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 Expanded Contents vii

4 7
The Poriferans The Ctenophores 135
and Placozoans 77 Introduction and General Characteristics 135
Phylum Porifera 77 Ctenophore Diversity 141
Introduction 77 Class Tentaculata 142
General Characteristics 79 Class Nuda 144
Poriferan Diversity 84
Class Calcarea 86
Class Demospongiae 86
Class Hexactinellida 87
8
Other Features of Poriferan Biology: Reproduction The Platyhelminths 147
and Development 88 Introduction and General Characteristics 147
Phylum Placozoa 89 Class Turbellaria 149
Introduction 89 Class Cestoda 156
General Characteristics 89 Class Monogenea 158
Class Trematoda 159
The Digeneans 159
Getting to a New Host 167
5 The Aspidogastreans (= Aspidobothreans) 169

Introduction to the
Hydrostatic Skeleton 95 9
The Mesozoans: Possible
Flatworm Relatives 179
6 The Mesozoans 179
Class Orthonectida 180
The Cnidarians 99 Class Rhombozoa 181
Introduction and General
Characteristics 99
Subphylum Medusozoa 102
Class Scyphozoa 102
10
Class Cubozoa 107 The Gnathifera: Rotifers,
Class Hydrozoa 109
Subclass Hydroidolina 109
Acanthocephalans, and
Order Siphonophora 112 Two Smaller Groups 183
The Hydrocorals 114 Introduction 183
An Unusual Group of Likely Medusozoans: Phylum Rotifera 184
the Myxozoa 115 Introduction and General Characteristics 184
A Medusozoan Sister Group? 116 Class Seisonidea 188
Class Staurozoa 116 Class Bdelloidea 191
Subphylum Anthozoa 117 Class Monogononta 191
Subclass Hexacorallia Other Features of Rotifer Biology 194
(= Zoantharia) 120 Digestive System 194
Subclass Octocorallia Nervous and Sensory Systems 195
(= Alcyonaria) 124 Excretion and Water Balance 195
viii Expanded Contents

Phylum Acanthocephala 196 Class Polychaeta 298

Phylum Gnathostomulida 198 Polychaete Reproduction 302
Phylum Micrognathozoa 198 Family Siboglinidae (formerly
the Pogonophora) 305

11 General Characteristics 305

Siboglinid Reproduction
and Development 311
The Nemerteans 205 The Echiurans 312
Introduction and General Characteristics 205 The Sipunculans 314
Other Features of Nemertean Biology 211 Class Clitellata 318
Classification 211 Subclass Oligochaeta 318
Protection from Predators 211 Oligochaete Reproduction 321
Reproduction and Development 211 Subclass Hirudinea 322
Leech Reproduction 325
Other Features of Annelid Biology 325

12 Digestive System 325

Nervous System and Sense Organs 327
The Molluscs 215 Circulatory System 328

Introduction and General Characteristics 215

Class Polyplacophora 218
Class Aplacophora 222
Class Monoplacophora 222 14
Class Gastropoda 224 The Arthropods 341
The Prosobranchs 231
Introduction and General Characteristics 341
The Opisthobranchia 233
The Exoskeleton 342
The Pulmonata 236
The Hemocoel 342
Class Bivalvia (= Pelecypoda) 237
Molting 343
Subclass Protobranchia 238
Nerves and Muscles 344
The Lamellibranchs 243
The Circulatory System 344
Subclass Anomalodesmata 251
Arthropod Visual Systems 345
Class Scaphopoda 254
Arthropod Reproduction 349
Class Cephalopoda 255
Classification Used in This Book 349
Other Features of Molluscan Biology 265
Subphylum Trilobitomorpha 350
Reproduction and Development 265
Class Trilobita 350
Circulation, Blood Pigments, and Gas
Subphylum Chelicerata 350
Exchange 266
Superclass Merostomata 351
Nervous System 266
Class Arachnida 352
Digestive System 269
Class Pycnogonida (= Pantopoda) 354
Excretory System 271
Subphylum Mandibulata 358
Superclass Myriapoda 358
Superclass Hexapoda 359
13 Class Insecta 359
Superclass Crustacea 373
The Annelids 295 Class Malacostraca 373
Introduction 295 Class Branchiopoda 379
Phylum Annelida 295 Class Ostracoda 381
General Annelid Characteristics 295 Class Copepoda 381
 Expanded Contents ix

Class Pentastomida 382

Class Cirripedia 389 18
Crustacean Development 389
Other Features of Arthropod Biology 392
Three Phyla of Uncertain
Digestion 392 Affiliation: Gastrotricha,
Excretion 395 Chaetognatha,
Blood Pigments 396
The Uncertainty of Arthropod
and Cycliophora 459
Introduction 459
Evolutionary Relationships 396
Phylum Gastrotricha 459
Phylum Chaetognatha 461
General Characteristics and Feeding 461
15 Chaetognath Reproduction 463
Chaetognath Lifestyles and Behavior 463
Two Phyla of Likely Arthropod Chaetognath Relationships 465
Relatives: Tardigrades Phylum Cycliophora 467

and Onychophorans 421

Introduction and General Characteristics 421
Phylum Tardigrada 422
19
Phylum Onychophora 424 The “Lophophorates”
(Phoronids, Brachiopods,
Bryozoans) and
16 Entoprocts 473
The Nematodes 431 Introduction and General Characteristics 473
Phylum Phoronida 474
Introduction and General Characteristics 431
Phylum Brachiopoda 476
Body Coverings and Body Cavities 432
Phylum Bryozoa (= Ectoprocta) 480
Musculature, Internal Pressure, and Locomotion 434
Class Phylactolaemata 482
Organ Systems and Behavior 436
Class Gymnolaemata 485
Reproduction and Development 438
Class Stenolaemata 488
Parasitic Nematodes 438
Other Features of “Lophophorate” Biology 489
Beneficial Nematodes 444
Reproduction 489
Digestion 490
Nervous System 491

17 Phylum Entoprocta (= Kamptozoa) 491

Four Phyla of Likely Nematode

Relatives: Nematomorpha, 20
Priapulida, Kinorhyncha, The Echinoderms 497
and Loricifera 451 Introduction and General Characteristics 497
Introduction 451 Class Crinoidea 500
Phylum Nematomorpha 452 Class Stelleroidea 503
Phylum Priapulida 454 Subclass Ophiuroidea 503
Phylum Kinorhyncha (= Echinoderida) 454 Subclass Asteroidea 505
Phylum Loricifera 456 Class Echinoidea 509
x Expanded Contents

