Animal Evolution: Interrelationships of the Living Phyla

421 Pages·2012·9.88 MB·English

by  Claus Nielsen

Animal Evolution
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Animal Evolution
Interrelationships of the Living Phyla
C l a u s N i e l s e n
Natural History Museum of Denmark,
University of Copenhagen
t h i r d e d i t i o n
1
1
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P reface to third edition
S tudies of molecular phylogeny have dominated the (University of California, Berkeley), G. Edgecombe
biological literature during the last decade, but impor- (The Natural History Museum, London), D.E.K.
tant new information from morphology and embryol- Ferrier (University of St Andrews), D. Gordon
ogy has been obtained, especially with the use of new (NIWA, Wellington), S. Harzsch (University of
immunostaining methods. For a period, the molecular Greifswald), L.Z. Holland (Scripps Research
results of higher animal phylogeny looked rather cha- Institute), T.W. Holstein (University of Heidelberg),
otic, with many conf icting trees based on studies of G. Jékely (University of Tübingen), S. Karpov
mitochondrial and ribosomal genes. However, the (University of Southampton), N. King (University
trees obtained through the new studies of large num- of California, Berkeley), R.M. Kristensen (University
bers of various sequences, of expressed sequence tags, of Copenhagen), B.S.C. Leadbeater (University of
and even on whole genomes, now seem to converge, Southampton), S.P. Leys (University of Alberta), C.J.
and a congruence with morphology-based trees seems Lowe (Hopkins Marine Station), C. Lüter (Museum
possible. This has inspired me to make this new für Naturkunde, Berlin), J. Olesen (University of
edition. Copenhagen), G. Purschke (University of
R evisions of larger texts are always in danger of Osnabrück), H. Ruhberg (University of Hamburg),
growing, because new material is added and old mate- E. Schierenberg (University of Köln), A. Schmidt-
rial not discarded. I have tried to avoid this, for exam- Rhaesa (University of Hamburg), G. Shinn (Truman
ple by using the excellent series Microscopic Anatomy of State University), A.B. Smith (The Natural History
Invertebrates, edited by Frederick W. Harrison (Wiley- Museum, London), T. Stach (Freie University,
Liss, New York, 1991–1999) as a general reference for Berlin), B.J. Swalla (University of Washington), J.M.
the anatomy of the various groups; references that can Turbeville (Virginia Commonwealth University),
be found in these volumes have generally been R.M. Woollacott (Harvard University). A special
excluded. My two papers on the development of tro- thank you to my ‘neighbour’ in the Zoological
chophora larvae (Nielsen, C. 2004, 2005. J. Exp. Zool. Museum, Dr Martin Vinther Sørensen, for many
(M ol. Dev. Evol. ) 3 02B: 35–68 and 304 B: 401–447) good discussions. All these colleagues (and others
have been used in the same way. that I may have forgotten in the list) have been of
M any colleagues have helped in various ways; great help, but don’t blame them for my mistakes.
some have sent me illustrations for use in the book M rs Birgitte Rubæk is thanked for f ne help with
and others have read one or more chapters and given the new illustrations.
good suggestions: S. Amano (Kanazawa University),
C openhagen
M. Blaxter (University of Edinburgh), M.J. Dayel
M arch 2011
P reface to second edition
D uring the years since the text for the f rst edition of O nce again it is my pleasure to thank the many
this book was concluded (in 1992), a wealth of new generous colleagues who have helped me in various
morphological information has become available, ways, especially those who have read drafts of various
including both histological/ultrastructural and embry- chapters and given many good comments: André
ological data, and new areas, such as numerical cladis- Adoutte (Paris), Wim J.A.G. Dictus (Utrecht),
tic analyses, DNA sequencing, and developmental Andriaan Dorresteijn (Mainz), Danny-Eibye-Jacobsen
biology, have become prominent in phylogenetic stud- (Copenhagen), Peter W.H. Holland (Reading),
ies. I have tried to update the information about mor- Reinhardt Møbjerg Kristensen (Copenhagen),
phology, but the other f elds have been more diff cult Thurston C. Lacalli (Saskatoon), George O. Mackie
to deal with; numerical cladistic analyses and molecu- (Victoria), Mark Q. Martindale (Hawaii), Rudolf
lar phylogeny are discussed in separate chapters, but Meier (Copenhagen), Edward E. Ruppert (Clemson),
following my conclusions in these two chapters, I have George L. Shinn (Kirksville), Nikolaj Scharff
in general refrained from discussing results obtained (Copenhagen), Gerhard Scholtz (Berlin), Ralf
through these methods. Sommer (Tübingen), Martin Winther Sørensen
I am fully aware that my coverage of molecular (Copenhagen), Gregory A. Wray (Durham), Russel L.
