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Developments in Aquaculture and Fisheries Science - 40

SCALLOPS: BIOLOGY, ECOLOGY,

AQUACULTURE, AND FISHERIES
THIRD EDITION
DEVELOPMENTS IN AQUACULTURE AND FISHERIES SCIENCE
The following volumes are still available:
22. FRONTIERS OF SHRIMP RESEARCH
Edited by P. F. DeLoach, W. J. Dougherty and M. A. Davidson 1991 xv+412 pages

24. MODERN METHODS OF AQUACULTURE IN JAPAN

Edited by H. Ikenoue and T. Kafuku 1992 xvi+274 pages

26. PROTOZOAN PARASITES OF FISHES

Edited by J. Lom and I. Dyková 1992 xii+316 pages

28. FRESHWATER FISH CULTURE IN CHINA: PRINCIPLES AND PRACTICE

Edited by J. Mathias and S. Li 1994 xvi+446 pages

29. PRINCIPLES OF SALMONID CULTURE

Edited by W. Pennell and B. A. Barton 1996 xxx+1040 pages

30. STRIPED BASS AND OTHER MORONE CULTURE

Edited by R. M. Harrell 1997 xx+366 pages

31. BIOLOGY OF THE HARD CLAM

Edited by J. N. Kraeuter and M. Castagna 2001 xix+751 pages

32. EDIBLE SEA URCHINS: BIOLOGY AND ECOLOGY

Edited by J. M. Lawrence 2001 xv+419 pages

33. GLOBAL SEAGRASS RESEARCH METHODS

Edited by F. T. Short and R. G. Coles 2001 viii+482 pages

34. BIOLOGY AND CULTURE OF CHANNEL CATFISH

Edited by C. S. Tucker and J. A. Hargreaves 2004 x+676 pages

36. THE KNOWLEDGE BASE FOR FISHERIES MANAGEMENT

Edited by L. Motos and D. Wilson 2006 xxi+476 pages

38. SEA URCHINS

Edited by J. M. Lawrence 2013 xviii+532 pages

39. THE SEA CUCUMBER APOSTICHOPUS JAPONICUS

Edited by H. Yang, J. F. Hamel, and A. Mercier 2015 xxiv+454 pages
Developments in Aquaculture and Fisheries Science - 40

SCALLOPS
Biology, Ecology, Aquaculture,
and Fisheries

THIRD EDITION
Edited by

SANDRA E. SHUMWAY
Department of Marine Sciences, University of Connecticut,
Groton, Connecticut, USA

G. JAY PARSONS
Fisheries and Oceans Canada, Aquaculture, Biotechnology
and Aquatic Animal Health Science Branch, Ottawa, Ontario, Canada

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For Andy Brand and Neil Bourne, scallop afficionados, mentors, and friends.
 Contents

List of Contributors xi Reproductive System 123

Preface xiii Nervous and Sensory Systems 135
Foot Byssal Complex 147
Preface from Second Edition xv
Acknowledgements 150
Prface from First Edition xvii References 150

1. Reconciling Morphological and Molecular 4. Scallop Adductor Muscles: Structure and Function
Approaches in Developing a Phylogeny for the PETER D. CHANTLER
Pectinidae (Mollusca: Bivalvia)
Introduction 161
JEANNE M. SERB
Structure of the Striated Adductor Muscle 162
Introduction 1 Structure of the Smooth Adductor Muscle 171
What Is a Scallop? 2 Myosin 177
Shell Morphology and Its Application in Taxonomic Function of the Striated Adductor 187
Classification Systems 5 Function of the Smooth Adductor 200
The Influence of Molecular Phylogenetics on Systematics 10 Achievements and Coda 206
Where Do We Go From Here? 19 Acknowledgements 207
Summary 24 References 207
Acknowledgements 25
Glossary 25 5. Neurobiology and Behaviour of the Scallop
References 25 DANIEL I. SPEISER AND LON A. WILKENS

2. Biology and Ecology of Scallop Larvae Introduction 219

SIMON M. CRAGG The Mantle Eyes of Scallops 219
Visually Influenced Behaviours in Scallops 231
Introduction 31 The Locomotory System 238
Early Life History Characteristics of Scallops 31 Acknowledgements 248
Larval Development 40 References 248
Comparative Anatomy 57
Functional Morphology 58 6. Reproductive Physiology
Physiology and Effects of Environmental Variables 60
BRUCE J. BARBER AND NORMAN J. BLAKE
Behaviour and Larval Distribution 62
Rearing Methods 70 Introduction 253
Future Directions 72 Gametogenic Cycles 254
Acknowledgements 73 Regulation of Gametogenic Cycles 268
References 73 Energy Metabolism 279
Applications to Aquaculture 287
3. Scallop Structure and Function Summary 290
PETER G. BENINGER AND MARCEL LE PENNEC Acknowledgements 292
References 292
Introduction 85
An Overview of the Scallop Body 85
7. Physiology: Energy Acquisition and Utilisation
The Mantle and Its Derivatives 87
BRUCE A. MACDONALD, V. MONICA BRICELJ, AND
Pallial Organs and Particle Processing 91 SANDRA E. SHUMWAY
Digestive System and Digestion 104
Cardio-vascular System 111 Introduction 301
Excretory System 119 Energy Acquisition 301

vii
viii CONTENTS

Energy Utilisation: Metabolic Expenditure 327 Further Study 512

Energy Utilisation 337 Acknowledgements 513
References 342 References 513
Further Reading 352

12. Swimming in Scallops

8. Nutrition in Pectinids HELGA E. GUDERLEY AND ISABELLE TREMBLAY
SIMONE SÜHNEL AND FRANCISCO J. LAGREZE-SQUELLA
Introduction 535
Introduction 355 Shell Characteristics and Their Influence Upon
Nutrients and Diet Improvement and Supplementation 355 Swimming Capacities 536
Summary 366 Biomechanics of Swimming in Scallops 538
References 366 Ligament Properties 539
Evaluation of Scallop Swimming Performance
9. Scallop Genetics and Genomics in Nature 540
MAUREEN K. KRAUSE AND ELISABETH VON BRAND Laboratory Tests of Escape Response Performance 541
Visual Assessment of Escape Response Performance
Introduction 371 in the Laboratory 542
Cytogenetics 371 Interspecific Differences in Laboratory Escape
Ploidy Manipulation 373 Response Performance 545
Molecular Markers of Genetic Variation 376 Physiology of Scallop Swimming 546
Applications 380 Biotic and Abiotic Influences on Muscle Use
Population Genetics 387 During Escape Responses 549
Scallop Genomics 397 Interspecific Differences in Patterns of Muscle Use 550
Additional Areas of Genetic Research 405 Metabolic Support of Swimming 552
Future Directions and Conclusions 412 Do Changes in Muscle Metabolic Capacities with
References 412 Reproductive Investment Explain Shifts in Escape
Response Behaviour? 556
Interspecific Differences in Muscle Metabolic
10. Diseases and Parasites of Scallops Capacities 557
RODMAN G. GETCHELL, ROXANNA M. SMOLOWITZ, Determinants of Interspecific Differences in
SHARON E. MCGLADDERY, AND SUSAN M. BOWER
Escape Response Strategies: An Indepth Study
Introduction 425 of Five Scallop Species 557
Microbial Diseases 425 References 562
Mycota 439
Protista 441 13. Scallops and Marine Contaminants
Platyhelminthes 448 ISLAY D. MARSDEN AND PETER J. CRANFORD
Polychaetes 452
Crustacea 454 Introduction 567
Gastropods 456 Trace Metals 568
Algae 457 Trace Organics 571
Foraminiferans 458 Nutrient Levels 573
Porifera 458 Offshore Oil and Gas Operations 574
Cnidaria 459 Health Risks 576
Non-Infectious Diseases 459 Scallops as Sentinel Organisms 577
Summary 460 Use of Biomarkers 578
Acknowledgements 460 Future Directions and Conclusions 580
References 460 References 580