Class Holothuroidea 513 Other Features of Urochordate Biology 545

Other Features of Echinoderm Biology 518 Reproduction 545
Reproduction and Development 518 Excretory and Nervous Systems 547
Nervous System 520 Subphylum Cephalochordata (= Acrania) 548

21 24
The Hemichordates 529 Invertebrate Reproduction
Introduction and General Characteristics 529 and Development—
Class Enteropneusta 530
Class Pterobranchia 534
An Overview 555
Introduction 555
Asexual Reproduction 556
22 Sexual Reproduction 558
Patterns of Sexuality 558
The Xenoturbellids: Gamete Diversity 560
Deuterostomes at Last? 537 Getting the Gametes Together 561
Larval Forms 565
Phylum Xenoturbellida 537
Dispersal as a Component of the Life-History
Pattern 577

23 Glossary of Frequently Used Terms 587

The Nonvertebrate Index 590
Chordates 539
Introduction and General Characteristics 539
Subphylum Tunicata (= Urochordata) 540
Class Ascidiacea 540
Class Larvacea (= Appendicularia) 542
Class Thaliacea 545
 Preface

About This Book level of detail to provide: I generalize wherever possible

to build a firm foundation without intimidation or con-
Invertebrate zoology is a fascinating but enormous field. fusion. Given a chance, the animals themselves soon win
More than 98% of all known animal species are inver- most students over.
tebrates, and that proportion is increasing with time as The most difficult part of writing this book has
more species are described. Invertebrates are distributed been deciding what to leave out. My decisions have been
among at least 30 phyla and a mind-boggling number reached largely by reading many dozens of research arti-
of classes, subclasses, orders, and families. The degree cles on the biology of each group and determining the
of morphological and functional diversity found within specific terminology and level of background information
some groups, even within single orders, can overwhelm that students will need to read those papers. Although
the beginning student. The enormous size of the field all phyla are covered, I have aimed for conciseness, not
and the great range of potential approaches to the subject exhaustiveness, and have emphasized unifying principles
make invertebrate biology challenging both to teach and rather than the diversity found within each group. Stu-
to learn. In preparing the seventh edition of this book, dents are best prepared to encounter the diversity of form
I have endeavored to make the tasks of both teaching and and function in lecture, laboratory, and field, once they
learning easier, and even enjoyable, while nevertheless have mastered basic concepts and terminology. I provide
presenting the latest thinking in the field. a sense of the ecological diversity encountered within
Too many people think of invertebrate zoology as an each group in a “Taxonomic Detail” section at the end of
exercise in memorizing terms and, perhaps, interesting but most chapters, a section that also adds to the value of the
trivial stories about interesting but irrelevant animals. Too book for reference.
many people think of invertebrate zoology as outdated, The text remains somewhat biased toward functional
and a field in which everything is already known. I hope morphology, bringing animals to life for students and
that this book alters those perceptions by presenting preparing them to make careful observations of living
invertebrate zoology as a lively area of ongoing and worth- animals in the laboratory and in the field. Most chapters
while modern biological inquiry. contain a section entitled “Topics for Further Discus-
Like previous editions, the seventh is designed as a sion and Investigation,” highlighting many of the major
nonintimidating, readable introduction to the biology research questions that have been and are being addressed
of each group, emphasizing those characteristics that set for the animals covered in each chapter. For each topic,
each group apart from all others. The book is intended I have selected references from the primary literature
to serve as the foundation for further learning—in lec- that should be intellectually accessible to any interested,
ture, laboratory, field, and library—a foundation that beginning student once he or she has read the relevant
is largely manageable by students. Instructors are then textbook chapter. I have had to exclude many excellent
free to embellish and expand on that foundation to suit papers because they were too advanced, were published
any desired focus: taxonomy and phylogeny, behavior, in less widely distributed journals, or were review papers
conservation, environmental biology, diversity of form rather than primary journal articles. The topics I have
and function, physiology, ecology, or current research in chosen, along with the accompanying references, could
any of those areas. The book whets the student’s appetite be used as a basis for lectures, class discussion, term
and provides the required background, buying instruc- papers, research proposals, or other writing assignments,
tors the time to discuss more fully whatever they feel are or simply as a convenient way of easing students into the
the most interesting and important aspects of the field. original literature by having them investigate topics that
This is the guiding principle in my decisions about the excite their curiosity. Students gain little by reading and

