Table Of ContentInvertebrate Ecology & Conservation Monographs E Ecological Traits and
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ISSN 1312-9082 lo
Senior editor: Josef Settele gic Genetic Variation in
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The main aim of the series is to attract the attention of both aca- r
a
demics and society to model studies in invertebrate ecology and it Amazonian Populations of the
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conservation. It is also intended to include in-depth studies sup- a
n
ported by detailed data, which nowadays are rarely published as d G Neotropical Millipede
complete works because of their volume, but which are often e
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much more sustainable than many high impact journal papers. e
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ic Poratia obliterata (Kraus, 1960)
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Volume 2 a
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ia
Ecological Traits and Genetic Variation in Amazonian Populations tio (Diplopoda: Pyrgodesmidae)
of the Neotropical Millipede Poratia obliterata (Kraus, 1960) n in
(Diplopoda: Pyrgodesmidae) (Brazil) A
by Natalie Bergholz m (Brazil)
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The periodic flood pulse of the Amazon River has been the main ia
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controlling factor in the riverine ecosystem for at least two P
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million years. Numerous adaptations, in some cases along with spe- p
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ciation, have evolved in the resident terrestrial invertebrates. la
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Having probably originated from the Andes, the small millipede io
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Poratia obliterata(Kraus, 1960) is currently known as bisexual s o
from all over Amazonia and as parthenogenetic from some f t
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European hothouses. Seasonal vertical migration to tree trunks, e
aggregation and a univoltine life cycle appear to be adaptive traits N
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of individuals living in Central Amazonian inundation forests. Such t
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developed survival strategies can either reflect ecological plasticity op
or implicate ecological speciation, i.e. ‘biotope-specific races’ or ica
ecotypes. To assess the causal mode of adaptation, ecological studies l M
were combined with genetic analyses. The allozyme data did not illip
support biotope-specific genetic adaptations of P. obliteratain e
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Central Amazonia. Comparing the distribution, biotope range, e
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population subdivision and genetic diversity of different millipedes, o
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the species P. obliterataappears to feature a generalist strategy. atia
The results show low divergence between Amazonian populations o
owfi tthh ivsa driiovuerss be ioantodp wesid aensdp rtehauds sspuecccieesss, fwuhlliyc ihn vsaedemeds steoascoonpael wineulln- bliter
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dation forests. The book is addressed to specialists in evolutionary ta
Natalie G. R. Bergholz
biology, ecological genetics, ecology and conservation of wetlands,
and millipede research and conservation.
The IECM series is established with the support of the
EU FP6 project ALARM (www.alarmproject.net)
For submitting of manuscripts,
ordering and subscriptions:
Pensoft Publishers, [email protected]
www.pensoft.net/iecm/index.html
Invertebrate Ecology and Conservation Monographs
Senior editor
Josef Settele
–— Volume 2 –—
Ecological Traits and Genetic Variation in
Amazonian Populations of the
Neotropical Millipede Poratia obliterata (K , 1960)
RAUS
(Diplopoda: Pyrgodesmidae) (Brazil)
by
Natalie G. R. Bergholz
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2
Ecological Traits and Genetic
Variation in Amazonian Populations
of the Neotropical Millipede
Poratia obliterata (K , 1960)
RAUS
(Diplopoda: Pyrgodesmidae) (Brazil)
N G. R. B
ATALIE ERGHOLZ
SOFIA–MOSCOW
2007
INVERTEBRATE ECOLOGY AND CONSERVATION MONOGRAPHS
Senior editor: Josef Settele
Editorial board: Joachim Adis (Germany, [email protected]), Sergei Golovatch (Russia, [email protected]),
Kongluen Heong (Korea, [email protected]), Marc Kenis (Switzerland, [email protected]), Timothy New (Australia,
[email protected]), Jari Niemelä (Finland, [email protected]), Lyubomir Penev (Bulgaria, [email protected]),
Alain Roques (France, [email protected]), Michael Samways (South Africa, [email protected]), John Spence
(Canada, [email protected]), Ingolf Steffan-Dewenter (Germany, [email protected]), Bernard Vaissiere
(France, [email protected]), Allan Watt (UK, [email protected]), Michal Woychiekowski (Poland, [email protected]),
Steve Wratten (New Zealand, [email protected])
The present volume is published with a partial financial support of the
Max-Planck-Institute for Limnology, Plön, Germany (www.mpil-ploen.mpg.de)
Volume 2
ECOLOGICAL TRAITS AND GENETIC VARIATION IN AMAZONIAN
POPULATIONS OF THE NEOTROPICAL MILLIPEDE PORATIA OBLITERATA (KRAUS, 1960)
(DIPLOPODA: PYRGODESMIDAE) (BRAZIL)
NATALIE G. R. BERGHOLZ
E-mail: [email protected]
Scientific readers: Prof. Dr. Joachim Adis, Dr. Sergei I. Golovatch,
Prof. Dr. Jost Kömpf & Prof. Dr. Jürgen Tomiuk
First published 2007
ISBN 978-954-642-290-3
Invertebrate Ecology and Conservation Monographs, Volume 2
ISSN 1312-9082
© PENSOFT Publishers
All rights reserved. No part of this publication may be reproduced, stored in a retrieval system or
transmitted in any form by any means, electronic, mechanical, photocopying, recording or
otherwise, without the prior written permission of the copyright owner.