studies, including the extremely promising evolution- Zimmer (Los Angeles). None of them should be held
ary developmental biology, is very incomplete. I have responsible for the ideas expressed here.
tried to select information from studies that appear to M y special thanks are due to Mr Gert Brovad
describe consistent phylogenetic signals, but my (Zoological Museum, Copenhagen) who has provided
choice is biased by my background as a morphologist. the new photos and to Mrs Birgitte Rubæk (Zoological
The interested reader is strongly advised to consult a Museum, Copenhagen) who has spared no effort in
recent textbook or review articles on the subject. preparing the many new drawings and diagrams.
P reface to first edition
N o naturalist can avoid being fascinated by the diver- T he ideas put forward in this book have developed
sity of the animal kingdom, and by the sometimes over a number of years, and during that period I have
quite bizarre specializations that have made it possible benef tted greatly from interactions with many col-
for the innumerable species to inhabit almost all con- leagues. Some have been good listeners when I have
ceivable ecological niches. felt the need to talk about my latest discovery; some
H owever, comparative anatomy, embryology, and have discussed new or alternative ideas, names or con-
especially molecular biology, demonstrate a striking cepts with me; some have provided eagerly sought
unity among organisms, and show that the sometimes pieces of literature or given me access to their unpub-
quite bewildering diversity is the result of variations lished results; and some have sent me photos for pub-
over a series of basic themes, some of which are even lication. To all these friends I extend my warmest
common to all living beings. thanks; no names are mentioned, because such a list
T o me, this unity of the animal kingdom is just as will inevitably be incomplete. A number of colleagues
fascinating as the diversity, and in this book I will try have read one to several chapters (the late Robert D.
to demonstrate the unity by tracing the evolution Barnes (Gettysburg) and Andrew Campbell (London)
of all of the 31 living phyla from their unicellular have read them all) and given very valuable and con-
ancestor. structive comments that I have often but not always
A ll modern books on systematic zoology empha- followed; I want to mention them all, not to make
size phylogeny, but space limitations usually preclude them in any way responsible, but to thank them for the
thorough discussions of the characteristics used to help and support that is necessary during an undertak-
construct the various phylogenetic trees. I will try to ing such as this: Quentin Bone (Plymouth), Kristian
document and discuss all the characters that have been Fauchald (Washington, DC), Gary Freemann
considered in constructing the phylogeny—both those (Austin), Jens T. Høeg (Copenhagen), Åse Jespersen
that corroborate my ideas and those that appear to (Copenhagen), Niels Peder Kristensen (Copenhagen),
detract from their probability. Margit Jensen (Copenhagen), Reinhardt Møbjerg
I n the study of many phyla, I have come across sev- Kristensen (Copenhagen), Barry S.C. Leadbeater
eral important areas in which the available informa- (Birmingham), Jørgen Lützen (Copenhagen), George
tion is incomplete or uncertain, and yet other areas O. Mackie (Victoria), Mary E. Petersen (Copenhagen),
that have not been studied at all; on the basis of this I Mary E. Rice (Fort Pierce), Edward E. Ruppert
have, for each phylum, given a list of some interesting (Clemson), Amelie H. Scheltema (Woods Hole),
subjects for future research, and I hope that these lists George L. Shinn (Kirksville), Volker Storch
will serve as incentives to further investigations. (Heidelberg), Ole S. Tendal (Copenhagen), and Russell
I t should be stressed that this book is not meant as L. Zimmer (Los Angeles).