11. Scallop Ecology: Distributions and Behaviour 14. Quantifying and Managing the Ecosystem
ANDREW R. BRAND Effects of Scallop Dredge Fisheries
BRYCE D. STEWART AND LEIGH M. HOWARTH
Introduction 469
Geographical Distribution 470 Introduction 585
Local Distribution 482 Types of Scallop Dredge Fishery 586
Scallop Behaviour Aspects Affecting Distribution 496 Effects on Scallop Populations 586
 CONTENTS ix

Effects on Marine Ecosystems 589 Bay Scallop (Argopecten irradians) 758

Effects on Different Seabed Types 594 Calico Scallop, Argopecten gibbus 761
Management 599 Aquaculture and Enhancement 765
Conclusion 603 Future 767
Acknowledgements 604 Acknowledgements 767
References 604 References 767

15. Dynamics, Assessment, and Management 19. The European Scallop Fisheries for Pecten
of Exploited Natural Scallop Populations maximus, Aequipecten opercularis, Chlamys islandica,
J.M. (LOBO) ORENSANZ, ANA M. PARMA, AND STEPHEN J. SMITH and Mimachlamys varia
PETER F. DUNCAN, ANDREW R. BRAND, ØIVIND STRAND, AND
Introduction 611 ERIC FOUCHER
Population Structure and Dynamics 613
Assessment of Abundance and Its Spatial Distribution 632 Introduction 781
The Fishing and Depletion Processes 640 Biology and Ecology 782
Overfishing and Ecological Effects of Fishing 654 Fisheries 801
Management 661 The Future 838
Acknowledgements 676 Acknowledgements 841
Acronyms used in the text 676 References 841
References 677
20. European Aquaculture
16. Scallops of the West Coast of North America ØIVIND STRAND, ANGELES LOURO, AND PETER F. DUNCAN
G. JAY PARSONS, RAYMOND B. LAUZIER, AND NEIL F. BOURNE
Introduction 859
Foreword 697 Historical Overview of European Scallop Aquaculture 859
Introduction 697 Spat Production 862
Fisheries 698 Grow-Out Culture 867
Aquaculture 709 Seabed Ranching and Stock Enhancement 875
Future 713 Biological Constraints to Scallop Culture 879
Acknowledgements 714 Market and Economics 881
References 714 Stock Protection Security and Legal Issues 882
Future Prospects 883
Acknowledgements 884
17. Fisheries Sea Scallop, Placopecten magellanicus References 884
KEVIN D.E. STOKESBURY, CATHERINE E. O’KEEFE, AND
BRADLEY P. HARRIS

Foreword 719
21. Scallop Fisheries and Aquaculture
Introduction 719 in Japan
Life History 721 YOSHINOBU KOSAKA

Ecosystem 723
Introduction 891
The Fishing Fleets of Canada and the United States 724
Patinopecten (Mizuhopecten) yessoensis 891
Stock Assessments 726
Future 926
Fisheries Management 729
Pecten albicans 931
Summary 731
Chlamys (Mimachlamys) nobilis 932
Acknowledgements 732
Acknowledgements 932
References 732
References 933

18. Scallop Aquaculture and Fisheries in Eastern

North America 22. Scallops and Scallop Aquaculture in China
SHAWN M.C. ROBINSON, G. JAY PARSONS, LESLIE-ANNE DAVIDSON, XIMING GUO AND YOUSHENG LUO
SANDRA E. SHUMWAY, AND NORMAN J. BLAKE
Introduction 937
Introduction 737 The Chinese Scallop 938
Sea Scallop (Placopecten magellanicus) 737 Other Scallop Species 943
x CONTENTS

Fishery 945 27. Scallop Aquaculture and Fisheries in Venezuela

Aquaculture 945 CÉSAR J. LODEIROS, LUIS FREITES, JOSÉ J. ALIÓ, MAXIMIANO NÚÑEZ,
AND JOHN H. HIMMELMAN
Harvest, Processing, and Marketing 951
Acknowledgements 951 Introduction 1073
References 951 Distribution, Habitat, and Reproduction 1073
Fisheries 1077
23. Scallops of Northwestern Pacific Russian Aquaculture 1078
Federation Perspectives for Culture 1084
VICTOR V. IVIN, OLGA G. SHEVCHENKO, AND Acknowledgements 1085
TATIANA YU. ORLOVA References 1085

Introduction 953
Biology and Ecology 953 28. Scallop Fishery and Culture in Peru
Fishing and Aquaculture 975 JAIME MENDO, MATTHIAS WOLFF, TANIA MENDO, AND LUIS YSLA

Acknowledgements 991
Introduction 1089
References 991
Species Description 1089
Production Chain and Marketing 1094
24. Aquaculture of the Scallop Fisheries and Aquaculture Production 1095
Nodipecten nodosus in Brazil Challenges and Projections of Peruvian Scallop Production 1104
GUILHERME S. RUPP AND G. JAY PARSONS Acknowledgements 1105
References 1105
Introduction 999
Aquaculture of Nodipecten nodosus in Brazil 1001
Acknowledgements 1015 29. Scallop Fisheries and Aquaculture in Mexico
References 1015 CÉSAR A. RUIZ-VERDUGO, VOLKER KOCH, ESTEBAN FÉLIX-PICO,
ANA ISABEL BELTRAN-LUGO, CARLOS CÁCERES-MARTÍNEZ,
JOSÉ MANUEL MAZON-SUASTEGUI, MIGUEL ROBLES-MUNGARAY,
25. Scallops Biology, Fisheries, and Management AND JORGE CACERES-MARTÍNEZ

