Structural attributes contributing to locomotor performance in the ostrich (Struthio camelus) [Elektronische Ressource] / presented by Nina U. Schaller

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Structural attributes contributing to locomotor performance in the ostrichNina U. Schaller2008Dissertation submitted to th e Combined F aculties for th e N atural S ciences an d M athematicsof th e R uperto-Carola U niversity of H eidelberg, G ermanyfor th e d egree ofDoctor of N atural S ciences Structural at tributes contributing t o loc omotor p erformance in t he os trich ( Struthio c amelus)presented b yDipl. Bi ol. N ina U . S challerBirthplace: K ronberg/Ts (Germany)thOral examination: 16 O ctober 2008 Referees: Prof. D r.Dr. hc . V olker S torch P rof. D r. T homas Bra unbeckIndexA A cknowledgementsB SummaryC Zusammenfassung1 I ntroduction 12 S tate of present knowledge and basic principles 72.1 Classification and e volutionary bac kground72.2 General phy siology – Metabolic r efinements f or e conomic c ost of t ransport 82.3 Statics and dy namics – G eneral s trategies and s tructural s pecialisations f or 10 economic l ocomotion3 Materials and me thods 21 3.1 Integrative appr oach 213.1.1Functional and c omparative morphology 213.1.2 Biomechanics and l ocomotor dyna mics 223.1.3 Integration3.2 Ostrich r aising, k eeping and t raining 233.2.1 Raising 243.2.2 Keeping3.2.3 Training 243.



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Structural attributes
contributing to
locomotor performance
in the ostrich
Nina U. Schaller
submitted to th e
Combined F aculties for th e N atural S ciences an d M athematics
of th e R uperto-Carola U niversity of H eidelberg, G ermany
for th e d egree of
Doctor of N atural S ciences

Structural at tributes
contributing t o loc omotor p erformance
in t he os trich ( Struthio c amelus)
presented b y
Dipl. Bi ol. N ina U . S challer
Birthplace: K ronberg/Ts (Germany)
thOral examination: 16 O ctober 2008 Referees: Prof. D r.Dr. hc . V olker S torch

P rof. D r. T homas Bra unbeckIndex
A A cknowledgements
B Summary
C Zusammenfassung
1 I ntroduction 1
2 S tate of present knowledge and basic principles 7
2.1 Classification and e volutionary bac kground7
2.2 General phy siology – Metabolic r efinements f or e conomic c ost of t ransport 8
2.3 Statics and dy namics – G eneral s trategies and s tructural s pecialisations f or 10
economic l ocomotion
3 Materials and me thods 21

3.1 Integrative appr oach 21
3.1.1Functional and c omparative morphology 21
3.1.2 Biomechanics and l ocomotor dyna mics 22
3.1.3 Integration
3.2 Ostrich r aising, k eeping and t raining 23
3.2.1 Raising 24
3.2.2 Keeping
3.2.3 Training 24
3.3 Itemisation of m ethods 254 Results: Analysis of locomotor system an d locomotor dynamics 29
4.1 F unctional and c omparative m orphology 29
4.1.1 Morphology 29
4.1.2 Morphometry (published a s: Locomotor characteristics of the os trich 43
[Struthio c amelus] – M orphometric and m orphological analyses,
N. U. Schaller, B. Herkner & R. Prinzinger )
4.1.3 Statics 55

4.2 Biomechanics and l ocomotor dy namics 63
4.2.1 Cons training qua lities of the pa ssive locomotor s ystem 64
4.2.2 The intertarsal joint of the os trich: 71
Anatomical examination a nd func tion of pa ssive structures in locomotion
(N. U. Schaller, B. Herkner, R. Villa & P. A erts,
submitted to the Journal of Anatomy)