xi
xii Preface

memorizing predigested summaries of the primary lit- session; it is always easier to assign additional sections of
erature; they gain much by reading and discussing that a text than to tell students what not to read within a larger
literature. Similarly, I have again decided against adding section. I find that periodic scheduled quizzes, designed
“end-of-chapter summaries,” arguing that students will to reward students for doing the assigned reading, pro-
learn far more of lasting value by writing and discussing vide excellent motivation. The final chapter (Chapter 24)
their own summaries than by memorizing mine. brings together all of the major phyla by considering
The excitement of invertebrate biology is found in the general principles of invertebrate reproduction and devel-
primary research literature, and a major goal of this book opment, providing students an opportunity to reminisce
is to motivate and prepare students to read that literature— about all the animals that they have encountered during
both the recent literature and that of past decades, before the the term and to begin moving beyond phylum-by-phylum
words “synapomorphy” and “Lophotrochozoa” were com- compartmentalization toward synthesis.
mon in the literature, and when most workers still thought This edition has a somewhat greater phylogenetic
that Echiura was a phylum and that myxozoans were mem- orientation than its predecessors, but it still avoids
bers of the Protozoa. The Research Focus Boxes scattered prolonged phylogenetic discussion. There is still no con-
throughout the book are based on individual papers drawn sensus concerning many invertebrate interrelationships:
from the primary literature to illustrate the range of questions Developmental studies, molecular techniques, and cladis-
that biologists have been asking about invertebrates and the tic analyses continue to revolutionize our thinking about
variety of approaches that have been used to address those evolutionary relationships, or at least to challenge many
questions. My goal here is to prepare students to read the treasured assumptions. Annelids and arthropods may or
primary literature by focusing on how questions are formu- may not be closely related, segmentation may be a derived
lated, how data are collected and interpreted, and how each rather than an ancestral character in molluscs, the ances-
study typically leads to further questions. Students inter- tral mollusc may have more closely resembled a bivalve
ested in the topic of a particular Research Focus Box might than a gastropod, nematodes are probably closely related
wish to read the original paper on which that Focus Box was to arthropods than to rotifers, insects may have evolved
based and then use that Focus Box as a model for summariz- from crustacean ancestors, leeches and earthworms may
ing other papers on related topics. For Focus Boxes based be derived polychaetes, and phoronids may be modified
on older research articles, interested students may wish to brachiopods. Similarly, nemertine worms, long consid-
follow the topic forward in time, using an indexing service ered acoelomates, actually may be unusual coelomates
such as the Web of Science. Many instructors now ask their with no direct flatworm affinities. Acoels may be primitive
students to write Research Focus Boxes modeled on the ones but may not belong in the phylum Platyhelminthes.
in this book; such assignments can teach students much of For some workers, even the definition of what it
lasting value, particularly if they include guided revision— means to be a protostome has shifted substantially over
not an onerous chore for either student or instructor since the past few years and now relies less on anatomical and
each Focus Box is only a few pages long. This assignment developmental criteria. Indeed, the usefulness of mor-
allows students to put their growing vocabulary to immedi- phology and ultrastructure in inferring phylogenetic
ate use, and it doesn’t seem to hurt, either, that students feel relationships has been seriously challenged in recent
that they’re learning useful skills in the process. years. “Lophophorates,” for example, appear to be deu-
As further incentive for both instructors and students, terostomes based on morphological and developmental
Bob Podolski at the College of Charleston has created criteria, but group unambiguously with protostomes in
a Research Focus Box (RFB) posting area on the Society most molecular analyses. Similarly, molecular data now
for Integrative and Comparative Biology website (Division align chaetognaths with protostomes, despite their many
of Invertebrate Zoology): http://www.sicb.org/dl/rfb.php. deuterostome developmental characteristics. The rela-
Now students can write for a real audience, not just for tionships between cephalochordates, chordates, and echi-
their instructors. Instructors are encouraged to post their noderms are also uncertain: Some recent molecular data
students’ best RFB’s on the website for others to read. suggest that cephalochordates are more closely related to
As with previous editions, chapters are self-contained echinoderms than to other chordates, implying that some
and can be assigned in whatever order best suits the orga- key chordate features were present in the ancestral deu-
nization of any particular course, once the introductory terostome and later lost in the evolution of echinoderms
chapters have been covered. In my own course, I cover and hemichordates. And molecular analyses, often in
some introductory material (particularly chapters 1 and 5) concert with careful ultrastructural studies, have mostly
and then begin with annelids and other protostomes. destroyed the idea that pseudopodia and flagella inform
Within each chapter, the material has been arranged in us about protozoan relationships.
manageable, readable units for the convenience of both Clearly, invertebrate systematics is a work in progress.
student and instructor. For example, a section entitled Although the phylogenetic atmosphere is charged with
“Introduction and General Characteristics” might be excitement, beginning students typically view textbook
assigned prior to a lecture on a particular group of organ- discussion of such controversies as simply another set of
isms, while a section called “Feeding and Digestion” might facts to be memorized. For this reason, such issues are
best be assigned prior to the accompanying laboratory best treated in lecture, where they can be used to animate
 Preface xiii

class discussion. Indeed, the controversies surrounding for some other animals, including the tardigrades; indi-
phylogenetic speculation are what make phylogeny inter- cations of an association with nematodes seems to have
esting, and can be used to make the animals themselves resulted from a long-branch attraction problem which,
interesting. Chapter 2 introduces students to the range once resolved, restores the tardigrades to the neighbor-
of approaches used in reconstructing phylogenies, and hood of arthropods and onychophorans. In addition, the
includes substantial discussion of cladistic analysis and the sponges now include 4 classes rather than 3—the Homo-
promise and potential pitfalls associated with the incorpo- scleromorpha have been moved out of the Demospongiae
ration of molecular data. The book provides the founda- to form a class of their own—Velella and Porpita are no
tion upon which instructors and students can build. longer in separate hydrozoan orders, and the Class Poly-
Whenever possible, I provide “Defining Character- chaeta may no longer be a valid taxonomic category. The
istics” as each new animal group is introduced, to help gnathostomulids have come to settle comfortably as close
students keep track of features separating each group of relatives of rotifers and acanthocephalans within the new
animals from other groups at the same taxonomic level. clade Gnathifera, and the xenoturbellids seem to be deu-
In essence, these defining characteristics are synapomor- terostomes, bringing the total number of extant deutero-
phies. For some groups there are no clear defining char- stome phyla to four.
acteristics, or characteristics that have been proposed are Several chapters have been largely rewritten. In
too controversial to be included at the present time. particular, I have reorganized the material on body cav-
Most chapters conclude with a section entitled ities in Chapter 2 to better emphasize the protostome-
“Search the Web,” guiding students to particularly good deuterostome distinction. I also have updated the section
websites associated with the group under discussion. “Inferring Evolutionary Relationships” in that chapter to
I have listed only those sites that speak with verifiable incorporate more of the modern molecular approaches.
authority and that are likely to be around and updated for In addition, I have updated the biological information
a number of years. about many of the animal groups discussed in this book.
For example, I now discuss recent information about the
Changes for the Seventh Edition role of sexual reproduction in the life cycle of cellular
slime molds in Chapter 3. Moreover, it seems clear that
When I finished the second edition of this book in 1984, the definitive host in the myxozoan life cycle is usually
I thought that it might need to be revised perhaps once or an invertebrate, with vertebrates usually serving as inter-
twice in my lifetime. A series of careful developmental stud- mediate hosts—the opposite of what occurs in the trema-
ies in concert with the increasing acceptance and use of tode life cycle. In Chapter 6, I now draw more attention
cladistic methodology and molecular data in phylogenetic to the Staurozoa, given their possible basal position
analyses have made the past 30 years far more exciting than within the Medusozoa. Throughout the book, I have
I had imagined they would be, and the changes to this edi- updated the General References sections and references
tion are substantial, despite the passage of only 4 years since listed in the “Topics for Further Discussion and Inves-
I completed the previous edition. tigation” sections, and have added new topics to several
Most chapters have been revised to reflect new dis- chapters. I also have updated material in the “Taxonomic
coveries and expanding research areas, including research Detail” sections of many chapters. For example, students
of commercial importance and environmental relevance. now can learn about a remarkable crustacean parasite
Remarkably, all of the rather dramatic organizational that destroys the tongue of its fish host and then takes the
changes that I made for the previous editions have held tongue’s place with its own body.
up, and indeed have generally been bolstered by additional The current edition deals somewhat more conspicu-
evidence. The pogonophorans, echiurans, and sipuncu- ously with biomedical relevance, biological invasions,
lans, for example, for many years treated as 3 separate habitat degradation, and other contemporary issues. I
phyla, are all now widely accepted as modified annelids have written several new Research Focus Boxes, added
based upon morphological, developmental, and molecular new “Topics for Further Discussion and Investigation”
data. Support has also generally increased for separating sections to some chapters, and included many new ref-
protostomes into at least two great groups—the Ecdysozoa erences to the recent primary literature, including many
and the Lophotrochozoa—although there is not yet com- published as recently as 2013. I also have added some won-
plete agreement about exactly which animals each group derful new websites to the Search the Web sections that
contains or how the animals within each group are related close each chapter, and have written one new invertebrate
to each other. riddle. In addition, since learning about invertebrates
This edition contains some impressive new changes. requires students to acquire a substantial new vocabulary,
For example, I now discuss the increasingly likely pos- I have added a new online resource to give students some
sibility that insects have evolved directly from crusta- ideas for including that new vocabulary in their everyday
cean ancestors, and that acoel and nemertodermatid conversation: Talking about Invertebrates (See left side
flatworms may belong in a separate phylum, the Acoe- of web page: http://ase.tufts.edu/biology/labs/pechenik/
lomorpha, which may have little relationship with other publications/); send your entries to me and I’ll post them
flatworms. I have also updated phylogenetic relationships for all to see.
xiv Preface

