Morphological Evolution, Adaptations, Homoplasies, Constraints, and Evolutionary Trends: Catfishes as a Case Study on General Phylogeny & Macroevolution

503 Pages·2004·34.34 MB·English

by  Rui Diogo

Morphological Evolution, Aptations,
Homoplasies, Constraints
and
Evolutionary Trends
Catfishes as a Case Study on General Phylogeny
and Macroevolution

Morphological Evolution, Aptations,
Homoplasies, Constraints
and
Evolutionary Trends
Catfishes as a Case Study on General Phylogeny
and Macroevolu tion
Rui Diogo
University of Liege
Liege, Belgium
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Library of Congress Cataloging-in-Publication Data
Diogo, Rui.
Morphological evolution, aptations, homoplasies, constraints and
evolutionary trends: catfishes as a case study on general phylogeny
and macroevolution/Rui Diogo.
p. cm.
Includes bibliographical references and index.
ISBN 1-57808-291-9
1. Catfishes--Phylogeny. 2. Macroevolution. I. Title
QL637.9.S5D56 2004
597l.49--dc22
ISBN 1-57808-291-9
O 2005, Copyright Reserved
All rights reserved. No part of this publication may be reproduced, stored in
a retrieval system, or transmitted in any form or by any means, electronic,
mechanical, photocopying or otherwise, without the prior permission.
Published by Science Publishers, Inc., NH, USA
Printed in India.
Preface
The catfishes, or Siluriformes, are included in the superorder Ostariophysi, a
major group of teleosts including, apart the Siluriformes, the
Gonorynchiformes, the Cypriniformes, the Characiformes and the
Gymnotiformes. With their 34 families, about 437 genera and more than 2700
species, catfishes represent about one third of all freshwater fishes and are
one of the most diverse and economically important groups of fresh and
brackish fishes in the world. They have a particularly wide and complex
geographical distribution, being found in North, Central and South America,
Africa, Eurasia, South-East Asia, Japan and Australia, with fossil catfishes
having even being reported in Antarctica.
The amazing diversity and complexity of order Siluriformes renders very
difficult the study of higher level phylogeny and thus general evolution of
this group as a whole. As highlighted by De Pinna (1998), in such a diverse
and complex group, historical processes such as phylogenetic convergences,
parallel adaptations to similar habitats or evolutionary reversions (e.g. in
subterranean catfishes) may expectedly be quite frequent. Such frequency of
alterations in historical processes raise serious problems in correctly inferring
the relationships among catfish groups from all continents, however, and
consequently from similar habitats within different regions of the globe. More-
over, such broad general studies are further complicated by another serious
drawback, related to the logistics of obtaining and including an appropriate
number of representatives of all major groups of catfishes in the phylogenetic
analysis. And not to be overlooked is the challenging and time-consuming
task of analysing and controlling, for each of these numerous terminal taxa,
the even more numerous characters necessarily obligatory to undertake such
a general phylogenetic analysis in a satisfactory manner.
The logistical difficulties probably explain why in spite of long-standing
scientific interest in the phylogeny of Siluriformes the only cladistic analyses
to date dealing with the interfamilial relationships of the order as a whole are
Mo's published work (1991) and De Pinna's unpublished thesis (1993).
vi Morphological Evolution, Aptations, Homoplasies, Constraints and Evolutionary Trends
Logistical difficulties might well account for neglect of muscular characters in
traditional studies of catfish interrelationships as these are often not readily
detectable nor readily documented and, as just mentioned, analysing and
controlling a large number of such features in each taxon constitutes a rather
difficult task. Moreover, museums are much more reluctant to loan the
specimens necessary for complete muscular dissection than specimens for
osteological observations.
Given all the obstacles mentioned above and the associated fact that the
only general cladistic studies on Sjluriformes are those of Mo (1991) and De
Pinna (1993), it is not surprising that the vast majority of catfish researchers
continue to view siluriform higher level phylogeny as a largely unresolved
issue (see Alves-Gomes, 2001; Chardon et al., 2003; Gayet and Meunier, 2003;
Ng, 2003; Teugels, 2003). One of the major purpose of this book is tc, analyse
the puzzling, intriguing, higher-level interrelationships of catfishes, and to
discuss the general evolution of these fishes. To achieve this purpose meant
devoting a large part of the work to a cladistic analysis of the higher level
phylogeny of the group. Some terminal taxa not included in previous analy-
ses, and particularly a large number of characters traditionally excluded from
those analyses, such as those concerning catfish myology, have been incorpo-
rated in this analysis, which lends particular importance to complex struc-
tures. The cladistic analysis of catfish general phylogeny and the focus on
their complex structures thus pave the way for a discussion on the evolution
of these complex structures within the whole order Siluriformes and on cat-
fish general evolution. Lastly, the focus on catfish higher level phylogeny and
evolution allows a broader, theoretical discussion on general phylogeny and
macroevolution.
Liege, Belgium RUI DIOGO
February 2nd, 2004
Acknowledgement
This project greatly benefited from the precious help of P. Vandewalle and
M. Chardon, to whom I owe a deep debt of gratitude. Since they accepted me