4.2.3 Dynamic analysis of pha langeal load a nd pre ssure distribution i n the os trich 95
(N. U. Schaller, K. D'Août, R. Villa B. Herkner & P. Aerts,
submitted to the Journal of Experimental Biology)
4.3 Contributions f rom a non-l ocomotor e lement 117
4.3.1 W ings 117
5 General discussion and integration 123
AppendixA Acknowledgements
Before mentioning all the people who advised on – and actively participated in – this study, I would like
to express that I feel very lucky to have been able to carry out this diversely-shaped project with its many
stimulating learning experiences. In light of this diversity, the sequence in which I offer thanks is not
intended t o r eflect an or der of importance – i n t he end, t he w hole of all the he lp m ade the w riting of these
following pa ges pos sible:
I deeply thank the CUSA NUSWE RK Foundation – Bischöfliche Studienförderung – in Bonn, Germany
for their generous PhD grant and the opportunity to meet many engaging personalities and gain new
friends. I would like to thank the DFG – German Research Foundation, Bonn – for a travel grant that
allowed a ttendance of a conference that pr oved ve ry i mportant for m y s ubsequent studies.
A t the University of Heidelberg, Germany, I am very happy to have Prof. Dr. Dr. hc. Volker Storch and
Prof. Dr. Thomas Braunbeck as reliable supervisors for my PhD project – especially considering that
they already had the 'pleasure' of supporting my aim to become a biologist more than a decade ago,
including f ield t rips w ith e ven-greater N ina-exposure than pos sible in t he sacred ha lls of our uni versity.
A t the Bayerische Staatssammlung in Munich, Germany, I was able to add valuable data to my research.
I particularly thank the technical assistant who helped me so much in carrying out morphometric
A t the ornithology department of the Natural History Museum, in Vienna, A ustria, I relied on great
specimen assistance from Dr. E rnst Bauernfeind, Dr. A nita Gamauf and Mr. Hans-Martin Berg but also
enjoyed my stays due to the very friendly atmosphere and – not surprising, considering the location – the
best coffee ever!
To Vienna I send another big thank you to Prof. Gerald Weissengruber at the Veterinärmedizinische
Universität who could not have been a better, nor more humorous, advisor and discussion partner in
specific S truthiomatic m atters.
Special thanks to the department of ornithology and palaeontology at the Royal Ontario Museum in
Toronto, Canada, for letting me explore their extensive collections and especially to Mr. Mark Peck who
arranged i mmediate access to t he collections w henever I ne eded a vian or theropod s keletal m aterial.
A t the Senckenberg Research Institute, Frankfurt/M., Germany – my major collaborating institution – I
thank Dr. Gerald Mayr and Prof. Dr. Dieter Stefan Peters from the ornithology department for their time
in discussion and Saskia Dimter for her commitment during the somewhat “animated” rhea dissection
sessions. I sincerely thank the paleoanthropology department for allowing periodic use of their facilities
and pa rticularly C hristine H emm for teaching m e – i n he r m otivating w ay – how to us e the 3D -scanner to
transfer solid ratite bones into a computer. I also would like to thank the staff, especially the
housekeepers and pr eparators, who w ere always w illing t o of fer assistance.
Hartelijk bedankt Prof. Dr. Peter A erts of the Laboratory for Functional Morphology at the Universiteit
A ntwerpen, Belgium, who took me under one of his many wings and provided guidance, suggestions and
deep insight at critical points in this project. I thank him also for his willingness to undertake a
spontaneous research trip to my ostriches where, besides packing a pressure plate, also brought Dr.
Kristiaan D'A oût – for me, the pre-eminent authority in footscan technology – who elucidated the
mysteries of all related conceptual and technical challenges. To this I can only say: “A wesome” job,
Kriske! Thank you both for many great discussions, telephone calls, reviews, emails and critiques. Your
ever-friendly hospitality in A ntwerpen introduced me to a beautiful city offering, in addition to
picturesque attractions, fantastisch l ekkere kr okettjes: S makelijk!
My deepest gratitude I send to the “Meister”, Dr. Bernd Herkner (Senckenberg Research Institute), my
“geistiger Ziehvater” and spiritual leader who guided me through this project with countless
philosophical observations, profound suggestions, revealing discussions and practical assistance. Besides
introducing me to the “holistic way of life”, he was (and remains!) a good friend and advisor in
sometimes stormy, but us ually f antastic, weather – H erzlichen D ank f ür alles!This project would not have been possible in the present form without the selfless participation and
