Evolutionary Developmental Biology of Invertebrates 2: Lophotrochozoa (Spiralia)

291 Pages·2015·19.672 MB·English

by  Andreas Wanninger (eds.)

Andreas Wanninger Editor
Evolutionary
Developmental
Biology of
Invertebrates
Vol. 2 Lophotrochozoa (Spiralia)
Evolutionary Developmental Biology
of Invertebrates 2

Andreas Wanninger
Editor
Evolutionary
Developmental Biology
of Invertebrates 2
Lophotrochozoa (Spiralia)
Editor
Andreas Wanninger
Department of Integrative Zoology
University of Vienna
Faculty of Life Sciences
Wien
Austria
ISBN 978-3-7091-1870-2 ISBN 978-3-7091-1871-9 (eBook)
DOI 10.1007/978-3-7091-1871-9
Library of Congress Control Number: 2015947925
Springer Wien Heidelberg New York Dordrecht London
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Cover illustration: Scanning electron micrograph of an early trochophore larva of the tusk shell,
Antalis entalis, a scaphopod mollusk. See Chapter 7 for details
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Pref ace
T he evolution of life on Earth has fascinated mankind for many centuries.
Accordingly, research into reconstructing the mechanisms that have led to the
vast morphological diversity of extant and fossil organisms and their evolu-
tion from a common ancestor has a long and vivid history. Thereby, the era
spanning the nineteenth and early twentieth century marked a particularly
groundbreaking period for evolutionary biology, when leading naturalists and
embryologists of the time such as Karl Ernst von Baer (1792–1876), Charles
Darwin (1809–1882), Ernst Haeckel (1834–1919), and Berthold Hatschek
(1854–1941) realized that comparing ontogenetic processes between species
offers a unique window into their evolutionary history. This revelation lay the
foundation for a research fi eld today commonly known as Evolutionary
Developmental Biology, or, briefl y, EvoDevo.
While for many of today’s EvoDevo scientists the principle motivation for
studying animal development is still in reconstructing evolutionary scenarios,
the analytical means of data generation have radically changed over the cen-
turies. The past two decades in particular have seen dramatic innovations
with the routine establishment of powerful research techniques using micro-
morphological and molecular tools, thus enabling investigation of animal
development on a broad, comparative level. At the same time, methods were
developed to specifi cally assess gene function using reverse genetics, and at
least some of these techniques are likely to be established for a growing num-
ber of so-called emerging model systems in the not too distant future. With
this pool of diverse methods at hand, the amount of comparative data on
invertebrate development has skyrocketed in the past years, making it increas-
ingly diffi cult for the individual scientist to keep track of what is known and
what remains unknown for the various animal groups, thereby also impeding
teaching of state-of-the-art Evolutionary Developmental Biology. Thus, it
appears that the time is right to summarize our knowledge on invertebrate
development, both from the classical literature and from ongoing scientifi c
work, in a treatise devoted to EvoDevo.
Evolutionary Developmental Biology of Invertebrates aims at providing
an overview as broad as possible. The authors, all renowned experts in the
fi eld, have put particular effort into presenting the current state of knowl-
edge as comprehensively as possible, carefully weighing conciseness
against level of detail. For issues not covered in depth here, the reader may
consult additional textbooks, review articles, or web-based resources,
v
vi Preface
p articularly on w ell-e stablished model systems such as C aenorhabditis
elegans ( w ww.wormbase.org ) or D rosophila melanogaster ( w ww.fl ybase.org ) .
Evolutionary Developmental Biology of Invertebrates is designed such
that each chapter can stand alone, and most chapters are dedicated to one
phylum or phylum-like taxonomic unit. The main exceptions are the hexa-
pods and the crustaceans. Due to the vast amount of data available, these
groups are treated in their own volume each (Volume 4 and Volume 5, respec-
tively), which differ in their conceptual setups from the other four volumes.
In addition to the taxon-based parts, chapters on embryos in the fossil record,
homology in the age of genomics, and the relevance of EvoDevo for recon-
structing evolutionary and phylogenetic scenarios are included in Volume
1 in order to provide the reader with broader perspectives of modern- day
EvoDevo. A chapter showcasing developmental mechanisms during regen-
eration is part of Volume 2 .
Evolutionary Developmental Biology of Invertebrates aims at scientists
that are interested in a broad comparative view of what is known in the fi eld
but is also directed toward the advanced student with a particular interest in
EvoDevo research. While it may not come in classical textbook style, it is my
hope that this work, or parts of it, fi nds its way into the classrooms where
Evolutionary Developmental Biology is taught today. Bullet points at the end
of each chapter highlight open scientifi c questions and may help to inspire
future research into various areas of Comparative Evolutionary Developmental
Biology .
I am deeply grateful to all the contributing authors that made E volutionary
Developmental Biology of Invertebrates possible by sharing their knowledge
on animal ontogeny and its underlying mechanisms. I warmly thank Marion
Hüffel for invaluable editorial assistance from the earliest stages of this proj-
ect until its publication and Brigitte Baldrian for the chapter vignette artwork.
The publisher, Springer, is thanked for allowing a maximum of freedom dur-
ing planning and implementation of this project and the University of Vienna
for providing me with a scientifi c home to pursue my work on small, little-
known creatures.
This volume covers the animals that have a ciliated larva in their life cycle
(often united as Lophotrochozoa), as well as the Gnathifera and the
Gastrotricha. The interrelationships of these taxa are poorly resolved and a
broadly accepted, clade-defi ning autapomorphy is lacking. Spiral cleavage is
sometimes assumed as the ancestral mode of cleavage of this grouping, and
therefore the clade is named Spiralia by some authors, although others prefer
to extend the term Lophotrochozoa to this entire assemblage. Aside from the
taxon-based chapters, this volume contains a chapter that highlights similari-
ties and differences in the processes that underlie regeneration and ontogeny,
using the Platyhelminthes as a case study.
Tulbingerkogel, Austria Andreas Wanninger
January 2015
Contents
1 Gnathifera. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 1
Andreas Hejnol
2 Gastrotricha . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 13
Andreas Hejnol
3 Platyhelminthes . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 21
Teresa Adell, José M. Martín-Durán,
Emili Saló, and Francesc Cebrià
4 Regeneration and Growth as Modes of Adult
Development: The Platyhelminthes as a Case Study . . . . . . . . 41
Francesc Cebrià, Emili Saló, and Teresa Adell
5 Cycliophora . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 79
Andreas Wanninger and Ricardo Neves
6 Entoprocta . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 89
Andreas Wanninger
7 Mollusca . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 103
Andreas Wanninger and Tim Wollesen
8 Nemertea. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 155
Jörn von Döhren
9 Annelida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 193
Christoph Bleidorn, Conrad Helm, Anne Weigert,
and Maria Teresa Aguado
10 Phoronida . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 231
Scott Santagata
11 Ectoprocta. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 247
Scott Santagata
12 Brachiopoda . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 263
Scott Santagata
Index . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 279
vii
1
Gnathifera
Andreas Hejnol