in the Laboratory of Functional and Evolutionary Morphology, in 1998, they
have introduced me to the anatomy, functional morphology, phylogeny and
systematics of Vertebrates in general and of Teleostei in particular. They have
shown me how interesting are the Siluriformes, not only in what concerns the
study of many different issues concerning the order itself, but also in what
refers to the ample and diverse implications that the analysis of these issues
has for a general discussion on theoretical Biology. They have not only spared
their time continuously to discuss various points concerning the project and
biological sciences in general, but also introduced me to Belgium and common
life in this beautiful country. I also want to thank very, very much my
laboratory's colleague, E. Parmentier. He is really one of the brightest young
scientists I have ever met, and his persistence, the remarkably ability that he
has to solve all the different type of problems, and the courage he has to enter
and to get deep involved in all type of scientific areas were really inspiring
for me.
A very special thanks to the late G. G. Teugels (Muske Royal de 1'Afrique
Centrale), for kindly providing several specimens studied in this work, for
participating in so many projects and for discussing catfish phylogeny and
systematics with me, and for allowing me to undertake bibliographical re-
search on his Museum, which revealed to be fundamental for this work. I am
also particularly thankful to R. Vari, as well as their colleagues S. Weitzman,
J. Williams and S. Jewett (National Museum of Natural History), for accept-
ing me in his amazing Museum during two academic years, for providing a
large part of the specimens analyzed in this work, and for reading and com-
menting several papers included in this work. I am thankful to I. Doadrio
(Museu Nacional de Ciencias Naturales) and F. Poyato-Ariza (Universidade
Autonoma de Madrid) for also receiving me in their laboratories and for
valuable scientific discussions.
viii Morphological Evolution, Aptations, Homoplasies, Constraints and Evolutio?~ya Trends
I acknowledge J. Cambray (Albany Museum of Grahamstown), P. Laleyk
(Universitk Nationale du Bknin), P. Duhamel (Muskum National D'Histoire
Naturelle), W. L. Fink, D. W. Nelson and H. H. Ng (Museum of Zoology,
University of Michigan), and M. Stiassny (American Museum of Natural
History) for providing numerous specimens studied in this work. Thanks to
R. Bills and P. Skelton (South African Institute for Aquatic Biodiversity) for
the kind donation of three austroglanidid specimens to the Laboratory of
Functional and Evolutionary Morphology, as well as to B. Hall, F. Galis,
T. Grande, T. Abreu, C. Ferraris, J. Lundberg, M. Brito, M.M. de Pinna,
P. Skelton, R. Reis, L. Soares-Porto, P. Bockmann, A. Zanata, E. Trajano,
B.G. Kapoor, F. Meunier, C. Oliveira, P. Peng, M. Hardman, S. He, D. Adriaens,
F. Wagemans, H. Gebhardt, M. Ebach, A. Wyss, J. Waters, B. Perez-Moreno,
G. Cuny, A. Choudhury, M. Vences, S.H. Weitzman, L. Cavin, F. Santini,
J.C. Briggs, L.M. Gahagan, M. Philiphe, J.G. Maisey, J. Alves-Gomes, T. Roberts,
M.J.L. Stiassny, R. Winterbottom and M. Gayet, P. Lecointre and L. Taverne
for their helpful criticism, advice and assistance.
I would like to thank very, very much G. Arratia. Being closely in contact
with her work while editing the book "Catfishes", seeing the seriousness and
the remarkable quality of her research, and hearing from her many and many
comments and, sometimes, I must say, hard critiques, has been extremely
important for me and for my personal formation as a scientist. Thank you so
much.
My special thanks to all my friends, particularly to Pedro Brito, Gregory
Piskula, Joao Malcato, Pedro Castro, Henry Evrard, Denoel Mathieu, Neo
Toumbos, Thomas Stokart and particularly Claudia Oliveira. Thank you very
much to you, Alejandrita Pelito Lindo. A very, very, very special thanks to
my parents, Valter and Fatima, to my brothers, Hugo and Luis, and to my
grandfather, Raul.
I really would also like to thank R. Primlani, who kindly invited me to
publish this work as a book of the prestigious company Science Publishers
Co. Thank you very much for the confidence in my work and for the several
projects we have together. I hope J will not disappoint you.
A particularly thorough review of the whole book was done by Margaret
Majithia, from Oxford & IBH Publishing Co., for which I am especially grateful.
Finally, thanks to all those who were involved in administering my PhD
scholarship (PRAXIS XXI/BD/19533/99, FundacZo para a Ciencia e a
Tecnologia, Portuguese Government) and other grants and/or awards received
during the last years, without whom this work would really not had been
possible.
Thanks to ALL of you!!!
Contents
Preface
Acknowledgement vii
1. Catfishes: Introduction
1. I Phylogenetic Position within Teleostei
1.2 Catfish Families
1.3 Historical Overview of Higher Level Phylogeny of Catfishes
1.4 Catfish, an Exceptional Biological Group
2. Methodology and Material
2.1 Phylogenetic Methodology
2.2 Delimitation of Terminal Taxa
2.3 Material, Techniques and Nomenclature
3. Phylogenetic Analysis
3.1 Character Description and Comparison
3.2 Cladistic Analysis, Diagnosis for Clades, and
Comparison with Previous Hypotheses
3.3 Character State Changes for Individual Genera
3.4 Results of Phylogenetic Analysis: Major Outlines
4. Higher-level Phylogeny and Macroevolution of Catfishes:
A Discussion
4.1 Structures Associated with Movements of the Mandibular
Barbels
4.2 Pectoral Girdle Complex
4.3 Adductor Mandibulae Complex
4.4 Palatine-maxillary System
4.5 Suspensorium and Associated Structures
4.6 Elastic Spring Apparatus
4.7 A Discussion on the Origin and Biogeographic Distribution
of Catfishes