The paradigm shifts that have occurred over the past Miller, for updates on tardigrade species numbers; and
25 years or so have been truly remarkable, and there is no end Jens Høeg and Gonzalo Giribet, for their helpful com-
in sight. If insects really evolved from crustacean ancestors, ments on the arthropod material.
for example, as recent molecular and morphological studies I would also like to thank the many people who con-
indicate, that will require a substantial redefinition of what tributed their expertise to previous editons of the text:
it means to be a crustacean. Similarly, there is now substan-
tial molecular evidence that oligochaetes and leeches evolved Frank E. Anderson, Southern Illinois University;
from polychaete ancestors, which would essentially make Christopher J. Bayne, Oregon State University; John
the Polychaeta the equivalent of the Annelida, and a study in F. Belshe, Central Missouri State University; Yehuda
2007 using expressed sequence tags has supported the inclu- Benayahu, Tel Aviv University; Bill Biggers, Wilkes
sion of Ectoprocta and Entoprocta within a single phylum, University; Robert Bieri, Antioch University; Chip
the Bryozoa, bringing us back to a classification established Biernbaum, College of Charleston; Brian L. Bingham,
over 100 years ago. Finally, although recent molecular stud- Western Washington University; Susan Bornstein-Forst,
ies using individual molecules continue to add support for Marian College; Kenneth J. Boss, Harvard University;
the Ecdysozoa-Lophotrochozoa dichotomy, as noted earlier, Barbara C. Boyer, Union College; Robert H. Brewer,
a recently reported whole-genome study of nine eukary- Trinity College; Maria Byrne, University of Sydney
otic species does not support that arrangement, but favors C. Bradford Calloway, Harvard University; Ron
instead the older Articulata hypothesis in which arthropods Campbell, University of Massachusetts at Dartmouth;
are more closely related to annelids than to nematodes. John C. Clamp, North Carolina Central University;
As more molecular data from more species and more Clayton B. Cook, Bermuda Biological Station for Research;
genes from each species are collected, we should see John O. Corliss, University of Maryland; Bruce Coull,
increasing stability in the accepted arrangements. There University of South Carolina
is good reason to hope. Ferenc A. deSzalay, Kent State University; Don Diebei,
As always, I welcome constructive criticism from all Memorial University of Newfoundland; Ronald V. Dimock,
readers, both instructors and students. Jr., Wake Forest University; William G. Dyer, Southern
[email protected] Illinois University, Carbondale
David A. Evans, Kalamazoo College
Daphne G. Fautin, University of Kansas; Paul Fell,
Acknowledgments Connecticut College; Ben Foote, Kent State University;
Bernard Fried, Lafayette College; Peter Funch, University
It is a great pleasure to thank the reviewers who have of Århus
helped to make this a better text. Audrey Gabel, Black Hills State University; James R.
Garey, University of South Florida; Ann Grens, Indiana
Douglas Lipka
University, South Bend
William Carey University
Kenneth M. Halanych, Woods Hole Oceanographic In-
John Lugthart stitution; G. Richard Harbison, Woods Hole Oceanographic
Dalton State University Institution; Norman Hecht, Tufts University; Gordon
Hendler, Natural History Museum of Los Angeles County;
Joshua Mackie
Anson H. Hines, Smithsonian Environmental Research Center;
San Jose State University
Edward S. Hodgson, Tufts University; Alan R. Holyoak,
Brian Steinmiller Manchester College; Duane Hope, Smithsonian Institution;
Waynesburg University William D. Hummon, Ohio University, Athens
William W. Kirby-Smith, Duke University; Robert E.
Janice Voltzow
Knowlton, George Washington University
University of Scranton
John Lawrence, University of South Florida; William
William Woods Layton, Dartmouth Medical School; Herbert W. Levi,
Tufts University Harvard University; Gail M. Lima, Illinois Wesleyan
University; B. Staffan Lindgren, University of Northern
I am also grateful to Gordon Hendler, for detailed advice British Columbia
on all things echinoderm; Tom Cavalier-Smith and Laura Robert Knowlton, Western Illinois University; James
Katz, for suggestions about how to best arrange the proto- McClintock, University of Alabama at Birmingham;
zoans in Chapter 3; Sally Leys, for detailed comments on Rachel Ann Merz, Swarthmore College; Nancy Milburn,
my revision of Chapter 4; Beth Okamura, for her patience Tufts University
in answering all of my questions about myxozoans; Gary Diane R. Nelson, East Tennessee State University; Claus
Rosenberg, for updates on gastropod species numbers; Nielsen, University of Copenhagen; Michele Nishiguchi,
Christer Erséus and Thorten Struck, for updates on an- New Mexico University -Las Cruces; Jon Norenburg, Smith-
nelid systematics and species numbers; William (Randy) sonian Institution
 Preface xv