support of ratite farming a nd ve terinary f acilities:
Thank you, Familie Gayer of the Rieser Straußenfarm, Hohenaltheim, Germany, who not only allowed
me to sample first-hand data but also provided me with my outstanding “research ostriches” and always
took t he time to a nswer m y que stions in de tail.
Thank you very much Mr. Schneider from the E mu farm Schneider, where I was provided the time and
space ga ther ki nematic and m orphometric da ta.
Great thanks I offer to Susanne and Horst E ngelhardt at the Donaumoos ostrich farm in Laipheim,
Germany, who allowed me to set up my data-acquisition system and actively helped me with some
“weighty” da ta collection. T heir hos pitality a nd pa rticipation w as gr eatly a ppreciated.
I also thank Mr. Kistner at the Mhou ostrich farm, Rülzheim, Germany, who provided me with a number
of os trich i ntertarsal joints, facilitating a n “ engaging” pa rt of m y s tudies.
'Doc' Bert Geier, Wiesbaden, Germany – Thank you so much for always being available for veterinary
cross-checks and providing answers to any and all questions concerning ostrich husbandry; it was a big
relief to be able to t rust in your expertise and e xtensive know ledge.
I owe a huge thank you und Danke to all my family and friends – old and new – who supported my work
both m entally a nd pr actically:
Special thanks to m y c omrade Dr. Heike S cherf for m any i nspiring di scussions and he r encouragement.
Thanks to m y de ar M eike K asten f or ke eping up m y s pirits w ith he r unpa ralleled s ense of hum or!
A ina and Mal“erator” Levin, my overseas family, for some editorial help and shelter offering relaxation
on a n a ppetizing a nd m usical scale.
Grazie tanto P appa V illa for a ni ce cottage retreat.
Olli, my very good friend, or, as he is officially referred to: Dr. Oliver Faulhaber, was always a source of
inspiration and encouragement and in his special, much-appreciated way, boosted my self-confidence
during rare episodes of “collapsed doctoral faith”. His practical assistance throughout the data sampling
sessions w ith m y os triches w ill remain unpa ralleled.
The goals of this project would have been unattainable without the on- going, selfless help from a large-
hearted man and his family. It started with the search for a suitable housing facility for my ostriches and
ended with finding myself yet another home, meiner Gass(t)familie: Jürgen, E del, Rosa-Li and Linda –
you all helped me so much in various ways throughout the last four years and the few lines I write here
simply cannot hope to reflect all you have done for me. I could always be sure that Tiffy and Frida were
happily kept in a safe and spacious haven. You lent practical assistance in the enclosure whenever
required and above all, you all welcomed me as a family member. It is fair to say that, with you around,
getting bored is hard but having a great time is really easy. “Prost” to many more years of shared
ostriches, Tintos and your gr eat pi zza!
One of the best lessons of this thesis was the confirmation that I will always be able to rely on my dear
parents, who I must also call my best friends. I sincerely thank Gisela and Rainer Schaller for their
absolute support, parental patience, friendly encouragement and vivid humour which turned some dark
hours pretty bright pretty fast! A bove and beyond the typical role of parents, they both actively
participated in ostrich transports and gave me a helping hand with my sometimes aesthetically-
challenged r esearch “ bits”. Speaking bot h c oncretely a nd a bstractly, I w ould not exist w ithout them!
I will never be able to thank enough my beloved partner, Rikk Villa, for his wide-ranging participation in
this thesis – but I will try to do so nevertheless! His engineering skills decisively facilitated data-
gathering e ven f rom across the bi g pond. I n num erous di scussions w e refined f undamental concepts and I
cannot imagine a more brilliant mental sparring partner. Finally, I happily admit that without his editorial
help and deep understanding of my way of thinking, I would not have been able to express my thoughts
in the present form. Thank you so very much and rest assured that in the next little while I will not bother
you w ith a nother P hD thesis!Last but certainly not least, I thank my feathered co-workers despite their obvious inability to read this –
I am simply in debt to them. Tiffy and Frida – with seemingly small brains – knew how to pull some
funny surprises: one should always beware of the ostrich beak! Without these friendly creatures I could
not have gained so much insight into their “locomotor performance”. Now that their career as research
ostriches ha s ended, t hey c an f ully c oncentrate on f ounding a family – G ood l uck f or your future!