an alternative to the several recent textbooks of sys- T he Danish Natural Science Research Council and
tematic zoology, but as a supplement, one that I hope the Carlsberg Foundation are thanked heartily for
will inspire not only discussions between colleagues their continued support covering travel expenses,
but also seminars on phylogeny—of the whole animal instrumentation and laboratory assistance; the
kingdom or of selected groups—as an integrated part Carlsberg Foundation has given a special grant to
of the teaching of systematic zoology. cover the expenses of the illustrations for this book.
viii Preface to First Edition
F inancial support from ‘Højesteretssagfører C.L. the typesetting of the book and for f ne work with the
Davids Legat for Slægt og Venner’ is gratefully layout and lettering of the illustrations.
acknowledged. D r Mary E. Petersen (Copenhagen) is thanked for
M rs Birgitte Rubæk and Mrs Beth Beyerholm are her meticulous reading of the f rst set of proofs.
thanked for their excellent collaboration on the artwork. F inally, my thanks go to Oxford University Press,
M y warmest thanks go to Kai and Hanne (Olsen & and especially Dr Cathy Kennedy, for a positive and
Olsen, Fredensborg) for a congenial undertaking of constructive collaboration.
C ontents
1 . I ntroduction 1 3 1. G NATHIFERA 180
2 . A NIMALIA (METAZOA) 7 3 2. P hylum Gnathostomulida 1 82
3 . P relude: Phylum Choanof agellata 1 3 3 3. P hylum Micrognathozoa 1 84
4 . E arly animal radiation 15 3 4. P hylum Rotifera 185
5 . P hylum Silicea 20 3 5. P OLYZOA (BRYOZOA s.l.) 1 95
6 . E URADICULATA 25 3 6. P hylum Entoprocta 199
7 . P hylum Calcarea 27 3 7. P hylum Cycliophora 204
8 . P ROEPITHELIOZOA 3 0 3 8. P hylum Bryozoa (Ectoprocta) 2 08
9 . P hylum Homoscleromorpha 3 1 3 9. B RACHIOZOA 220
1 0. E UMETAZOA (GASTRAEOZOA) 3 3 4 0. P hylum Phoronida 2 24
1 1. P hylum Placozoa 39 4 1. P hylum Brachiopoda 230
1 2. N EURALIA 42 4 2. E CDYSOZOA 238
1 3. P hylum Cnidaria 45 4 3. P ANARTHROPODA 2 40
1 4. T RIPLOBLASTICA 53 4 4. P hylum Arthropoda 242
1 5. P hylum Ctenophora 55 4 5. P hylum Onychophora 2 59
1 6. B ILATERIA 62 4 6. P hylum Tardigrada 265
1 7. A COELOMORPHA 64 4 7. C YCLONEURALIA 2 70
1 8. P hylum Acoela 66 4 8. N EMATOIDEA 275
1 9. P hylum Nemertodermatida 6 9 4 9. P hylum Nematoda 276
2 0. P hylum Xenoturbellida 7 1 5 0. P hylum Nematomorpha 2 86
2 1. E UBILATERIA 74 5 1. S CALIDOPHORA 290
2 2. P ROTOSTOMIA 87 5 2. P hylum Priapula 293
2 3. S PIRALIA (LOPHOTROCHOZOA) 1 04 5 3. P hylum Kinorhyncha 2 97
2 4. S CHIZOCOELIA 112 5 4. P hylum Loricifera 300
2 5. P hylum Annelida 114 5 5. P hylum Chaetognatha 3 04
2 6. P hylum Sipuncula 134 5 6. D EUTEROSTOMIA 3 11
2 7. P hylum Mollusca 140 5 7. A MBULACRARIA 316
2 8. P hylum Nemertini 156 5 8. P hylum Echinodermata 3 21
2 9. P hylum Platyhelminthes 1 66 5 9. H EMICHORDATA 3 35
3 0. P hylum Gastrotricha 174 6 0. P hylum Enteropneusta 3 36