Steve Palumbi, University of Hawaii; Lloyd Peck, Jon D. Witman, Marine Science Institute–Northeastern
British Antarctic Survey; John F. Pilger, Agnes State University; William Woods, Tufts University; Robert
College; William J. Pohley, Franklin College of Indiana; Woollacott, Harvard University
Gary Polis, Vanderbilt University; Rudolph Prins, Russel L. Zimmer, University of Southern California
Western Kentucky University
Mary Rice, Smithsonian Institute; Robert Robertson, I am also indebted to many other colleagues around
The Academy of Natural Sciences; Pamela Roe, California the world who cheerfully provided their stunning photo-
State University, Stanislaus; Frank Romano, Jacksonville graphs for this and previous editions, answered specific
State University queries in person or by telephone and e-mail, or com-
Amelia H. Scheltema, Woods Hole Oceanographic mented on drafts of new Research Focus Boxes that
Institution; J. Malcolm Shick, University of Maine; Owen I based on their original research papers.
D. V. Sholes, Assumption College; Terry Snell, University I am grateful to all of these people: their devotion to,
of Tampa; Eve C. Southward, Plymouth Marine Labora- enthusiasm for, and knowledge about invertebrate biology
tory (U.K.) is both inspiring and humbling.
John Tibbs, University of Montana, Missoula Finally, I am happy to thank Steve Wainwright for
Steve Vogel, Duke University getting me off to a good start. Chapters 1 and 24 of this
J. Evan Ward, Salisbury State University; Daniel book are based on papers I wrote as an undergraduate for
Wickham, University of California–Bodega Marine his Animal Diversity course at Duke University. Students:
Laboratory; Edward O. Wilson, Harvard University; never throw your schoolwork out; you just never know!
Visit https://testbankfan.com
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 2013 Guide to the Major Animal Groups
(aschelminth groups are shaded blue)
(P = Platyzoa)
Invertebrates Animalia Protostomia Spiralia Lophotrochozoa Panarthropoda Ecdysozoa Cycloneuralia Deuterostomia
Protozoan
Protists
Porifera
Placozoa
Cnidaria
Ctenophora P
Gastrotricha P
Rotifera P
Acanthocephala P
Gnathostomulida P
Platyhelminthes P
Mesozoa P
Nemertinea
Mollusca
Sipuncula
Annelida
Bryozoa
Brachiopoda
Phoronida
Entoprocta
Cycliophora
Arthropoda
Tardigrada
Onychophora
Nematoda
Nematomorpha
Kinorhyncha
Priapulida
Loricifera
Chaetognatha
Echinodermata
Hemichordata
Chordata
Xenoturbellida
 The Geologic Time Scale (MYA)
(major global extinctions are shaded blue)

Estimated
% of genera
Eons Eras Periods Epochs going extinct*

Quaternary Holocene
Pleistocene MYA
1.6
Pliocene
5
Miocene

Cenozoic
24
Tertiary Oligocene
37
Eocene
58
MYA Paleocene
66 47% ± 4%
Cretaceous
Mesozoic

190
Jurassic
205 53% ± 4%
Triassic
Phanerozoic

250 82% ± 4%
Permian
290
Pennsylvanian
(Late Carboniferous)
325
Mississippian
Paleozoic

(Early Carboniferous)
355
57% ± 3%
Devonian
410
Silurian
438 60% ± 4%
Ordovician
510
Cambrian
544
Precambrian

*Based on Sepkoski, J. J. Jr. 1996. In: O. H. Walliser, ed. Global Events and Event Stratigraphy.
Springer: Berlin, pp. 35–51.
MYA = Millions of years ago
This page intentionally left blank
 1
Introduction and Environmental
Considerations
Introduction: The Importance
of Research on Invertebrates
“We need invertebrates, but they don’t need us.”
E. O. Wilson. 1987. The little things that run the
world (the importance and conservation of
invertebrates).
Conservation Biology 1:344–346

It surprises me that people don’t want to learn more about

invertebrates. For one thing, the diversity of invertebrate
form and function is truly astounding. In addition, much
fascinating and important research has been conducted
and continues to be conducted using invertebrates.
Many diseases of humans and of the animals and
plants upon which we depend are caused by invertebrates,
either directly or indirectly, and invertebrates play critical
roles in most food webs in all habitats. Studies on vari-
ous invertebrate species have taught us much of what we
presently know about the control of gene expression,
mitosis, meiosis, and regeneration; the design of gene
regulatory networks in embryonic development; aging,
programmed cell death, wound repair, and regeneration;
the mechanisms of pattern formation during embryonic
development; the control and consequences of phenotypic
plasticity, in which a single genotype can produce differ-
ent phenotypes under different environmental conditions;
the evolutionary history of hemoglobin and ecdysteroid
function; fertilization and chemoreception; the transmis-
sion of nerve impulses; the biochemical basis of learning
and memory; the biology of vision; and the biochemical
and genetic basis for predisposition to some major dis-
eases (e.g., type II diabetes). Much of what we know about
the mechanisms by which genetic diversity originates, is
maintained, and is transmitted to succeeding generations
also comes through the study of invertebrates, as do many
basic principles of animal behavior, development, physi-
ology, ecology, and evolution. Similarly, molecular studies
on various invertebrate species are rapidly increasing our
2 Chapter 1

