Table Of ContentEdmundas Lekevičius
Biodiversity
Maintenance, Function, Origin, and
Self-Organisation into Life-Support
Systems
Biodiversity
Edmundas Lekevičius
Biodiversity
Maintenance, Function, Origin, and Self-
Organisation into Life-Support Systems
Edmundas Lekevičius
Nature Research Centre
Vilnius, Lithuania
ISBN 978-3-031-11581-3 ISBN 978-3-031-11582-0 (eBook)
https://doi.org/10.1007/978-3-031-11582-0
© The Editor(s) (if applicable) and The Author(s), under exclusive license to Springer Nature
Switzerland AG 2022
This work is subject to copyright. All rights are solely and exclusively licensed by the Publisher, whether
the whole or part of the material is concerned, specifically the rights of translation, reprinting, reuse of
illustrations, recitation, broadcasting, reproduction on microfilms or in any other physical way, and
transmission or information storage and retrieval, electronic adaptation, computer software, or by similar
or dissimilar methodology now known or hereafter developed.
The use of general descriptive names, registered names, trademarks, service marks, etc. in this publication
does not imply, even in the absence of a specific statement, that such names are exempt from the relevant
protective laws and regulations and therefore free for general use.
The publisher, the authors, and the editors are safe to assume that the advice and information in this book
are believed to be true and accurate at the date of publication. Neither the publisher nor the authors or the
editors give a warranty, expressed or implied, with respect to the material contained herein or for any
errors or omissions that may have been made. The publisher remains neutral with regard to jurisdictional
claims in published maps and institutional affiliations.
This Springer imprint is published by the registered company Springer Nature Switzerland AG
The registered company address is: Gewerbestrasse 11, 6330 Cham, Switzerland
In memory of Edward Osborne Wilson, one
of the greatest biologists of our time.
Foreword
Modern evolutionary biology has been tremendously effective at understanding and
dissecting the mechanisms at work in the evolution of species, but this progress has
largely come at the expense of explaining the broader patterns and processes of
diversification of life on Earth. Indeed, the so-called modern synthesis of evolution
focuses almost entirely on basic evolutionary mechanisms such as mutation, natural
selection, drift, and gene flow, while it has little to say about broader questions such
as: Why are there so many species on Earth? Why and how have ecosystems evolved
to their present structure, functional organisation, and level of complexity? What is
the functional role of genetic, phenotypic, and species diversity in the functioning
of ecosystems and the biosphere as a whole? Yet these questions are clearly of the
utmost importance in the current era of massive anthropogenic environmental
changes, which some have called the ‘Anthropocene’.
Based on this observation, Ed Wilson (1992), in his book The Diversity of Life,
predicated that ‘the greatest advances in evolutionary biology will be made in ecol-
ogy, explaining more fully in time why the diversity of life is of such and such a
nature and not some other’. In the present book, Edmundas Lekevicius takes up this
challenge and proposes a synthetic view of the ecological patterns and processes
that are involved in the evolution and diversification of life on Earth, from the small-
scale interplay of competition and cooperation between species to the large-scale
self-organisation of biodiversity and ecosystems through Earth’s history. Although
Lekevicius ironically refers to himself as a ‘sceptical maverick’ in modern evolu-
tionary biology, I believe his book asks the right questions and is fully in line with
Darwin, who did not hesitate to devote entire books to the emergence of sociality
and morality in humans or to the role of earthworms in the economy of nature.
Almost inevitably, on a subject as vast as the maintenance, function, origin, and
self-organisation of biodiversity, Lekevicius’ book raises more questions than it
answers. To take a few examples among many, his book argues convincingly that
some sort of trophic or functional division of labour is necessary for an ecosystem
to function, but it does not say roughly how many species or phenotypes would be
needed to do so. Similarly, it argues convincingly that a diversity of species or phe-
notypes is necessary to cope with spatially and temporally varying environments,
vii
viii Foreword
but it does not provide even a rough estimate of the number of species or phenotypes
needed for this purpose. Ecology simply does not yet have the theoretical and
empirical elements to answer such questions quantitatively. As a result of this lack
of knowledge, it is still impossible to properly test many of the predictions derived
from either theoretical deduction or empirical induction regarding the level of bio-
diversity on our planet, from the scale of a tiny microcosm to the scale of the entire
biosphere. The subtitle of Lekevicius’ book, ‘Towards a general theory of biodiver-
sity’, aptly recognises the fact that we do not yet have all the elements of a truly
general theory of biodiversity.
