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Floral biology, flowering phenology and floral visitors of five insect pollinated tree species in a tropical lowland rainforest remnant of Pernambuco, Brazil [Elektronische Ressource] / vorgelegt von Leonhard Krause

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242 pages
Institut für Systematische Botanik und Ökologie Universität Ulm Floral Biology, Flowering Phenology and Floral Visitors of Five Insect-Pollinated Tree Species in a Tropical Lowland Rainforest Remnant of Pernambuco, Brazil Dissertation zur Erlangung des Doktorgrades Dr. rer. nat. der Fakultät für Naturwissenschaften der Universität Ulm vorgelegt von Leonhard Krause aus Luckenwalde Jahr der Promotion: 2008 Amtierender Dekan: Prof. Dr. Klaus-Dieter Spindler 1. Gutachter: Prof. Dr. Gerhard Gottsberger 2. Gutachter: Tag der Promotion: Table of contents I Table of contents TABLE OF CONTENTS.............................................................................................. I Acknowledgements.................................................................................................................................................I Index of scientific terms and abbreviations used in the text (including non-SI units) .....................................I 1 SUMMARY........................................................................................................... I 2 ZUSAMMENFASSUNG ....................................................................................... I 3 INTRODUCTION................................................................................................. 1 3.1 Tree reproduction in tropical lowland rainforests............
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Institut für Systematische Botanik und Ökologie
Universität Ulm






Floral Biology, Flowering Phenology and Floral Visitors
of Five Insect-Pollinated Tree Species in a Tropical
Lowland Rainforest Remnant of Pernambuco, Brazil



Dissertation
zur Erlangung des Doktorgrades Dr. rer. nat.
der Fakultät für Naturwissenschaften der Universität Ulm






vorgelegt von
Leonhard Krause aus
Luckenwalde

Jahr der Promotion: 2008
























Amtierender Dekan: Prof. Dr. Klaus-Dieter Spindler
1. Gutachter: Prof. Dr. Gerhard Gottsberger
2. Gutachter:

Tag der Promotion:

Table of contents I

Table of contents
TABLE OF CONTENTS.............................................................................................. I
Acknowledgements.................................................................................................................................................I
Index of scientific terms and abbreviations used in the text (including non-SI units) .....................................I
1 SUMMARY........................................................................................................... I
2 ZUSAMMENFASSUNG ....................................................................................... I
3 INTRODUCTION................................................................................................. 1
3.1 Tree reproduction in tropical lowland rainforests...............................................................................1
3.2 General species characterisation and state of the art .......................................................................... 1
3.2.1 Tapirira guianensis Aubl., Anacardiaceae........................................................................................... 1
3.2.2 Ocotea glomerata (Nees) Mez, Lauraceae 1
3.2.3 Trichilia lepidota Mart., Meliaceae ..................................................................................................... 1
3.2.4 Sclerolobium densiflorum Benth., Caesalpinioideae (Fabaceae) ......................................................... 1
3.2.5 Eschweilera ovata (Cambess.) Miers, Lecythidaceae 1
4 MATERIAL AND METHODS .............................................................................. 1
4.1 Study area................................................................................................................................................ 1
4.1.1 Forest vegetation at the Usina São José S/A sugar company............................................................... 1
4.1.2 Forest patch characterization................................................................................................................ 1
4.2 Methods ................................................................................................................................................... 1
4.2.1 Flower and inflorescence morphology................................................................................................. 1
4.2.2 Anthesis ............................................................................................................................................... 1
4.2.3 Sexual and reproductive system........................................................................................................... 1
4.2.4 Nectar................................................................................................................................................... 1
4.2.5 Floral scent........................................................................................................................................... 1
4.2.6 Flowering Phenology ........................................................................................................................... 1
4.2.7 Floral visitors ....................................................................................................................................... 1
5 RESULTS 1 Table of contents II