understanding of the genetic basis for evolutionary shifts This book, together with lectures and laboratory
in morphology and life history, including the possible role sessions, opens the door to the great and growing litera-
of horizontal gene transfer, in which sets of genes may be ture of invertebrate biology.
transferred intact from one species to another, and the
role that such transfer may play in evolution. Certain in- Environmental Considerations
vertebrate species recently have become key models for
understanding the evolution of the vertebrate brain. The organisms considered in this book are grouped into
In addition, modern research on invertebrates is more than 30 phyla. The members of almost half of these
helping to unravel the story of how immune recognition phyla are entirely marine, and the members of the remain-
systems evolved and how they work. Interest in certain ing phyla are found primarily in marine and, to a lesser
invertebrates as biological agents for controlling various extent, freshwater habitats. Excluding the arthropods,
agricultural pests and as sources of unique chemicals of invertebrates generally have been far less successful in in-
potential biomedical and commercial importance also is vading terrestrial environments. Even those invertebrate
increasing. Some of the substances isolated from marine species that are terrestrial as adults often have aquatic
sponges, for example, promise to be potent antitumor developmental stages. It is worthwhile, therefore, to con-
agents, and others isolated from certain spiders and ven- sider some of the physical properties of water—both fresh
omous snails are providing neurobiologists with highly and salt—to discover why so many species are aquatic for
specific chemical probes for studying key aspects of all or part of their lives. The physical properties of salt
nerve and muscle function, such as how ion channels are water, freshwater, and air play major roles in determining
opened and closed. Still other substances derived from the structural, physiological, and behavioral characteris-
invertebrates show considerable promise as instant ad- tics displayed by animals living in various habitats.
hesives (glues produced by onychophorans and barnacles
and by some spider and bivalve species, for example) and Air Is Dry, Water Is Wet
anticorrosion agents (e.g., barnacle cements). Detailed
studies of crustacean and insect navigation and locomo- Air is dry, whereas water is wet. As trivial as this statement
tion, and how that locomotion is controlled and coordi- may seem, the repercussions with respect to morphology,
nated, may lead to the design of new robots, both flying respiratory physiology, nitrogen metabolism, and repro-
and crawling, macro and micro; studies on the optical ductive biology are tremendous, as seen in Table 1.1.
properties of certain sponge fibers may lead to the manu- Because aquatic organisms are in no danger of dry-
facture of more effective fiber optic cables; and detailed ing out, gas exchange can be accomplished across the
studies of how echinoderms form their remarkable cal- general body surface. Thus, the body walls of aquatic in-
cite crystals may have similarly sophisticated engineering vertebrates are generally thin and water permeable, and
applications. any specialized respiratory structures that exist may be
Invertebrates also have become widely used to evalu- external and in direct contact with the surrounding me-
ate and monitor pollutant stress in aquatic environments, dium. Gills, which can be structurally quite complex, are
and the rapid loss of invertebrate species from both terres- simply vascularized extensions of the outer body wall.
trial and aquatic habitats is gaining increasing attention in These extensions increase the surface area available for
biodiversity studies. gas exchange and, if they are especially thin walled, may
The recently documented phenomenon Colony also increase the efficiency of respiration (measured as
Collapse Disorder, in which hundreds of thousands of the volume of gas exchanged per unit time per unit area).
honeybees simply abandon their hives and disappear, is In contrast to the minimal complexity required for
worrisome: apples, almonds, and approximately 90 other aquatic respiratory systems, terrestrial organisms must cope
crops in the United States depend on honeybees for pollina- with potential desiccation (dehydration). Terrestrial species
tion, accounting for some $15 billion in annual sales. Bum- relying on simple diffusion of gases through unspecialized
blebees, which pollinate some 15% of commercial crops in body surfaces must have some means of maintaining a
the U.S., are also in serious decline. Similarly ominous is moist outer body surface, as by the secretion of mucus in
the recently documented increase in the acidity of seawater earthworms. Truly terrestrial invertebrates generally have a
in the world’s oceans, as discussed in the next section. water-impermeable outer body covering that prevents rapid
Finally, there is a growing concern about the increased dehydration. Gas exchange in such species must be accom-
spread of various invertebrate species into nonnative habi- plished through specialized, internal respiratory structures.
tats, and increasing attention is being paid to the mecha- The union of sperm and egg, and the subsequent de-
nisms of transport and to the ecological impact of such velopment of a zygote, can be achieved far more simply by
biological invasions. Of course, nobody yet knows the aquatic invertebrates than by terrestrial species. Marine
consequences of continued pollution, biological invasions, organisms, in particular, may shed sperm and eggs freely
and global climate change on food web function, in either into the environment. Because the gametes, embryos, and
aquatic or terrestrial food-webs; probably there is only larvae of marine species are not subject to dehydration
one way to do the experiment, and we’re all participating. or to osmotic stress, fertilization and development can
Random documents with unrelated
content Scribd suggests to you:
 Why, then, without doubt
It was Vivien! But yet do you know
‘Tis the Eve of Saint John, and here, last night,
I dreamed that I saw my dream again!

[The hand and arm of the statue fall, broken, to the

ground at the feet of The Marquise.

The Marquise

Ah!

De Vardes (pushes the marble aside with his foot)

It is nothing! The stone was cracked last night.

Some crack-brained peasant had no better mark!

The Marquise

‘Tis a présigne!—I feel it.—

De Vardes

You shudder!

The Marquise

One trod near my grave! I’m suddenly cold!

De Vardes

The sun never shines on this terrace!

 The Marquise

No!
‘Twas an air from the Forest of Paimpont
Came over me!

[Voices within. De L’Orient sings.

De L’Orient

In Ys they did rejoice,

In Ys the wine was free;
The Ocean lent its voice
Unto that revelry!

The Marquise

Oh, come away!

Let us find the violins and the sun!
There are other woods than Paimpont. Come away!

[Exeunt De Vardes and The Marquise.

Yvette (leaves the shadow of the statue)

‘Twas he! That horseman who did waken me

That Saint John’s Eve I strayed in Paimpont Wood!
O Our Lady—

Séraphine (from the statue)

Saint Yves! There is bread!

 [Yvette takes from the table a loaf of bread and
throws it to Séraphine, who springs upon it like a
famished wolf.

Ah—h—h!

[Setting her teeth in the loaf.

[Yvette, about to lay her hand upon another round of

bread, sees the fan lying upon the cloth. She leaves
the bread and takes up the fan. It opens in her hand.

Yvette

Oh!—

[She sits in the great chair and waves the fan slowly
to and fro.

Were I a lady fair and free,

I would powder my hair with dust of gold,
I would clasp a necklace around my throat,
Of jewels rare, and a gown I would wear,
Blue silk like Our Lady of Toute Remède!
My shoes should be made of golden stuff,
And a broidered glove should dress my hand,
My hand so white that a lord might kiss!
I would spin fine flax from a silver wheel,
I would weave a web for my bridal sheets,
I would sing of King Gradlon under the sea,
Were I a lady fair and free!

Enter Grégoire.

Séraphine (from the statue)

 Yvette!
Yvette!

Yvette

Peace, peace!

Grégoire

What have you there?

Yvette

A fan.
So long I’ve wanted one!

Grégoire

A fan, forsooth!
You cannot eat a fan, drink it, wear it!

Yvette

I would look on’t.

One day at Vannes the deputy’s sister
Showed me a fan, but it was not like this!
Oh, not like this with these wreaths of roses,
These painted clouds, this fairy ship!

Grégoire
 The price
Would keep a peasant from starvation!
And belike it fell from the lifted hand
Of Madame la Marquise de Blanchefôret!

[The fan breaks in Yvette’s hand.

Séraphine (leaving the statue)

Thou evil-starred!

Yvette

What have I done?

Grégoire

Diantre!
Now you will be beaten as well as hanged!

Yvette

She called us miserable brigands!

Enter De Vardes.

Séraphine

Saint Yves! Saint Hervé! Saint Herbot!

De Vardes (to Grégoire)

 Voices?

Grégoire

Monseigneur?

De Vardes

The fan of Madame la Marquise.

Grégoire

Monseigneur?

De Vardes (perceiving Yvette and Séraphine)

What will you have, good people?

Séraphine

Saint Guenolé! Saint Thromeur! Saint Sulic!—

He did not see us in the dark last night!

[De Vardes regards them more closely.

Grégoire

Séraphine Robin—Yvette Charruel—

They are not bad folk, monseigneur!

Séraphine
 No, faith!

[De Vardes studies the name written upon a playing

card which he holds in his hand.

De Vardes (to Grégoire)

Say to Monsieur the Deputy from Vannes

That I await him here.

[Exit Grégoire. De Vardes looks intently at

Yvette.

Yvette

It was so beautiful,
The fan—I took it in my hand—it broke!