in Argentina Introduction 1111

GASPAR SORIA, J.M. (LOBO) ORENSANZ, ENRIQUE M. MORSÁN, Description of Mexican Scallops 1112
ANA M. PARMA, AND RICARDO O. AMOROSO
Fisheries 1114
Introduction 1019 Aquaculture 1119
The Tehuelche Scallop, Aequipecten tehuelchus 1019 References 1123
The Patagonian Scallop, Zygochlamys patagonica 1032
Acknowledgements 1041 30. Scallop Fisheries, Mariculture, and Enhancement
References 1041 in Australasia
MICHAEL C.L. DREDGE, ISLAY D. MARSDEN, AND JAMES R. WILLIAMS
26. Scallop Fishery and Aquaculture in Chile:
Introduction 1127
A History of Developments and Declines
Biology 1129
ELISABETH VON BRAND, ALEJANDRO ABARCA,
GERMÁN E. MERINO, AND WOLFGANG STOTZ Fisheries and Fishery Management 1140
Culture 1161
Introduction 1047 Future Directions and Conclusions 1164
Species Description 1047 Acknowledgements 1166
Fisheries 1050 References 1166
How Scallop Aquaculture Started in Chile 1053
Aquaculture Production 1055
Acknowledgements 1068 General Index 1171
References 1068 Species Index 1189
 List of Contributors
Alejandro Abarca Departamento de Acuicultura,
Universidad Católica del Norte, Coquimbo, Chile
José J. Alió Instituto Nacional de Investigaciones Agrı́colas,
Cumaná, Estado Sucre, Venezuela
Ricardo O. Amoroso School of Aquatic and Fishery
Sciences, University of Washington, Seattle, WA, USA
Bruce J. Barber Eckerd College, St. Petersburg, Florida,
USA
Ana Isabel Beltran-Lugo Departamento de Ingenierı́a en
Pesquerı́as, Universidad Autónoma de Baja California Sur
Ap., La Paz, Baja California Sur, México
Peter G. Beninger Laboratoire de Biologie Marine, Faculté
des Sciences, Université de Nantes, Nantes, France
Norman J. Blake College of Marine Science, University of
South Florida, St. Petersburg, Florida, USA
Neil F. Bourne Fisheries and Oceans Canada, Pacific
Biological Station, Nanaimo, British Columbia, Canada
Susan M. Bower Fisheries and Oceans Canada, Pacific
Biological Station, Nanaimo, British Columbia,
Canada
Andrew R. Brand University of Liverpool, Port Erin, Isle of
Man, United Kingdom
V. Monica Bricelj Department of Marine and Coastal
Sciences and Haskin Shellfish Research Laboratory, School
of Environmental and Biological Sciences, Rutgers
University, Port Norris, NJ, USA
Jorge Caceres-Martı́nez Centro de Investigación Cientı́fica y
Educación Superior de Ensenada (CICESE), Ensenada,
Baja California, México; Instituto de Sanidad Acuı́cola,
Ensenada, Baja California, México Instituto de Sanidad
Acuı́cola, Ensenada, Baja California, México
Carlos Cáceres-Martı́nez Departamento de Ingenierı́a en
Pesquerı́as, Universidad Autónoma de Baja California Sur
Ap., La Paz, Baja California Sur, México
Peter D. Chantler Unit of Veterinary Molecular and Cellular
Biology, Comparative Biomedical Sciences, Royal Veterinary
College, University of London, London, United Kingdom
Simon M. Cragg Institute of Marine Sciences, University of
Portsmouth, Portsmouth, United Kingdom
Peter J. Cranford Fisheries and Oceans Canada, Coastal
Ecosystem Sciences Division, Bedford Institute of
Oceanography, Dartmouth, Nova Scotia, Canada
xi
Leslie-Anne Davidson Fisheries and Oceans Canada,
Moncton, New Brunswick, Canada
Michael C.L. Dredge Tasmania, Australia
Peter F. Duncan Faculty of Science, Health, Education and
Engineering, University of the Sunshine Coast,
Queensland, Australia
Esteban Félix-Pico Centro Interdisciplinario de Ciencias
Marinas, La Paz, Baja California Sur, México
Eric Foucher IFREMER Laboratoire Halieutique de Port
en Bessin, Port en Bessin, France
Luis Freites Instituto Oceanográfico de Venezuela,
Universidad de Oriente, Cumaná, Estado Sucre,
Venezuela
Rodman G. Getchell Department of Microbiology and
Immunology, College of Veterinary Medicine, Cornell
University, Ithaca, NY, USA
Helga E. Guderley Département de Biologie, Université
Laval, Québec City, Québec, Canada
Ximing Guo Haskin Shellfish Research Laboratory,
Department of Marine and Coastal Sciences, Rutgers
University, Port Norris, NJ, USA
Bradley P. Harris Fisheries, Aquatic Science and
Technology Laboratory, Alaska Pacific University,
Anchorage, AK, USA
John H. Himmelman Département de Biologie, Université
Laval, Québec City, Québec, Canada
Leigh M. Howarth School of Ocean Sciences, Bangor
University, Menai Bridge, Anglesey, United Kingdom
Victor V. Ivin Institute of Marine Biology, Far Eastern
Branch of the Russian Academy of Sciences, Vladivostok,
Russia
Volker Koch Deutsche Gesellschaft für Internationale
Zusammenarbeit (GIZ), La Paz, Baja California Sur, México
Yoshinobu Kosaka Aomori Prefectural Industrial
Technology Research Center, Food Research Institute,
Aomori, Japan
Maureen K. Krause Department of Biology, Hofstra
University, Hempstead, NY, USA
Francisco J. Lagreze-Squella Federal University of Paraná,
Pontal do Paraná, Paraná, Brasil
Raymond B. Lauzier Fisheries and Oceans Canada, Pacific
Biological Station, Nanaimo, British Columbia, Canada
xii LIST OF CONTRIBUTORS
Marcel Le Pennec Institut Universitaire Européen de la
Mer, Université de Bretagne Occidentale, Plouzané, France
César J. Lodeiros Instituto Oceanográfico de Venezuela,
Universidad de Oriente, Cumaná, Estado Sucre, Venezuela
Angeles Louro Instituto Español de Oceanografı́a, A
Coruña, Spain
Yousheng Luo Marine Fisheries Research Institute, Dalian,
Liaoning, PR China
Bruce A. MacDonald Biology Department, University of
New Brunswick, Saint John, New Brunswick, Canada
Islay D. Marsden School of Biological Sciences, University
of Canterbury, Christchurch, New Zealand
José Manuel Mazon-Suastegui Centro de Investigaciones
Biologicas del Noroeste, La Paz, Baja California Sur,
México
Sharon E. McGladdery Fisheries and Oceans Canada, St.
Andrews Biological Station, St. Andrews, New Brunswick,
Canada
Jaime Mendo Dpto. Manejo Pesquero y Medio Ambiente,
Facultad de Pesquerı́a, Universidad Nacional Agraria La
Molina, Lima, Peru
Tania Mendo Institute for Marine and Antarctic Studies,
University of Tasmania, Taroona, Tasmania, Australia
Germán E. Merino Departamento de Acuicultura,
Universidad Católica del Norte, Coquimbo, Chile
Enrique M. Morsán Instituto de Biologı́a Marina y
Pesquera ‘Alte. Storni’, Universidad Nacional del
Comahue, San Antonio Oeste, Rı́o Negro, Argentina
Maximiano Núñez Instituto Oceanográfico de Venezuela,
Universidad de Oriente, Cumaná, Estado Sucre,
Venezuela
Catherine E. O’Keefe School for Marine Science and
Technology, University of Massachusetts, Fairhaven, MA,
USA
J.M. (Lobo) Orensanz Centro Nacional Patagónico-
CONICET, Puerto Madryn, Chubut, Argentina
Tatiana Yu. Orlova Institute of Marine Biology, Far Eastern
Branch of the Russian Academy of Sciences, Vladivostok,
Russia
Ana M. Parma Centro Nacional Patagónico-CONICET,
Puerto Madryn, Chubut, Argentina
G. Jay Parsons Fisheries and Oceans Canada, Aquaculture,
Biotechnology and Aquatic Animal Health Science Branch,
Ottawa, Ontario, Canada
Shawn M.C. Robinson Fisheries and Oceans Canada, St.
Andrews, New Brunswick, Canada
Miguel Robles-Mungaray Acuacultura Robles, La Paz,
Baja California Sur, México
César A. Ruiz-Verdugo Departamento de Ingenierı́a en
Pesquerı́as, Universidad Autónoma de Baja California Sur
Ap., La Paz, Baja California Sur, México
Guilherme S. Rupp EPAGRI Empresa de Pesquisa
Agropecuária e de Extensão Rural de Santa Catarina,
Florianópolis, Santa Catarina, Brazil
Jeanne M. Serb Department of Evolution, Ecology
and Organismal Biology, Iowa State University, Ames,
IA, USA
Olga G. Shevchenko Institute of Marine Biology, Far
Eastern Branch of the Russian Academy of Sciences,
Vladivostok, Russia; Research and Educational Center
Primorsky Aquarium, Ostrov Russkii, Vladivostok,
Russia
Sandra E. Shumway Department of Marine Sciences,
University of Connecticut, Groton, Connecticut, USA
Stephen J. Smith Fisheries and Oceans Canada, Bedford
Institute of Oceanography, Dartmouth, Nova Scotia,
Canada
Roxanna M. Smolowitz Aquatic Diagnostic Laboratory,
Feinstein College of Arts and Sciences, Roger Williams
University, Bristol, RI, USA
Gaspar Soria Centro Nacional Patagónico (CONICET),
Puerto Madryn, Chubut, Argentina
Daniel I. Speiser Department of Biological Sciences,
University of South Carolina, Columbia, SC, USA
Bryce D. Stewart Environment Department, University of
York, Heslington, North Yorkshire, United Kingdom
Kevin D.E. Stokesbury School for Marine Science
and Technology, University of Massachusetts, Fairhaven,
MA, USA
Wolfgang Stotz Departamento de Biologı́a Marina,
Universidad Católica del Norte, Coquimbo, Chile
Øivind Strand Institute of Marine Research, Bergen,
Norway
Simone Sühnel Federal University of Santa Catarina,
Florianópolis, Santa Catarina, Brasil.
Isabelle Tremblay Département de Biologie, Université
Laval, Pavillon Alexandre-Vachon, Québec City, Québec,
Canada; Ressources Aquatiques Québec, Institut des
sciences de la mer de Rimouski, Université du Québec à
Rimouski, Rimouski, Québec, Canada
Elisabeth von Brand Departamento de Biologı́a Marina,
Facultad de Ciencias del Mar, Universidad Católica del
Norte, Coquimbo, Chile
Lon A. Wilkens Center for Neurodynamics and
Department of Biology, University of Missouri-St. Louis,
St. Louis, MI, USA
James R. Williams National Institute of Water and
Atmosphere, Viaduct Harbour, Auckland, New Zealand
Matthias Wolff Theoretical Ecology and Modelling, Leibniz
Center for Tropical Marine Ecology, Bremen, Germany
Luis Ysla Dpto. Manejo Pesquero y Medio Ambiente,
Facultad de Pesquerı́a, Universidad Nacional Agraria La
Molina, Lima, Peru
 Preface