5.1 Floral biology, phenology and floral visitors of Tapirira guianensis .................................................. 1
5.1.1 Morphology.......................................................................................................................................... 1
5.1.2 Anthesis ............................................................................................................................................... 1
5.1.3 Sexual and reproductive system........................................................................................................... 1
5.1.4 Nectar................................................................................................................................................... 1
5.1.5 Floral scent........................................................................................................................................... 1
5.1.6 Phenology ............................................................................................................................................ 1
5.1.7 Floral visitors ....................................................................................................................................... 1
5.2 Floral biology, phenology and floral visitors of Ocotea glomerata...................................................... 1
5.2.1 Morphology.......................................................................................................................................... 1
5.2.2 Anthesis ............................................................................................................................................... 1
5.2.3 Sexual and reproductive system........................................................................................................... 1
5.2.4 Nectar................................................................................................................................................... 1
5.2.5 Floral scent........................................................................................................................................... 1
5.2.6 Phenology ............................................................................................................................................ 1
5.2.7 Floral visitors ....................................................................................................................................... 1
5.3 Floral biology, phenology and floral visitors of Trichilia lepidota ...................................................... 1
5.3.1 Morphology.......................................................................................................................................... 1
5.3.2 Anthesis ............................................................................................................................................... 1
5.3.3 Sexual and reproductive system........................................................................................................... 1
5.3.4 Nectar................................................................................................................................................... 1
5.3.5 Floral scent........................................................................................................................................... 1
5.3.6 Phenology ............................................................................................................................................ 1
5.3.7 Floral visitors ....................................................................................................................................... 1
5.4 Floral biology, phenology and floral visitors of Sclerolobium densiflorum ........................................ 1
5.4.1 Morphology.......................................................................................................................................... 1
5.4.2 Anthesis ............................................................................................................................................... 1
5.4.3 Sexual and reproductive system........................................................................................................... 1
5.4.4 Floral scent........................................................................................................................................... 1
5.4.5 Phenology ............................................................................................................................................ 1
5.4.6 Floral visitors ....................................................................................................................................... 1
5.5 Floral biology, phenology and floral visitors of Eschweilera ovata..................................................... 1
5.5.1 Morphology.......................................................................................................................................... 1
5.5.2 Anthesis ............................................................................................................................................... 1
5.5.3 Sexual and reproductive system........................................................................................................... 1
5.5.4 Nectar................................................................................................................................................... 1
5.5.5 Floral scent........................................................................................................................................... 1
5.5.6 Phenology ............................................................................................................................................ 1 Table of contents III

5.5.7 Floral visitors ....................................................................................................................................... 1
6 DISCUSSION ...................................................................................................... 1
6.1.1 Morphology.......................................................................................................................................... 1
6.1.2 Anthesis ............................................................................................................................................... 1
6.1.3 Sexual system, reproductive system, and pollen/ovule-ratio .............................................................. 1
6.1.4 Nectar................................................................................................................................................... 1
6.1.5 Floral scent........................................................................................................................................... 1
6.1.6 Flowering phenology ........................................................................................................................... 1
6.1.7 Floral visitors ....................................................................................................................................... 1
7 REFERENCES.................................................................................................... 1
8 APPENDIX .......................................................................................................... 1
Acknowledgements IV