Séraphine

All that she touches breaks!

De Vardes (to Yvette)

Wast ever thou

In the Forest of Paimpont?

Yvette

Oh, monseigneur!
Last Eve of Saint John, by the Druid Stone!

De Vardes
Ah!—

[He takes the fan from Yvette’s hand and examines it.

Beyond all remedy!—Well, ‘tis done.

Do not tremble so!

Yvette

I tremble not!

Enter Lalain.

Séraphine (to Yvette)

Here’s Monsieur Lalain!

Yvette

I care not, I!

De Vardes

Ah,
Rémond Lalain!

Lalain (stiffly)

Monsieur—

De Vardes
 A moment, pray,
Until I’ve spoken with these worthy folk!

Lalain (coldly)

Monsieur the Baron’s pleasure!

[He moves aside, but in passing speaks to Yvette.

Yvette! Yvette!

Yvette

Monsieur the Deputy?

Lalain

Too fair art thou!

Beware! This is the Seigneur of Morbec!

Yvette

I know.

Lalain

He is the foe of France!

Yvette

I know.
 De Vardes (to Séraphine)

Your business, well?

Séraphine (stammering)

Our business, monseigneur?—

Oh, give me help, Saint Yves le Véridique!—
Our business?—Saint Michel!—Well, since we’re here!—
Monseigneur, was the pullet plump and sweet?

De Vardes

The pullet?

Yvette

Our pullet, monseigneur.

Lalain

Distrained for rent!

Séraphine

And Lisette, monseigneur?

May we enquire for Lisette’s health?

De Vardes

Lisette?
 Yvette

Our cow, monseigneur.

Lalain

Taken for taxes!

Séraphine

It was the best Lisette!

Yvette

She followed me
Through the green lanes, and o’er the meadows salt.
Her breath was sweet as May!

De Vardes

It would please you

To have your cow again?

Yvette

Oh, monseigneur!
Monseigneur, I’m the herd girl of Morbec!

Lalain (aside)

They gaze into each other’s eyes!

 De Vardes

What is
Thy name?

Yvette

Yvette.

Séraphine

Ay, ay, ‘tis so!—Yvette.

Called also The Right of the Seigneur!—

De Vardes

The Right of the Seigneur!

Séraphine (nodding)

Just so.

Lalain (aside)

Recall
Just one of a great seigneur’s privileges!
Baiser des mariées, in short, my friend!

Séraphine
O holy Saints! the night that she was born!
The thunder pealed, the sea gave forth a cry,
The forked lightnings played, the winds were out
And in the hut her mother lay and wailed,
And called on all the saints, the while Jehan
(That was her mother’s husband, monseigneur),
He stood and struck his heel against the logs.
Up flew the sparks, for all the wood was drift,
Salt with the sea, and every flame was blue.
I held the babe—Yvette, show monseigneur
The mark beneath the ear!

Yvette

No!

Séraphine

Stubbornness!
‘Tis there!

Lalain

A birthmark—a small blue flower!

De Vardes

Ah!

Séraphine
 Ay! a little mark.—Jehan Charruel!
He was a violent man,—the sea breeds such!
He cursed Yvonne upon her pallet there,
So pale she was, and dying with the tide!
He cursed the saints, the purple mark, the babe,
And some one else I dare not name—

Lalain

I dare!
Henri-Etienne-Amaury de Vardes,
Late Baron of Morbec!

Séraphine
 Then out he goes,
A-weeping hard—Jehan—into the night.
Ouf! how it blew!—
The sea ran high, he met it in the dark,
Was drowned! Yvonne went with the ebb. Behold
Yvette!

[Séraphine retreats to the table, where she furtively

drinks from a half-emptied wineglass. Lalain follows
her and the two talk together.

De Vardes

That purple flower, that violet

By nature limned upon thy slender throat,—
From north to south, from east to west ‘tis known!
A De Vardes bore that mark at Poitiers.
The marshal, Hugues the Fair, and black Arnaud,
The late baron—Why, what hast thou to do
With burning down châteaux to make a light
To show the Morbihan that purple flower?

Yvette

O Our Lady of Thorns!

De Vardes

Herd girl too fair!

And vision of Paimpont, fair as I dreamed!
How fair was thy errand last night?

Yvette

Monseigneur!
 De Vardes

In the ashes of Morbec what shouldst thou find?

Yvette

We only wished to make a little light—

A little light to let the neighbours know
That we were hungry!

De Vardes

What neighbours hast thou?

Yvette

Normandy and Maine, Anjou and Poitou,

The sea, the sky, and somewhat far away,
The Club of the Jacobins at Paris.

De Vardes

Thy father was a nobleman of France!

Yvette

I never had a father, monseigneur!

I had a mother, and she loved, they say,
She dearly loved the fisherman Jehan!
When for the dead I pray, I pray for them.

De Vardes

How old art thou?

 Yvette

How old? Ah, let me see!

[She counts upon her fingers.

The year the hailstones fell and killed the wheat;

The year the flax failed and we made no songs;
The year I begged for bread; the bitter year
We buried Louison who died of cold,
And Jacques was hanged who shot the seigneur’s deer;
The Pardon of Sainte Anne I had a gown;
Came Angélique from Paris, told us how
The wicked Queen was smiling, smiling there;
Justine pined away, they shot Michel If,
Down fell the Bastille, I learned Ça ira;
The deputy came to the curé’s house,
Beside the deep blue sea I walked with him.
A day there was at Vannes, a glorious day,
When music played, and every banner waved,
And all the folk went mad and rang the bells!
Vive la Révolution! Vive Mirabeau!
Vive Rémond Lalain! I wept when ‘twas o’er,
Last summer was so fair! I wandered far,
One day I wandered through a darksome wood—
‘Twas on the Eve of good Saint John, I know!

De Vardes

Ah—

Yvette
 The summer fled, the light, the warmth did go,
The winter came that was so cruel cold,
Cold as the dead! And hunger, monseigneur,
With bread at the château!—Died Baron Henri.—
The summer came again, the roses bloomed,
The roses bloomed, but they were not for us!
For us the dank seaweed, the thorny furze.
The lark sang well, but ah, it sang too high!
We could not lift our hearts to heaven’s gate;
We only heard the wind moan at our door.
We cried to the saints, but they took no heed!
One told us what they did at Goy and Vannes,
At Goy and Vannes, pardieu! they helped themselves!
We heard there had come a new lord to Morbec,
A soldier and a stranger to us all!
Three days have gone since I did sit alone
Upon the cliff edge in the waving grass;
The mew and curlew cried, the night wind blew,
And in the sunset glow red turned Morbec!
I thought of my mother, I thought of France,
I looked at the château cruel and high,
And as I was hungry I ate my black bread!—
I think, monseigneur, that I am nineteen.