From Sandro Botticelli’s The Birth of Venus to the corporate symbol for Shell Oil, scallops are one of the most
widely recognised shellfish worldwide. They are of global economic importance and support both commercial
fisheries and mariculture efforts. They have been the subject of numerous research efforts and their high eco-
nomic stature encourages aquaculture efforts by academicians and industrial researchers. The rapid growth, early
maturity and high economic value of many scallop species offer special inducement to mariculturists. Scallops
are also key in the structure and function of the communities of which they are characteristic components.
Interest in all aspects of scallop biology, aquaculture, ecology and fisheries continues; however, it is the field of
genetics that has seen the greatest surge in research efforts over the past decade.
The First Edition, published in 1991, represented the first comprehensive treatise on scallops. It was prepared
as camera-ready copy using the very earliest of ‘desk-top’ publishing programmes and only through the partici-
pation of Scott Siddall was publication made possible. Technology has thankfully progressed and word-proces-
sing has made projects such as this more tenable but still not without frustrations and roadblocks.
The original version was followed 15 years later with the Second Edition in 2006. Since that printing, five
International Pectinid Workshops have been held: Halifax, Nova Scotia, Canada; Santiago de Compostela, Spain;
Quingdao, China; Florianopolis, Brazil; and most recently, Galway, Ireland. These meetings provided a unique
and focused forum for discussion of ‘all things scallop’ and we believe that both directly and indirectly the con-
tributions presented here have benefitted greatly from those open and often lively discussions.
Many of the authors from the First and Second Editions have since retired or taken on other responsibilities
and readers will note a number of new contributors. Some have joined forces with prior authors, while other con-
tributors have penned new submissions.
The present volume presents a comprehensive overview of the biology, ecology, aquaculture, and fisheries
prospects, both present and future, for scallops worldwide. The inclusion of fisheries in the title is a testament to
increased research and the importance of the commercial fleets and habitats. Some chapters have been completely
rewritten, some have been revised, and new chapters have been added. While the subject matter is specifically
concerned with scallop species, the book should be of interest and relevance to a number of readers including
advanced undergraduate/graduate students, mariculturists, researchers and fisheries managers, and scallop
enthusiasts. Chapters have been prepared by some of the foremost authorities in their respective fields and repre-
sent 24 countries. Manuscripts submitted by authors whose first language is not English have been corrected for
gross grammatical errors and clarity of presentation only. Their efforts to write a manuscript in a foreign lan-
guage are to be applauded and should not be clouded by editorial license. Scallops are of more economic impor-
tance in some countries than in others and this is reflected in the specific coverage of the individual species.
The book is comprised of 30 chapters: the first 15 spanning topics from phylogeny, biology of larvae, structure
and function, genetics, physiology, nutrition, swimming, ecology, populations dynamics, etc. And the second
half of the book is a comprehensive country or regional overview of the fishery and aquaculture of scallops from
all the key areas of production.
We have dedicated this edition to our friends and colleagues, Andrew (Andy) Brand and Neil Bourne. From
his home base on the Isle of Man, Andy devoted his career to mentoring students, and bringing top-tier science
to fishermen, government managers, aquaculturists, and most recently to the sustainability certification efforts of
the Marine Stewardship Council. Andy has always been generous with this knowledge and time, and unofficially
recognised as the senior statesman of the International Pectinid Workshops. Thank you, Andy, for all you have
contributed. And from his scenic base in Nanaimo, British Columbia, Canada Neil, too, has spent an illustrious
career as a shellfish researcher working with managers, scientists, students, and fishing and aquaculture industry
members to discover and communicate a vast array of knowledge of scallops and other bivalve species. In addi-
tion to being a mentor, teacher and researcher par excellence, Neil was instrumental in laying the foundation for

xiii
xiv PREFACE

the scallop aquaculture industry in British Columbia. Neil, thank you for your friendship, insights and
contributions.
The current edition has benefitted greatly from the efforts of Noreen Blaschik who has tirelessly helped with
proofreading, reference cross-checking and other thankless tasks. The talents of Eric Heupel were key in redraft-
ing many of the figures and are gratefully acknowledged. Getting authors to agree to participate in ventures such
as this is much easier than extracting the final texts and we extend our deepest gratitude to the authors for their
skill, patience and perseverance. Thanks to Sharon, Michael and Christopher for their support, cajoling and
patience. Gus and Zeus spent many evenings pawing over texts, and their assistance made the tasks at hand
more tolerable.
A special thanks to Patricia Osborne, Elsevier Science Publishers who initiated and encouraged the effort for a
Third Edition and to Jaclyn Truesdell, Debbie Clark and Karen Miller for their patience and guidance during the
production.
For successful culture and management, a good knowledge of the biology, ecology, physiology and fisheries of
the species is necessary and we hope that this Third Edition of Scallops: Biology, Ecology, Aquaculture, and
Fisheries will serve as a solid reference base for years to come.

Sandra E. Shumway1 and G. Jay Parsons2

1
Department of Marine Sciences, University of Connecticut, Groton, CT, USA
2
Fisheries and Oceans Canada, Aquaculture, Biotechnology and Aquatic Animal Health Science Branch,
Ottawa, ON, Canada
 Preface from Second Edition

From their continued appearance in art and culture to their savoured place at the dinner table, scallops have
not lost their general appeal and continue to garner the attention of scientists, aquaculturists, chefs, shell collec-
tors and others. Discussion during the Sixth International Pectinid Workshop in 1987 provided the impetus for
the First Edition of Scallops: Biology, Ecology and Aquaculture which was published in 1991 and has been out of
print for nearly a decade. Talk of a Second Edition began almost immediately, but was gently ignored for some
years because of the time and effort required of all contributors and a sense on the part of the original editor that
there was not enough new information to warrant a second volume.
Interest in all aspects of scallop biology and ecology has continued to increase, culture efforts have expanded
globally and scallops continue to bring high prices from both the commercial fisheries and aquaculture ventures.
Potential authors began to enquire enthusiastically about the potential for a Second Edition, Elsevier Science
Publishers were eager to add the volume to their series and there was a renewed sense of need coupled with
availability of new information to proceed with publishing a Second Edition.
The present volume represents an updated and revised version of the First Edition. It is a comprehensive over-
view of the biology, ecology and aquaculture for scallop species worldwide. Some chapters have been completely
rewritten by previous or new authors and two chapters have been reprinted as in the original volume
(see Chapter 5 Neurobiology and Behaviour of the Scallop by Wilkens and Chapter 8 Physiological
Integrations and Energy Partitioning by Thompson and MacDonald) as they are as pertinent now as they were in
1991. Other chapters have been revised and new ones added (Nutrition and accounts by country of aquaculture
and fisheries Brazil, Venezuela and the Russian Federation now that the former Soviet Union has transformed
since the First Edition!). This work would not have been possible without the assistance of several individuals.
We thank Linda Kallansrude, Manon Chouinard and Dinah Helpert who provided secretarial and administrative
support, Maille Lyons who took on the arduous task of developing the indices, Mara Vos-Sarraiento (Elsevier)
for assistance, guidance and support, and a special thanks to Sharon, Christopher, Michael and Max for their
support and patience.
Finally, the authors are to be congratulated for their efforts and thanked profusely for their patience during
the publication process.
We have dedicated this volume to Sam Naidu, friend and colleague, who devoted his career to working with
fishermen, aquaculturists and government managers to ensure a sustainable fishery for future generations. Sam
was a gentleman, generous with his knowledge and expertise, accepted no social barriers and was a pillar of the
Pectinid Workshops. Moreover, he had perfected the art of living well and appreciating life. Since publication of
the First Edition, there have been 7 subsequent International Pectinid Workshops, the 15th held in Mooloolaba,
Australia and the 16th held in Halifax, Nova Scotia, Canada in 2007.
We hope this volume will continue to foster interest in scallops, especially aquaculture for a sustainable future,
for years to come.

