contrast, oil palm plantations have been shown to not only dramatically alter the species composition and reduce biodiversity, but also curtail many crucial biotic and abiotic ecosystem functions. The focus of this dissertation was to investigate the response of anuran species richness and community composition to logging and conversion to oil palm plantation in northern Borneo (chapter II). I analysed the diet of various frog species and their change with habitat degradation. Furthermore, I assessed the shift in the trophic position of the anuran community as well as the response of anuran phylogenetic, dietary, and functional diversity to logging and conversion to oil palm plantations (chapter III). Finally, the resilience of the predator-prey interaction between an ant-specialist toad and its ant prey was analysed using shifts in species-level interactions (chapter IV).

II. This part of the study compares the species richness, relative abundance and community composition of stream anuran assemblages among primary forests, repeatedly logged forests and oil palm plantations. I used a highly standardised sampling setup applying transect-based sampling. Surprisingly, most of the anuran species native to primary forests were able to survive in logged forest streams. In contrast, on average only one third of the forest species richness was found in oil palm plantation streams. However, a high percentage of canopy cover above the plantation streams was able to mitigate this loss substantially. This study demonstrates the high conservation value of logged forests for Southeast Asian anurans. In contrast, the conversion to oil palm plantations leads to a dramatic decline of forest species.

However, they have a mainly unused potential to contribute to the protection of parts of the regional anuran biodiversity if conservation-oriented management options are implemented.

Summary (English)

III. In this part, I analysed the shifts in trophic position and multiple diversity layers of Southeast Asian stream-dependent anuran species across a gradient of disturbance from primary forest through intensively logged forest to oil palm plantation. For this purpose, I identified the diet composition of 59 anuran species by means of stomach flushing.

Furthermore, I use diet composition of frog species as well as species traits to calculate dietary and functional diversity, respectively. I found that the trophic position of the entire anuran community is elevated in heavily disturbed habitats. Furthermore, species diversity, phylogenetic species variation, dietary diversity, and functional diversity were reduced.

However, beyond the effect of the decreased species richness, only phylogenetic species variability and functional diversity were significantly impacted by land conversion, indicating a non-random loss of phylogenetic groups and functionally unique species. Overall, the observed changes to species interactions and functional composition suggest a greatly modified role of anurans in altered habitats and major foodweb reorganisation. Such farreaching changes to the way species groups interact are likely to threaten local biodiversity and ecosystem functioning in natural and particularly modified habitats. However, I could also show, that small-scale habitat quality, provided by riparian reserves, is able to mitigate the negative consequences of land conversion considerably.

IV. Here I assess how logging of rain forest and conversion to oil palm plantations affect the populations of the ant-specialist giant river toad (Phrynoidis juxtaspera), and availability and composition of its ant prey. I measured canopy cover as an estimate for the degree of disturbance. I found that toad abundance decreased with increasing disturbance. At the same time, ant community composition was altered, and local ground-foraging ant species richness increased with disturbance. However, for a given amount of canopy cover, primary forest supported more ant species than altered habitats. Despite these changes, composition of ants consumed by toads was only weakly affected by habitat change, with the exception of the invasive yellow crazy ant (Anoplolepis gracilipes), which was positively selected in oil palm plantations. This suggests that predator-prey interactions can be mostly maintained with habitat disturbance despite shifts in community composition, and even that some predators are capable of exploiting new prey sources in novel ecosystems.

V. I could show that anuran diversity and their trophic interaction is negatively impacted by logging and in particular by conversion to oil palm plantations. From species richness and community composition, my study expanded to phylogenetic, dietary and functional diversity.

Furthermore, I investigated the interaction of a particular toad species with its preferred prey

Summary (English)

(ants), on species level. This increasing degree of detail in my study provided comprehensive results, beyond the detail of many related studies. Overall, conservation of the remaining forest in Southeast Asia is urgently required to protect anuran biodiversity and their trophic interactions.

Zusammenfassung (German) Zusammenfassung (German) I. Durch den stetig steigenden Bedarf an vielfältigen Ressourcen stehen heutzutage vor allem tropische Ökosysteme unter enormem Druck hin zu intensiver Landwirtschaft und der Umwandlung von natürlichen Lebensräumen. Getrieben durch die hohe Nachfrage an Tropenhölzern und dem politisch gewollten Anstieg der Palmölproduktion, etablierte sich in Südostasien eine weit verbreitete Landnutzungsumwandlung charakterisiert durch wiederkehrende Holznutzung und die anschließende Umwandlung zu Ölpalmplantagen.

Manche Tiergruppen und vor allem Baumarten sind dadurch stark gefährdet. Dennoch können andere Organismengruppen wie Vögel oder Säuger auch in genutzten Wäldern überleben.

Auch Ökosystemdienstleistungen, wie Kotbeseitigung oder die Aktivität von Aasfressern bleiben vielfach in genutzten Wäldern erhalten. Im Gegensatz dazu stellen Ölpalmplantagen keinen geeigneten Lebensraum für viele Tier und Pflanzenarten dar und viele Ökosystemserviceleistungen können in diesen Plantagen nicht aufrecht erhalten werden. Ziel dieser Arbeit war die Auswirkungen dieser massiven Habitatumwandlung auf die Artenzahl und die Artenzusammensetzungen von Fröschen und Kröten in Nordborneo zu untersuchen (Kapitel II). Ich untersuchte die Nahrungszusammensetzung von mehreren Froscharten und wie sich diese mit Habitatumwandlung verändert. Zusätzlich habe ich die Stellung von Fröschen in der Nahrungskette sowie ihre phylogenetische, nahrungs- und funktionelle Diversität untersucht (Kapitel III). Schließlich habe die Interaktion von Fröschen mit ihrer Beute (in diesem Fall Ameisen) auch auf Artebene untersucht (Kapitel IV).

II. Dieser Teil der Arbeit vergleicht die Artenzahl, relative Häufigkeit und die Artenzusammensetzung der bachlebenden Froscharten zwischen Primärwäldern, genutzten Wäldern und Ölpalmplantagen. Dazu nahm ich die Froschbestände mit einer standardisierten Transektmethode auf. Ich fand heraus, dass die meisten Froscharten aus Primärwäldern auch in forstwirtschaftlich intensiv genutzten Wäldern überleben konnten. Im Gegensatz dazu wiesen Ölpalmplantagen nur durchschnittlich ein drittel der Froscharten aus Wäldern auf.

Jedoch konnte dieser Artenschwund durch einen hohen Kronenschluss über den Plantagenbächen reduziert werden. Diese Studie belegt den hohen naturschutzfachlichen Wert von genutzten Wäldern in Südostasien und die geringe Bedeutung von Ölpalmplantagen in diesem Zusammenhang. Jedoch zeigt es auch Wege auf, wie Ölpalmplantagen naturschutzfachlich aufgewertet werden können.

Zusammenfassung (German)

III. Dieser Teil der Arbeit befasst sich mit den Auswirkungen der Habitatumwandlung auf die trophischen Interaktionen und mehrere Ebenen der Froschbiodiversität. Dazu untersuchte ich die Nahrungszusammensetzung von 59 Froscharten mittels Magenspülung. Darüber hinaus nutzte ich diese Daten um die Nahrungsvielfalt zu untersuchen und Daten aus der Literatur und die funktionelle Diversität zu erfassen. Dabei fand ich heraus, dass Frösche bei zunehmender Habitatzerstörung eine höhere Stellung in der Nahrungskette einnahmen.

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