Acknowledgements
• to Prof. Dr. Gerhard Gottsberger for the release and conception of the
interesting research theme within his project, project leadership, support,
teaching, incentive, countless helpful discussions during my field work and
writing, comments on earlier drafts, and preparing of the first expertise
• to Prof. Dr. Marian Kazda for integrating me in the Institute of Systematic
Botany and Ecology, providing working space, many helpful advice, and
preparing of the second expertise
• to my colleagues Dr. Daniel Piechowski, Dr. Michael Schessl, and Dr. Holger
Teichert for project elaboration and administration and for all the help and
support provided in the field, during discussions and in difficult moments
• to the counterparts Prof. Dra. Maria Jesus Nogueira Rodal and Prof. MSc. Ana
Carolina Borges Lins e Silva (UFRPE-Recife-Brazil) for their invitation,
friendly reception, collaboration, patience, scientific and logistical help, and
for the comprehension of the special needs of this work, muito obrigado!
• to the Usina São José S/A sugar company, especially to its director Frederico
Augusto Cavalcanti de Petribu Vilaça for kindly providing accommodation,
climatic data, research permission and infrastructure
• to CNPq and IBAMA for plant collecting and export permissions
• to Evelin Schäfer for the project administration and all the help in possible and
impossible situations
• to Dr. Ilse Silberbauer-Gottsberger, Dr. Dawn Frame, Prof. Dr. Manfred
Ayasse, Dr. Martin Freiberg, Dr. Albert-Dieter Stevens, Dr. Andreas Jürgens,
Dr. Christoph Knogge, Dr. Stefan Dötterl, Dr. Stefan Jarau, Dr. Heiko
Bellmann, Dr. Marco Tschapka, Kordula Heinen and Heiko Hentrich for their
valuable support, sharing opinions and for teaching their skills
• to the staff of the Institute of Systematic Botany and Ecology: Dr. Jürgen
Hoppe, Dr. Hermann Muhle, Dr. Martin Werth, Ellen Salzer, Eliana
Schacher,Hans Malchus,, and all its graduate and postgraduate students for
their help, joy and advice Acknowledgements V

• to Thomas, Ute, Marcus, and Sebastian for their support and company in
many situations, soon you will have your title too!
• to all the LEVE, LAFIT, and LECA students and researchers for collaboration,
help, joy, and support in countless occasions
• to Edgar Caliento Barbosa, Dr. Pille Urbas, and Marcelle Almeida da Silva for
assistance during field work and help in data gathering
• to the specialists for identifying insects: Prof. Dr. Clemens Schlindwein (bees),
Prof. Dr. Brett C. Ratcliffe (scarab beetles), M.Sc. Carlos E.B. Nobre (butterflies)
and M.Sc. Ana Gabriela Delgado Bieber (ants)
• to Dr. Stefan Dötterl for analysing all the scent samples
• to my friends Mauricéa and Tschá for their warm and cordial company every
time I went to Brazil
• to the environmentalists, colleagues, and friends Alexandre and Domingos
(R.I.P.), Roberto, Daniel, Lula, Paulo, Airton, Mariana, Úrsula, Bráulio, Eva,
Michele, Raymundo, Gabi, Paulinho, Janaína, Cibele, Cecília and Marcelo for
providing priceless, unique insights into the Brazilian way of nature
conservation and life
• to Fina and Tom for their interest in the work and the research movies of their
dad
• to Hanni and Mirko for their interest in Brazil, Pernambuco, and my work
• to my parents for their continuous support, comprehending my long absence,
and to Eneide
• especially to Patricia, for giving me all support and love I needed, having so
much patience with me, believing in me, and much more! Te amo!

This study was part of the project “Sustainability of remnants of Atlantic rainforest in
Pernambuco and its implications for conservation and regional development” within
the program “Mata Atlântica – Science and Technology for the Atlantic Rainforest”
financed by the German Ministry of Education and Research (BMBF) and the
Brazilian National Council for Scientific and Technological Development (CNPq). Index of used terms and abbreviations VI

Index of scientific terms and abbreviations used in the text
(including non-SI units)

AQC 6-aminoquinolyl-N-
Hydroxysuccinimidyl carbamate
a.s.l. above sea level
C nectar concentration (weight/weight)
d nectar density
dbh diameter in breast height
df degrees of freedom
n number individuals ind
ninflorescences infl
P/O pollen/ovule ratio
rpm rotations per minute
r Spearman rank correlation coefficient s
sd standard deviation
SEM scanning electron microscope
USJ Usina São José S/A sugar factory
VPD water vapour pressure deficit
arithmetic mean
median
1. Summary VII