Pensoft Publishers
Geo Milev Str. 13a, Sofia 1111, Bulgaria
[email protected]
www.pensoft.net
Printed in Bulgaria, January 2007
4
E
DITORIAL
Biodiversity and its conservation have become key issues for science and society. Enor-
mous efforts have been devoted in recent decades worldwide to study the functional role
of biodiversity in natural and human-transformed ecosystems, patterns in biodiversity
and processes determining its variation and induced changes over space and time, and
ways of conserving biodiversity. The integration of science, both at European and world
level has led to numerous remarkable studies on biodiversity, often performed by large
international teams of specialists.
The usual practice of publishing in science is somewhat biased to offering the scientific
results to international journals. This practice is understandable but such publications are
usually limited in volume and their lifespan is often limited to a few years. This phenome-
non forces researchers to publish their results in several separate papers, often spread in
different journals. The integration of science, however, demands also an integrative approach
in publishing and disseminating the scientific results to the academic community, stake-
holders and society. To respond to this obvious need, Pensoft Publishers in cooperation
with an international team of specialists from around the world, is pleased to launch a new
series entitled “Invertebrate Ecology and Conservation Monographs”. The series will be
published in 4 issues per year. Each issue may be published in a volume of a maximum of
250 printed pages. Larger monographs will be published as supplements to the series.
The main aim of the series is to attract the attention of both academics and society to
model studies in invertebrate ecology and conservation. It is also intended to include in-
depth studies supported by detailed data, which nowadays are rarely published as com-
plete works because of their volume, but which are often much more sustainable than
many high impact journal papers.
The first two volumes appear in January 2007 and are devoted to two well-designed popula-
tion biology studies on a European butterfly and an Amazonian millipede. The first vol-
umes are: Ecology of an Almost Extinct Population of the Scarce Swallowtail (Iphi-
clides podalirius (Linnaeus, 1758)) by Roland Steiner, Gabriel Hermann and Josef Settele
(volume 1; in German with English summary) and Ecological Traits and Genetic Varia-
tion in Amazonian Populations of the Neotropical Millipede Poratia obliterata (Kraus,
1960) (Diplopoda: Pyrgodesmidae) (Brazil) by Natalie Bergholz (volume 2).
The Editorial Board hopes that the new series will quickly become an authoritative, well-
recognised forum for all those who want to publish their results as a whole in a distinct,
monograph-style, book. For the submission of manuscripts, either the Publisher, the Se-
nior Editor or any of the members of the Editorial Board can be contacted.
The Editors
5
EDITORIAL
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2
This work is dedicated to my family,
as well as to my best friends,
Kathleen Katscher and Swetlana Siniza
Whilst standing in the midst of the grandeur of a Brazilian forest,
it is not possible to give an adequate idea of the higher feelings of wonder,
admiration, and devotion which fill and elevate the mind.
(Charles R. Darwin, 1832)
It is not the strongest of the species that survive, nor the most intelligent,
but the one most responsive to change.
(Attributed to Charles R. Darwin)
8
C
ONTENTS
Editorial — 5
1 Introduction — 13
1.1 Study Area — 13
1.1.1 Geology of the Amazon Basin — 13
1.1.2 Climate — 15
1.1.3 Water Level Regime — 16
1.1.4 Seasonal Inundation Forests — 16
1.2 The Millipede Poratia obliterata — 17
1.2.1 Classification and Biology — 17
1.2.1.1 Systematics — 17
1.2.1.2 Basic Anatomy — 18
1.2.1.3 Characterisation — 19
1.2.1.4 Development — 20
1.2.1.5 Ecology — 21
1.2.1.6 Dispersal Ability — 23
1.2.2 Origin and Range of P. obliterata Populations — 23
1.2.3 Biotope-Related Adaptation and Speciation? — 24
1.3 Survival Strategies in Central Amazonian Floodplains — 26
1.4 Molecular Markers in Population Genetics — 29
1.4.1 Definition of Molecular Markers — 29
1.4.2 Subject of Population Genetics — 29
1.4.3 Application of Molecular Markers — 29
1.4.3.1 Classification and Speciation — 29
1.4.3.2 Adaptive Divergence among Populations — 31
1.4.4 Technique — 31
1.4.5 Appraisal: Choice of an Appropriate Marker — 32
2 Thesis Outline — 33
2.1 Part I: Ecological Traits — 33
2.2 Part II: Genetic Variation — 33
3 Ecological Traits — 35
3.1 Material and Methods — 35
3.1.1 Study Sites — 35
3.1.1.1 Várzea — 35
3.1.1.2 Igapó — 38
3.1.1.3 Várzea & Igapó — 41
3.1.1.4 Terra firme — 44
9
CONTENTS