Chapter vignette artwork by Brigitte Baldrian.
© Brigitte Baldrian and Andreas Wanninger.
A. Hejnol
Sars International Centre for Marine Molecular
Biology , University of Bergen ,
Thormøhlensgate 55 , Bergen 5008 , Norway
e-mail: [email protected]
A. Wanninger (ed.), Evolutionary Developmental Biology of Invertebrates 2: Lophotrochozoa (Spiralia) 1
DOI 10.1007/978-3-7091-1871-9_1, © Springer-Verlag Wien 2015
2 A. Hejnol
INTRODUCTION internal relationships using molecular data
(Witek et al. 2008; Wey-Fabrizius et al. 2014 ).
Figure 1.1 illustrates the likely phylogenetic
T he taxon Gnathifera was erected based on relationships as a consensus phylogeny that is
morphological data by Ahlrichs ( 1995 , 1997 ). based on recent molecular as well as morpho-
The taxon comprises the Gnathostomulida logical data. In all gnathiferans, fertilization is
and Syndermata (which unites Rotifera, internal and the development direct. The para-
Acanthocephala, Seisonida) (Fig. 1 .1 ). With the sitic acanthocephalans have evolved additional
discovery of L imnognathia maerski (Kristensen dispersal stages that allow infection and transi-
and Funch 2000 ), the taxon Micrognathozoa has tions between hosts.
been included into the Gnathifera. The name
Gnathifera is based on the presence of a com-
plex jaw apparatus in the pharynx of all groups, GNATHOSTOMULIDA
except Acanthocephala (Sørensen 2 003 ; Funch
et al. 2 005) . Gnathifera are tiny, bilaterally Gnathostomulids are wormlike, microscopic,
symmetric animals that live in aquatic habitats. marine, interstitial animals that are covered with
Only the parasitic acanthocephalans reach body a monociliary epidermis. There are about 100
lengths of up to 80 cm. The acanthocephalans described species that are ordered into two
have lost many morphological characters as taxa, the elongated Filospermoidea and the
adaptations to their p arasitic lifestyle, includ- more compact Bursovaginoida (Sterrer 1 972 ;
ing the jaw apparatus and the digestive tract. Sørensen et al. 2 006 ). The animals have a mouth
Gnathifera has been placed in the Spiralia, opening that contains the pharyngeal bulb with
often affi liated with the Platyzoa (Funch et al. the cuticular jaw structure. Gnathostomulids
2005; Dunn et al. 2008; Witek et al. 2008 , 2 009 ; have a blind gut – some species possess a “tem-
Hejnol et al. 2009; Wey-Fabrizius et al. 2014 ). poral anus” (Knauss 1979 ) – an anterior brain, a
However, the sister group of Gnathifera remains ventral ganglion that is affi liated with the mouth,
unclear. Since gene sequences of most of the and one to three pairs of basiepithelial nerves
gnathiferan species seem to evolve fast, it (Kristensen and Nørrevang 1977 ; Müller and
remains diffi cult to resolve with confi dence the Sterrer 2 004 ).

Fig. 1.1 Gnathiferan phylogenetic relationships. Phylogenetic relationships of gnathiferan taxa according to recent
molecular and morphological studies (Images modifi ed from Funch et al. 2 005 )