Lekevicius’ philosophical and methodological stance, however, is sometimes
problematic in this respect. For instance, he argues that coping with such a complex
phenomenon as biological diversity requires conceptual modelling, not mathemati-
cal modelling. As a theoretical ecologist who has developed theory on many of the
issues discussed in this book, I respectfully disagree with this view. On the contrary,
my experience has shown me that formalising concepts into mathematical models is
of paramount importance in clarifying concepts, turning them into testable predic-
tions, and then testing these predictions either experimentally or by using an appro-
priate combination of observational data and clever statistical analyses. In doing so,
scientific progress is greatly accelerated at relatively low cost (Loreau, 2010b).
Lekevicius is also highly critical of evolutionary biologists who have argued that
the history of life is not necessarily progressive and that evolution has a large ele-
ment of chance. In contrast to this view, he argues that life on Earth has a goal, to
increase the total biomass of the biosphere, and that evolution is constantly moving
life towards this goal. I have explained in detail elsewhere (Loreau, 2010a) why we
should not expect ecosystems and the biosphere to have a single goal because of the
tension between different levels of natural selection, and why we should carefully
distinguish between large-scale trends in the development of ecosystems or life in
general and true goals or optimum principles. The current disruptions brought about
by our own species, which are reducing the total biomass of the biosphere, clearly
illustrate that life does not always proceed gradually or progressively. More gener-
ally, I strongly believe that chance and necessity are only abstract concepts whose
relevance depends heavily on the spatial and temporal scale considered, and thus
that a complete theory of biodiversity requires an explicit consideration of the issue
of scale and an explicit integration of processes across scales.
Despite these reservations, I thoroughly enjoyed reading Lekevicius’ book. His
book is pleasantly provocative, full of stimulating ideas and established ecological
facts, and it shakes up neo-Darwinism in a refreshing way. It reminds us that some
of the biggest questions that evolutionary biology should be answering remain
unanswered, even though these questions are essential to overcome the current
global biodiversity crisis.
Centre for Biodiversity Theory Michel Loreau
and Modelling in Moulis (France)
Moulis, France
04.03.2022
Foreword ix
References
Loreau, M. (2010a). From populations to ecosystems: Theoretical foundations for a new ecologi-
cal synthesis (Monographs in population biology). Princeton University Press.
Loreau, M. (2010b). Linking biodiversity and ecosystems: Towards a unifying ecological theory.
Philosophical Transactions of the Royal Society B, 365, 49–60.
Wilson, E. O. (1992). The diversity of life. W. W. Norton.
Acknowledgments
This book would hardly have been completed without the support of my colleagues
from the Nature Research Centre of Lithuania. My foremost thanks go to Vincas
Būda, the director of this Centre, and Ričardas Paškauskas, the head of the
Laboratory of Algology and Microbial Ecology at this Centre, who both have not
only kept me going and would not allow me to sink into desperation but also man-
aged to organise funding for the preparation of this book. I also highly appreciate
the especially valuable mediation and advice provided by Gediminas Valkiūnas, the
head of the P. B. Šivickis Laboratory of Parasitology at the Centre. Finally, I would
like to express deep gratitude to my colleague Yuri Gleba from Nomad Bioscience
GmbH in Germany for his recommendations and comments.
I am equally grateful to my translator Mantas Zurba – I was impressed by his
dedication and diligence. He did more than just translate the book, as he helped me
to correct numerous inconsistencies in the original text.
Edmundas Lekevičius
xi
Contents
Part I In Search for a Functional Explanation
1 Introduction: Contemporary Biology Struggles
to Explain the Great Diversity of Genotypes and Species . . . . . . . . . . 3
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 9
2 Methodological Excursion: An Apology for Theoretical
Synthesis, Simplicity, Deductive Method,
and Conceptual Modelling . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 11
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 24
3 Species Do Not Just ‘Struggle’ . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 37
4 Species Diversity as Trophic Specialisation . . . . . . . . . . . . . . . . . . . . . 39
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 45
5 Producer Diversity as Reflection of Variation
in the Abiotic Environment . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 47
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 57
6 Do Consumers Maintain Diversity of Their Food Sources? . . . . . . . . 61
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
7 Species Diversity Ensures Higher Total Biomass
and Helps to Stabilise It . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 71
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
8 Conspecifics Do Not Just ‘Struggle’ Either . . . . . . . . . . . . . . . . . . . . . 79
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
9 Genotype Diversity as Even More Delicate Specialisation . . . . . . . . . 83
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 92
10 Environmental Fluctuations and the Daily Role of Selection . . . . . . . 95
References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 102
xiii