Sandra E. Shumway and G. Jay Parsons

Groton, CT, USA
Ottawa, ON, Canada

xv
 Preface from First Edition

Scallops are among the better known shellfishes and are widely distributed throughout the world. They are of
worldwide economic importance and support both commercial fisheries and mariculture efforts. They have been
the subject of numerous research efforts and their high economic stature encourages aquaculture efforts by both
academicians and industrial researchers. The rapid growth, early maturity and high economic value of many
scallop species offer special inducement to mariculturists. Scallops are already cultured successfully in a number
of geographic locations and emphasis is increasing globally. Further, scallops play an important role in the struc-
ture and function of the communities of which they are a characteristic and important component.
Not since the publication of Marine Mussels (Bayne, 1975: IBP Handbook) has the current knowledge of a par-
ticular group of molluscs received a comprehensive coverage. His effort was a welcome addition to the scientific
literature and provided an opportunity to synthesise a wealth of information. For successful culture and manage-
ment, a good knowledge of the biology of the species is necessary. While scallops have been the subject of a con-
siderable amount of research, to date no integrated account exists of the biology.
Discussions regarding the need for a comprehensive treatise on scallops first took place in 1987 during the
Sixth International Pectinid Workshop in Menai Bridge, Wales. The level of enthusiasm was high and convincing
the individual authors to undertake writing the chapters was an easy task. The subsequent task of collecting and
integrating those manuscripts proved to be considerably more arduous. There were further discussions regarding
the final versions of manuscripts during the Seventh International Pectinid Workshop held in Portland, Maine, in
1989. These meetings provided a unique opportunity for specialists in the field of scallop biology representing
over 20 countries to meet and discuss their individual views and I believe that the final product has benefited
greatly from those open and often lively discussions.
The present volume presents a comprehensive overview of the biology, ecology, and aquaculture prospects,
both present and future, for scallops worldwide. Although the subject matter is specifically concerned with the
biology, ecology and aquaculture of scallop species, the book should be of interest and relevance to a number of
readers including advanced undergraduate/graduate students, mariculturists and research workers. Contributed
chapters have been prepared by some of the foremost authorities in their respective fields and represent some 18
countries. Manuscripts submitted by authors whose first language is not English have been corrected for gross
grammatical errors and clarity of presentation only. Their efforts to write a manuscript in a foreign language are
to be applauded and should not be clouded by editorial licence.
The book is divided into two main sections: the first dealing with general biology, ecology, and physiology
and the second dealing with fisheries and culture efforts in specific countries. Obviously, scallops are of more
economic importance in some countries than in others and this is reflected in the specific coverage of specific spe-
cies in Chapter 12.
The phylogenetic relationship of scallop species of commercial interest is examined by Waller in Chapter 1.
The available information on larval requirements and ranges of tolerance of environmental variables, both prime
importance to aquaculture efforts, are discussed by Cragg and Crisp in Chapter 2. Beninger and LePennec pro-
vide an elegant account of the internal anatomy of scallops in Chapter 3. The importance of scallop adductor
muscles to the field of muscle physiology is clearly defined by Chantler in Chapter 4. Energy acquisition and uti-
lisation are discussed by Bricelj and Shumway in Chapter 5 and the physiological integrations and partitioning of
this energy are discussed in Chapter 6 by Thompson and MacDonald. Reproductive biology, another area of
prime importance to culturists, is reviewed by Barber and Blake in Chapter 7. Two of the most striking features
of scallops, eyes and their ability to swim, are discussed by Wilkens in Chapter 8. Diseases and parasites, major
threats to aquaculture ventures, are discussed by Getchell in Chapter 9, and Gould and Fowler review the effects
of pollutants on the commercial species of scallops in Chapter 10. General ecology, distribution and behaviour
are given in Chapter 11 by Brand. Beaumont and Zouros review the application of genetic tools to scallops and

xvii
xviii PREFACE FROM FIRST EDITION

the importance of genetics to fisheries and aquaculture of scallops in Chapter 12. In Chapter 13, Orensanz, Parma
and Iribane discuss population dynamics and the management of natural stocks. The commercial species of scal-
lops are covered in Chapter 14 according to geographic location. Much of the data presented in Chapter 14 is
being reported for the first time, including such elementary measures as growth, meat weight and landings.
This work would not have been possible without the efforts of Dr. Scott Siddall. His expertise in the field of
desktop publishing coupled with his knowledge of shellfish biology made my task possible. His continued inter-
est, support and interjections of humour when most needed made the task enjoyable. Jan Barter, Martha
Hernandez-Davis, John Hedley and Jim Rollins helped in various ways and their willingness to undertake such
arduous tasks as proofreading, crosschecking references, retyping of manuscripts and redrawing and repair of
submitted diagrams are gratefully acknowledged. A special thanks must go to Bob Goodman, Elsevier Science
Publishing Company, for his considerable patience in ‘meeting’ deadlines and his insight and support. Finally,
I would like to thank the authors for their skill, patience, and perseverance.

Sandra E. Shumway
Boothbay Harbor, ME, USA
 C H A P T E R

1
Reconciling Morphological and Molecular
Approaches in Developing a Phylogeny for the
Pectinidae (Mollusca: Bivalvia)
Jeanne M. Serb

INTRODUCTION

Scallops (Pectinidae Rafinesque, 1815) are one of the most morphologically, behaviourally, and biologically
diverse family of bivalves. As a major molluscan clade of Mesozoic and Cenozoic eras, the Pectinidae has had a
tumultuous 245-million-year history, surviving two mass extinction events at the Triassic-Jurassic and
Cretaceous-Paleogene boundaries (Hautmann, 2010; Newell and Boyd, 1995; Waller, 2006a). Over the intervening
quarter of a billion years, it is likely that scallops underwent several adaptive radiations that played a major role
in generating the biological diversity seen today. Currently, about 270 extant species of scallops are recognised
(Dijkstra, 2014) and are distributed across polar, temperate, and tropical seas, with the greatest biological diver-
sity in the tropical Caribbean and Indo-Pacific oceans (Raines and Poppe, 2006). Most scallop species are found
in the shallows of sublittoral reefs, sandy bays, and sea grass beds of the continental shelves, with a smaller num-
ber of species restricted to deeper water including the abyssal depths at 7000 m. While the typical scallop shell
shape is the familiar rounded disk and straight hinge line formed by two ear-like auricles, scallops exhibit a wide
range of modifications on this generalised phenotype, with striking shape, texture, and colour variants of the
shell. This variety has made scallops a favourite of amateur shell collectors and scientists alike and, as a result,
their shells are well represented in the collections of natural history museums (e.g. Dijkstra, 1999; Dijkstra and
Koehler, 2008). The on-going challenge for scallop biologists is to organise the impressive biological diversity of
the Pectinidae in a phylogenetic context.
Under a phylogenetic classification system, taxa are organised hierarchically into groups that reflect a hypothe-
sis of evolutionary relationship derived from a phylogenetic tree. A phylogenetic tree (also called a ‘phylogeny’
or ‘topology’) is a bifurcating diagram that depicts the relationships between taxa based on shared history
through common descent (Gregory, 2008). Not only can the phylogeny help identify groups of taxa (clades), but
phylogenetic trees indicate relationships between these groups, for example, how tribes are related to one another
in a subfamily. Phylogenetic trees also provide a historical context and a relative order for when lineages arose
and diversified. Moreover, the phylogeny can be used to study the evolution of traits, such as whether a charac-
ter state is ancestral (plesiomorphic) or derived (i.e. trait polarity), or to test hypotheses regarding the rate or
mode of trait evolution.
Basing a classification system on phylogeny incorporates the evolutionary history of those taxa and all the
associated information into the classification. Species or lineages that are descendants of a single ancestor repre-
sent ‘natural groups’, and the recognition of natural groups gives predictive power to a classification system
because the shared characters may also indicate similarities in life history traits, habitat requirements, or ecologi-
cal function due to shared history of the organisms. Thus, modern classification systems are information retrieval
resources, where an individual species can be used to represent a broader set of species in some contexts, such as

Scallops: Biology, Ecology, Aquaculture, and Fisheries.