1 Summary
The highly diverse Atlantic forest ranges from the northeastern tip of Brazil more
than 3500 km to the Brazilian-Paraguayan-Argentinean border, covering originally
15% of the country’s territory. 500 years after the beginning of the European
colonisation, only 7.6%, and in Northeast Brazil, less than 5% of the original forest
extension escaped in small forest patches from deforestation for, e.g., sugar cane
fields, pasture land, or human settlements. Key reproductive processes of tree species
like pollination or seed dispersal in those fragments are often negatively affected by
the loss of its pollinators and dispersers.
Prolonged deforestation in Northeast Brazil suggests that tree species which
today are frequent should be well-adapted to the adverse conditions resulting from
deforestation. However, the uncertain age of many forest fragments and lack of
studies on reproductive biology of these frequent tree species let plenty of doubt on
their invulnerability.
The present study aims to analyse the functionality of floral biology and
phenology of five frequent insect-pollinated tree species as well as diversity and
main behavioural patterns of its floral visitors, in order to detect potential bottlenecks
that may limit successful pollination and fruit set. The study was conducted in a 319
ha forest fragment on the property of a private sugar cane company of Northeast
Brazil.
Tapirira guianensis, Anacardiaceae is a frequent canopy tree species in Neotropical lowland
rainforests occurring in wide parts of Central and South America. In the study area,
its sex expression was clearly dioecious, with no gender switch detected. In the dry
season, flower buds were formed and entered into dormancy. Its length was
unpredictable and lasted up to 106 days in some individuals. 10.7 days after
triggering by rain fall events, male and female individuals entered synchronously
into a short explosive flowering peak, lasting up to three days in male and up to
seven days in female individuals. Both in single individuals and in the surveyed
population, multiple flowering events per flowering period were registered. The 1. Summary VIII

flowers were organized in loose axillary panicles, grouped together in clearly
distinguishable inflorescence clusters at the end of the branches. Male clusters were
much more conspicuous due to their 24.3-fold excess in terms of flower number
compared to female clusters, and due to their more exposed position at the branches.
In single male Tapirira guianensis flowers, functionality was observed up to 40
hours after opening, in single female flowers up to 6 days. The flowers were small
(diameter: 4.2 mm in male, 4.6 mm in female flowers), disc-shaped, greenish-white
and permitted easy, practically unobstructed access to the floral resources. In female
flowers, pollination occurred when insects searched for nectar or pollen. The
2stigmatic surface of female flowers was 0.28 mm .
Tapira guianensis had a pollen-ovule ratio typical for obligate outcrossing
species of 9,341:1 when assuming a 1:1 ratio between male and female flowers, or of
227,920:1 when considering the observed male excess.
Beside pollen in male flowers, main floral resource was nectar in both flower
sexes. In bagged flowers at the time of first insect visits at 05:50 h, the nominal
volume and concentration of the openly presented nectar was 2.27 µl and 23.1%
(w/w), respectively. It attained up to 72.7% and very low volumina due to water
evaporation in the course of the day. Per flowering event, a median nectar
production of 1.96 l per female and 5.09 l per male tree individual was estimated. The
nectar sugars were composed of 1% sucrose, 52% fructose and 48% glucose.
Therefore, the nectar was hexose-dominated which is known to difficult evaporation
compared to sucrose-dominated nectars. Four amino acids were detected at a total
-1 -1concentration of 1248 ng·µl , being proline the highest concentrated at 899 ng·µl .
The floral scent of Tapirira guianensis was emitted during daylight, had a dull,
heavy, not very sweet but still pleasant note and was largely composed of common
isoprenoids. 21 scent compounds were detected, 19 in that of male and 11 in that of
female flowers, with α-pinene, (E)-ocimene, and β-caryophyllene being the highest
concentrated with approx. 15% each. The relative amount of β-caryophyllene
decreased in the course of the day. 5 ng scent emission per flower and hour were
estimated.
Bagged female flowers formed no fruits, indicating the need for pollen vectors
and male individuals for fruit set.

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