De Vardes

Pauvre petite!

Yvette

Ah, poor indeed!

De Vardes

How dark
Thine eyes!
 Yvette

My mother’s were darker, they say!

De Vardes

Thy face is the face of a picture there.

Yvette

I know—the Duchess Jeanne, who died for love.

De Vardes

Did Vivien teach thee magic in the wood?

Yvette

Monseigneur?

De Vardes

Pauvre petite!

Yvette

O Our Lady!
The roses smell so sweet—

[Lalain comes forward.

Lalain
 I pardon crave,
But I must sup to-night at Rennes. Please you,
Release this peasant girl! Affairs there are
Of which I’d speak—

De Vardes

Ay, presently!

Lalain

Now!

De Vardes

Monsieur!

Lalain

Citoyen René-Amaury Vardes—

De Vardes

Is that, monsieur, the latest Paris mode?

Citoyen René-Amaury Vardes,
The De left off, our hats (Glances at Lalain) left on!

Lalain (removing his hat)

Monsieur
The Baron of Morbec!

De Vardes (bowing)
 Monsieur
The Deputy for Vannes!

[Laughter and voices within.

Enter from the château The Marquise and Mlle. de

Château-Gui with De L’Orient and De Buc.

De L’Orient (sings)

Then spake the king of Ys

Above the song and shout,
Bring here the golden key
That keeps the ocean out!

The Marquise

Monsieur le Baron,
My lost fan!

Yvette (aside)

Oh me!

De Vardes

Madame la Marquise,
I will give you a fan that’s to my taste;
By Watteau painted, mounted by Laudet,
Fragile and fine, an Adonis of fans!
This that I broke I will keep for myself.

[Pockets the fan.

Forgive the mere accident!

Yvette
 Ah!

Séraphine (from the table)

Ah—h—h!

Lalain (aside)

Gods!
If I forgive!

The Marquise

At Blanchefôret, monsieur,
The Watteau, Laudet, Adonis of fans,
I’ll take from your hand—

De Vardes

I ride there anon,

(Aside.) But not through the Forest of Paimpont
And not on the Eve of Saint John.

The Marquise

Come soon,
My garden is sweetest in June.

De L’Orient (sings)

In Ys they sing no more,

In Ys the city old!
The waves are rolling o’er
The king and all his gold.
 Mlle. de Château-Gui

Look at my fan, Monsieur le Baron!

[Lalain crosses to Yvette.

Lalain

Hast thou forgot, hast thou forgot, Yvette,

Thy part, thy lot, the very name they give thee?
This is Morbec, this is the brazen castle!
There are no roses here.

Yvette

So generous
He was!

Lalain

Generous! Oh, well are you called

The Right of the Seigneur!

Yvette (passionately)

Give me not that

Detestable name!

Lalain

So meek under wrongs—

Yvette

Oh!
 Lalain

So quick to forget—

Yvette

Oh!

Lalain

La patrie—
Sworn oaths—the tricolour—

Yvette

Anger me not!

Lalain

On your lips Ça Ira! but in your heart

O Richard, O mon Roi!

Yvette

‘Tis false!

Lalain

And I—and I—Yvette!

Yvette

Speak not to me!

Lalain
You gaze at that man! I tell you he wooes
Madame la Marquise de Blanchefôret!

[Yvette crosses to The Marquise, De Vardes,

and the guests.

Yvette (to The Marquise)

Madame!
I broke the fan! I would pay if I might.
I would keep your cows, or spin your flax—

The Marquise

The fan!
You broke the fan—not monsieur there!

Yvette

No, I!

The Marquise

Sainte Geneviève!

Enter Count Louis, The Vidame, Mme. de Vaucourt,

etc.

Séraphine

Yvette!

Count Louis

La belle Marquise!
 [Séraphine draws Yvette back to the base of the
statue. Count Louis, The Marquise, and the
guests talk together. Lalain crosses to De Vardes.

Lalain

René de Vardes!

De Vardes

Rémond Lalain!

Lalain

This day I bury our friendship of old!

De Vardes

So!

Lalain

I owe to you a thousand louis

Which I’ll repay, monsieur!

De Vardes

I doubt it not.

Lalain

Touch not the girl Yvette!

De Vardes
At last the heart of the matter! I see
You have been through the Forest of Paimpont.

Lalain

Or touch at your peril!

De Vardes

Monsieur!

Lalain

Oh, if
You lay your hand upon your sword, monsieur,
I’m for you there!

De Vardes

Art mad, or drunk with power,

Monsieur the favourite of the Jacobins?

Lalain

There’ll come a day when to be Jacobin

Is something more, monsieur, than to be king!

De Vardes

Indeed!

[A Sergeant of Hussars appears on the terrace and

salutes.

Sergeant!
 The Sergeant

My Colonel!

De Vardes

Well, your report.

The Sergeant

My Colonel, wood and shore we’ve searched since dawn,

And twenty bitter rogues we’ve found, no less!
They crouched behind the tall grey stones, or lay
Prone in the furze, or knelt at Calvaries!
Two women remain—

[He stares at Yvette and Séraphine.

Séraphine

O Saint Thégonnec!
Saint Guirec! Saint Servan!

Yvette

O Our Lady!

Enter The Abbé.

The Abbé

De Vardes, your precious peasants—

[He sees Yvette.

 Who is here?
The De Méricourt, the mænad, I swear!
Who wounded De Vardes!

Yvette

Oh!—

Mme. de Vaucourt

The Egyptian!

Séraphine

Monseigneur, monseigneur, she’s none of mine!

Mlle. de Château-Gui

The poor girl!

Séraphine

Ah, mademoiselle, it is
The innocentest creature!

The Abbé (touches Yvette upon the cheek)

Good-morning,
My dear!

Count Louis

Hm—m—m!—pretty!
 The Vidame

Certainly the gallows

Should be thirty feet high.

Count Louis

Hm—m—m! Something less,

Monsieur le Vidame!

Lalain

Diable!

De Vardes (to the sergeant)

Where are your captives?

The Sergeant

My Colonel,
I have them safely here! Ha! you within!

[Enter from the hall of the château soldiers and

huntsmen with peasants, men and women; some
sullenly submissive, others struggling against their
bonds. They crowd the terrace before the great
doors. The guests of De Vardes to the right and
left upon the terrace, the stairs, and in the garden.
Yvette and Séraphine beside the statue; Lalain
near them; De Vardes with his hand upon the
great chair.

Mme. de Vaucourt

Oh, the brigands!

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