S.E. Shumway and G.J. Parsons (Editors). 1 © 2016 Elsevier B.V. All rights reserved.
2 1. RECONCILING MORPHOLOGICAL AND MOLECULAR APPROACHES IN DEVELOPING A PHYLOGENY FOR THE PECTINIDAE (MOLLUSCA: BIVALVIA)

the ‘surrogate approach’ for conservation management (Wiens et al., 2008) or as an organismal model for a
research programme (Jenner and Wills, 2007). For the Pectinidae, a better understanding of the phylogeny of scal-
lops could guide aquaculturalists when targeting which species would be best suited for culturing under particu-
lar conditions. In addition, a common nomenclature for scallops will facilitate communication among workers
from different scientific disciplines or countries.
Furthermore, the Pectinidae has great potential to be a model system in evolutionary biology. Scallops repre-
sent a large and diverse group of animals with a superb fossil record, and so can tell us much about evolutionary
processes like speciation, adaptation, and extinction. Already the Pectinidae has demonstrated utility in the study
of morphology (Serb et al., 2011; Sherratt et al., n.d.), behaviour (Alejandrino et al., 2011), sensory biology (Pairett
and Serb, 2013; Serb et al., 2013), genome evolution (Wang and Guo, 2004), and extinction risk (Smith and Roy,
2006; Harnik, 2011). But in order to understand the processes underlying these trends, we need an accurate phy-
logeny and, preferably, work towards an inclusive phylogeny that accounts for both living and extinct species.
With this phylogeny in hand, researchers can not only organise biological diversity into a classification scheme
that reflects relatedness but also study the processes that generated this biological diversity.
This review examines the conflicts between morphological-based and molecular phylogenetic hypotheses of
the Pectinidae. The first section describes the characteristics of scallops by defining the traits unique to the family
and provides background on the ecological and phenotypic diversity of the group. Through a review of previous
work, the second section explains how this morphological diversity was used to develop a classification system
for the Pectinidae and shows how these systems are affected by convergent evolution of the shell. The third sec-
tion describes how molecular phylogenetics can add to our understanding of scallop relationships and evolution,
reviews the last decade of phylogenetic study on the Pectinidae, and provides a case study. The chapter ends
with some thoughts about what still needs to be done to address long-standing issues in scallop phylogenetics
including determining the evolutionary origin of the Pectinidae, how to better estimate biological diversity in the
family, and methods to incorporate extant and extinct taxa in a comprehensive phylogenetic hypothesis for the
scallops.

WHAT IS A SCALLOP?

Defining the Pectinidae

Scallops belong to the Pectinoidea Rafinesque, 1815, a superfamily that includes the Pectinidae; Entoliidae
Teppner, 1922; Propeamussiidae Abbott, 1954; Spondylidae Gray, 1826; and the extinct Entolioidesidae Kasum-
Zade, 2003; Neitheidae Sobetski, 1960; Pernopectinidae Newell, 1938; Tosapectinidae (Bieler et al., 2010). All
members of Pectinoidea are united by having a triangular resilium with a non-mineralised medial core that func-
tions below the hinge line (Waller, 1978, 2006b). In early ontogeny, all species exhibit a pectiniform growth stage
complete with a byssal notch, and presumably these species can attach to substrates using a byssus (Waller,
2006b). After the Pectinidae, the second largest extant family of the Pectinoidea is the Propeamussidae or ‘glass
scallops’ (B200 species). Shells of these small, deep-water bivalves are thin, colourless, and resemble some pecti-
nid species. The family Spondylidae or ‘thorny oysters’ (B70 species) is distributed throughout the tropical and
subtropical oceans and all members cement to substrates. The family with the fewest extant species is the
Entoliidae, or ‘smooth pectens’ (two species). As amended by Waller (2006b), this family contains two extant spe-
cies. Other morphological differences among the pectinoidean families include the hinge dentition and the min-
eral composition of the valves (see Waller, 2006b).
The Pectinidae is the largest pectinoidean family and its members can be identified by two key structures: (i) a
particular hinge-ligament system that connects the two valves of the shell and (ii) the presence of a ctenolium
(Waller, 1984; Hautmann, 2010). The hinge-ligament system of the scallop is the alivincular-alate ligament condition
(as described in Hautmann, 2004), where a triangular-shaped, fibrous ligament (the resilium) is secreted in a depres-
sion (the resilifer) on internal surface of a long, straight hinge margin of the dorsal portion of the valve. A second
ligament extends laterally along the outer length of the straight hinge margin, in a wing-like or ‘alate’ manner.
The lateral ligament aligns and connects the left and right valves of the shell together, while compression of
the uncalcified centre of the resilium performs like a spring, acting in opposition to the action of the adductor mus-
cles that draw both valves together. Thus, it has been hypothesised that the elastic hinge was key to development of
shell clapping and swimming behaviours exhibited by scallops (Trueman, 1953; Marsh and Olson, 1994; Dickinson
et al., 2000). All scallops, and a few extinct families (e.g. Pernopectinidae and Entiolioididae; Korobkov, 1960),

SCALLOPS
 WHAT IS A SCALLOP? 3
possess the alivincular-alate ligament. In contrast, the ctenolium is unique to the Pectinidae and is present at least in
the early post-larval (dissoconch) growth stages. The ctenolium is a row of hook-shaped denticles along the ventral
edge of byssal notch (Waller, 1978, 1984) (Figure 1.1). As a protein byssus is secreted by the byssal gland within the
muscular foot, it passes over the denticles of the ctenolium which keeps the array of threads flat along the edge of
the shell (Waller, 1984: Figure 4b, c). The combination of a byssal thread sheet and the two-point contact of shell to a
surface (see Stanley, 1972) stabilises the animal’s connection to the substrate and greatly reduces how much the shell
can rotate around the byssus. As a result, much more torque must be applied to break the byssal attachment (Waller,
1984) and this may provide an additional level of protection from predators that must dislodge and manipulate prey
items before consumption (Caro et al., 2008).

The Relationship Between Life Habit and Shell Shape

Scallops metamorphose from a swimming pediveliger to a crawling, benthic post-larva. At this time, the man-
tle begins secretion of the dissoconch shell (Hodgson and Burke, 1988), whose microsculpture patterns have been
used as characters in some classification systems (Waller, 1991). All newly metamorphosed scallops possess a cte-
nolium and can produce a byssus to attach to a substrate. This attachment is temporary as the animal can release
the byssus and then secrete a new byssus to attach to another substrate. As the animal grows and matures, the
ctenolium may be overgrown by the advancing front of the anterior auricle (Waller, 1984: Figures 2a c, 3b). This
results in the progressive loss of the ability to attach using a byssus as body size increases (Caddy, 1972). Thus,
some scallop species only employ a byssus as juveniles and lose the ability to produce a byssus as adults, becom-
ing ‘free-living’.
Based on the ability to attach and mode of locomotion, adult scallops can be placed into six functional groups
or life habits (sensu Stanley, 1970). These life habits are associated with specific shell forms of scallops (Stanley,
1970). The majority of scallop species have the byssal attaching life habit, where ability to attach temporarily to a
substrate via byssus is maintained after maturity. Byssal attachers have a chlamydoid form, where the shell has a
strong asymmetry along the anterior posterior axis such that the auricles are of different sizes and the shell
outline is oblique (Figure 1.2A). Additionally, the right valve has a deep byssal notch with a well-developed
ctenolium (Figure 1.1). It has been hypothesised that the chlamydoid form is the plesiomorphic condition for the
Pectinidae (Waller, 1993). A few byssal attaching species, like Pedum spondyloidium (Gmelin, 1791), exhibit the
nestling life habit. These species are obligate associates with living scleractinian corals, where the post-larva pref-
erentially settles on, and byssally attaches to, coral polyps which grow around the scallop encasing it (Yonge,
1967; Waller, 1972a).
Other species, such as Crassadoma gigantea (Gray, 1825), permanently attach to hard, often abiotic, substrates
by secreting new shell material. This cementing life habit is associated with a shell that transitions from the

Post. auricle Umbo

Ant. auricle

Byssal
Ctenolium
notch

Right valve

FIGURE 1.1 Some morphological features of the pectinid shell. Right valve of Volachlamys tranquebaria oriented with anterior (ant.) towards
right and posterior (post.) towards left.

SCALLOPS
Another random document with
no related content on Scribd:
 Table of the Twelve Houses.
Astrologers draw their table of the TWELVE HOUSES into a triple
quadrangle prepared for the purpose, of which there are four
principal angles, two of them falling equally upon the horizon, and
the other two upon the meridian, which angles are sudivided into 12
triangles for the 12 houses, in which they place the 12 signs of the
Zodiac, to each of which is attributed a particular quality,—viz.

1.— Aries, denoted by the sign ♈︎, is, in their extravagant opinion,
a masculine, diurnal, cardinal, equinoctial, easterly sign, hot
and dry,—the day house of Mars.
2.— Taurus, ♉︎, is a feminine, nocturnal, melancholy, bestial,
furious sign—cold and dry.
3.— Gemini, ♊︎, is a masculine sign, hot and moist, diurnal, aërial,
human, double-bodied, &c.
4.— Cancer, ♋︎, is a feminine, nocturnal, phlegmatic sign, by
nature cold and moist, the only house of Luna.
5.— Leo, ♌︎, is a sign, masculine, diurnal, bestial, choleric and
barren; a commanding, kingly sign—hot and dry, the only
house of the sun.
6.— Virgo, ♍︎, is a feminine, nocturnal, melancholy, and barren
sign.
7.— Libra, ♎︎, is a sign masculine, cardinal, equinoctial, diurnal,
sanguine and human, hot and moist.
8.— Scorpio, ♏︎, is a feminine, nocturnal, cold and phlegmatic
northern sign.
9.— Sagittarius, ♐︎, is a sign masculine, choleric, and diurnal, by
nature hot and dry.
 10. Capricorn, ♑︎, is a feminine, nocturnal, melancholy, solstitial,
— moveable, cardinal, and southern sign.
11.— Aquarius, ♒︎, is a masculine, diurnal, fixed, sanguine, and
human sign.
12. Pisces, ♓︎, is a feminine, nocturnal, phlegmatic, northerly
— double-bodied sign, the last of the twelve.
Having thus housed their signs and directed them in their
operations, they afterwards come to enquire of their tenants, what
planet and fixed stars they have for LODGERS, at the moment of the
nativity of such person; from whence they draw conclusions with
regard to the future incident of that person’s life. For if at the time of
that person’s nativity they find Mercury in 27° 52 min. of Aquarius,
and in the sextile aspect of the horoscope, they pretend to foretel that
that infant will be a person of great sagacity, genius, and
understanding; and therefore capable of learning the most sublime
sciences.
Astrologers have also imagined, for the same ridiculous purpose,
to be in the same houses different positions of the signs and planets,
and from their different aspects, opposition and conjunction, and
according to the rules and axioms they have prescribed to themselves
and invented, have the sacrilegious presumption to judge, in dernier
resort, of the fate of mankind, though their pretended art or science
is quite barren either of proofs or demonstrations.
 Signs to the Houses of the Planets.
The planets have allowed themselves each, except Sol and Luna,
two signs for their houses; to Saturn, Capricorn and Aquarius; to
Jupiter, Sagittarius and Pisces; to Mars, Aries and Scorpio; to Sol,
Leo; to Venus, Taurus and Libra; to Mercury, Gemini and Virgo;
and to Luna, Cancer.
 Angles or Aspects of the Planets.
By their continual mutations among the twelve signs, the planets
make several angles or aspects; the most remarkable of which are the
five following, viz.—
☌ Conjunction.—Δ Trine.—☐ Quadrate.—⚹ Sextile.—☍
Opposition.
A Conjunction is when two planets are in one and the same
degree and minute of a sign; and this, according to Astrological cant,
either good or bad, as the planets are either friends or enemies.
A Trine is when two planets are four signs, or 120 degrees distant,
as Mars in twelve degrees of Aries, and Sol in twelve degrees of
Leo. Here Sol and Mars are said to be in Trine Aspect. And this is
an aspect of perfect love and friendship.
A Quadrate Aspect is when two planets are three signs, or 90
degrees distant, as Mars in 10 degrees, and Venus in 10 degrees of
Leo. This particular aspect is of imperfect enmity, and Astrologers
say, that persons thereby signified, may have jars at sometime, but of
such a nature as may be perfectly reconciled.
A Sextile Aspect, is when two planets are two signs, or 60 degrees
distant, as Jupiter in 15 degrees of Aries; and Saturn in 15 degrees
of Gemini; here Jupiter is in a sextile aspect to Saturn. This is an
aspect of friendship.
An Opposition is, when two planets are diametrically opposite,
which happens when they are 6 signs, or 180 degrees (which is one
half of the circle) asunder; and this is an aspect of perfect hatred.
A Partile Aspect, is when two planets are in a perfect aspect to
the very same degree and minute.
Dexter Aspects, are those which are contrary to the succession of
signs; as a planet, for instance, in Aries, casts its sextile dexter to
Aquarius.
Sinister Aspect, is with the succession of signs, as a planet in
Aries, for example, casts its sextile sinister in Gemini.
In addition to these, Astrologers play a number of other diverting
tricks; hence we read of the Application—Prohibition—
Translation—Refrenation—Combustion—Exception—
Retrogradation, &c. of planets.
 The Application of Planets.
Application of the planets is performed by Astrologers in three
different ways.
1. When a light planet, direct and swift in its motion, applies to a
planet more ponderous and slow in motion; as Mercury in 8° of
Aries, and Jupiter in 12° of Gemini, and both direct; here Mercury
applies to a sextile of Jupiter, by direct application.
2. When they are both retrograde, as Mercury in 20° of Aries, and
Jupiter in 15° of Gemini; here Mercury, the lighter planet, applies to
the sextile aspect of Jupiter; and this is by retrogradation.
3. When one of the planets is direct, and the other retrograde; for
example, if Mercury were retrograde in 18° of Aries, and Jupiter
direct in 14° of Gemini; in this case Mercury applies to a sextile of
Jupiter, by a retrograde motion.
 Prohibition,
is when two planets are applying either by body or aspect; and
before they come to their partile aspect, another planet meets with
the aspect of the former and prohibits it.
 Separation,
is when two planets have been lately in conjunction, or aspect, and
are separated from it.
 Translation of Light and Virtue,
is when a lighter planet separates from the body or aspect of a
heavier one, and immediately applies to another superior planet, and
so translates the light and virtue of the first planet to that which it
applies to.
 Refrenation,
is when a planet is applied to the body or aspect of another; and,
before it comes to it, falls retrograde, and so refrains by its
retrograde motion.
 Combustion.
A planet is said to be combust of Sol, when it is within 8° 30″ of
his body, either before or after his conjunction: but Astrologers
complain, that a planet is more afflicted when it is applying to the
body of Sol, than when it is separating from combustion.
 Reception,
is when two planets are in each other’s dignities, and it may either
be by house, exultation, triplicity, or term.
 Retrogradation,
is when a planet moves backward from 20° to 9°, 8°, 7°, and so out
of Taurus into Aries.
 Frustration,
is when a swift planet applies to the body or aspect of a superior
planet; and before it comes to it, the superior planet meets with the
body or aspect of some other planet.
 The Dragon’s Head and Tail.
To the seven planets, viz. Saturn, Jupiter, Mars, Sol, Venus,
Mercury, and Luna; Astrologers add, two certain nodes or points,
called the Dragon’s head, distinguished by this sign ☋, and the
Dragon’s tail by ☊. In those two extremities of the beast, our students
in Astrology place such virtues, that they can draw from thence
wealth, honour, preferments, &c. enough to flatter the avarice,
ambition, vanity, &c. of the fools who follow them. Sensible,
however, that the admirers of this art support their principles and
defend their doctrines by examples founded on their own experience
and on the authority of history; there is no necessity for us here to
expose the weakness and futility of their arguments. Tully’s proof
will suffice; who, amidst the darkest clouds of superstition and
ignorance, and in the very heyday of paganism and idolatry, and
whilst religion itself seemed to countenance Astrology, inveighs
severely against it in Lib. 2, de devinat. “Quam multa ego Pompeis,
quam multa Crasso, quam multa huic ipsi Cæsari a Chaldæis dicta
memini, neminem eorum nisi senectute, nisi domi, nisi cum clantate
esse moriturum? ut mihi per Mirum videatur quem quam extare,
qui etiam nunc credastis, quorum predicta quotidie videat re et
eventis refelli[9].”
 Climacteric.
Astrologers have used their best artifices, and employed all the
rules of their art, to render those years of our age, which they call
climacterics, dangerous and formidable.
Climacterick from the Greek, κλιμακτης, which means by a scale or
ladder, is a critical year, or a period in a man’s age, wherein,
according to Astrological juggling, there is some notable alteration to
arise in the body; and a person stands in great danger of death. The
first climacterick, say they, is the seventh year of a man’s life; the rest
are multiples of the first, as 21, 49, 56, 63, and 84; which two last are
called the grand climactericks, and the danger more certain.
Marc Ficinus accounts for the foundation of this opinion: he tells
us there is a year assigned for each planet to rule over the body of a
man, each in his turn; now Saturn being the most maleficent
(malignant) planet of all, every seventh year, which falls to its lot,
becomes very dangerous; especially those of 63 and 84, when the
person is already advanced in years. According to this doctrine, some
hold every seventh year an established climacteric; but others only
allow the title to those produced by the multiplication of the
climacterical space by an odd number, 3, 5, 7, 9, &c. Others observe
every ninth year as a climacterick.
There is a work extant, though rather scarce, by Hevelius, under
the title of Annus Climactericus, wherein he describes the loss he
sustained by his observatory, &c. being burnt; which, it would
appear, happened in his grand climacterick. Suetonius says, that
Augustus congratulated his nephew upon his having passed his first
grand climacterick, of which he was very apprehensive.
Some pretend that the climacterick years are fatal to political
bodies, which perhaps may be granted, when they are proved to be so
to natural ones; for it must be obvious that the reason of such danger
can by no means be discovered, nor what relation it can have with
any of the numbers above-mentioned. Though this opinion has a
great deal of antiquity on its side; Aulus Gellius says, it was borrowed
from the Chaldeans, who, possibly, might receive it from Pythagoras,
whose philosophy turned much on numbers, and who imagined an
extraordinary virtue in the number 7.
The principal authors on the subject of climactericks, are Plato,
Cicero, Macrobius, Aulus Gellius, among the ancients; Argol,
Magirus, and Salmatius, among the moderns. St. Augustine, St.
Ambrose, Beda, and Bœtius, all countenance the opinion.
 Lucky and Unlucky Days.
Astrologers have also brought under their inspection and controul
the days of the year, which they have presumed to divide into lucky
and unlucky days; calling even the sacred scriptures, and the
common belief of Christians, in former ages, to their assistance for
this purpose. They pretend that the 14th day of the first month was a
blessed day among the Israelites, authorised therein, as they pretend,
by the several following passages out of Exodus, c. xii. v. 18, 40, 41,
42, 51. Leviticus, c. xxiii. v. 5. Numbers, c. xxviii. v. 16. “Four
hundred and thirty years being expired of their dwelling in Egypt,
even in the self same day departed they thence.”
With regard to evil days and times, Astrologers refer to Amos, c. 5,
v. 13, and c. vi. v. 3. Ecclesiasticus, c. ix. v. 12. Psalm, xxxvii. v. 19.
Obadiah, c. xii. Jeremiah, c. xlvi. v. 21, and to Job cursing his birth
day, chap. iii. v. 1 to 11. In confirmation of which they also quote a
calendar, extracted out of several ancient Roman catholic prayer
books, written on vellum, before printing was invented, in which
were inserted the unfortunate days of each month, as in the following
verses;—
January.—Prima dies mensis, et septima truncat ensis.
February.—Quarta subit mortem, prosternit tertia fortem.
March.—Primus mandentem, disrumpit quarta bibentem.
April.—Denus et undenus est mortis vulnere plenus.
May.—Tertius occidit, et Septimus ora relidit.
June.—Denus Pallescit, quindenus fædera nescit.
July.—Ter denus mactat, Julii denus labefactat.
August.—Prima necat fortem, perditque secunda cohortem.
September.—Tertia Septembris, et denus fert mala membris.
October.—Tertius et denus, est sicut mors alienus.
November.—Scorpius est quintus, et tertius est vita tinctus.
December.—Septimus exanguis, virosus denus ut Anguis.

This poetry is a specimen of the rusticity and ignorance at least of

the times; and is a convincing proof that Christianity had yet a very
strong tincture of the Pagan superstitions attached to it, and which
all the purity of the gospel itself, to this very day, has not been able
entirely to obliterate.
That the notion of lucky and unlucky days owes its origin to
paganism, may be proved from Roman history, where it is mentioned
that that very day four years, the civil wars were begun by Pompey
the father; Cæsar made an end of them with his son, Cneius
Pompeius being then slain; and that the Romans accounted the 13th
of February an unlucky day, because on that day they were
overthrown by the Gauls at Allia; and the Fabii attacking the city of
the Recii, were all slain with the exception of one man: from the
calendar of Ovid’s “Fastorum,” Aprilis erat mensis Græcis
auspicatissimus; and from Horace, lib. 2, ode 13, cursing the tree
that had nearly fallen upon it; ille nefasto posuit die.
The number of remarkable events that happened on some
particular days have been the principal means of confirming both
Pagans and Christians in their opinion on this subject. For example,
Alexander the Great, who was born on the 6th of April, conquered
Darius and died on the same day. The Emperor Bassianus Caracalla
was born and died on a sixth day of April. Augustus was adopted on
the 19th of August, began his Consulate, conquered the Triumviri,
and died the same day.
The Christians have observed that the 24th of February was four
times fortunate to Charles the Fifth. That Wednesday was a fortunate
day to Pope Sixtus V. for on a Wednesday he was born, on that day
made a Monk, on the same day made a General of his order, on that
day created a Cardinal, on that day elected Pope, and also on that day
inaugurated. That Thursday was a fatal day to Henry VIII. King of
England, and his posterity, for he died on a Thursday; King Edward
VI. on a Thursday; Queen Mary on a Thursday; and Queen Elizabeth
on a Thursday. The French have observed that the feast of Pentecost
had been lucky to Henry III. King of France, for on that day he was
born, on that day elected king of Poland, and on that day he
succeeded his brother Charles IX. on the throne of France.
 Genethliaci.
(From γενεθλη, origin, generation, nativity.)
These, so called in Astrology, are persons who erect Horoscopes;
or pretend what shall befal a man, by means of the stars which
presided at his nativity[10]. The ancients called them Chaldæi, and by
the general name mathematici: accordingly the several civil and
canon laws, which we find made against the mathematicians, only
respect the Genethliaci, or Astrologers; who were expelled Rome by a
formal decree of the senate, and yet found so much protection from
the credulity of the people, that they remained unmolested. Hence an
ancient author speaks of them as hominum genus, quod in civitate
nostra sempe et vetabitur, et retinebitur.
Genethliacum, (Genethliac poem,)

Is a composition in verse, on the birth of some prince, or other

illustrious person; in which the poet promises him great honours,
advantages, successes, victories, &c. by a kind of prophecy or
prediction. Such, for instance, is the eclogue of Virgil to Pollio,
beginning
Sicelides Musæ, paulo majora Canamus.

There are also Genethliac speeches or orations, made to celebrate a

person’s birth day.