Crime Analysis with Crime Mapping 4th

Edition, (Ebook PDF)

Visit to download the full and correct content document:
https://ebookmass.com/product/crime-analysis-with-crime-mapping-4th-edition-ebook
-pdf/
More products digital (pdf, epub, mobi) instant
download maybe you interests ...

Justice, Crime, and Ethics (Ebook PDF)

https://ebookmass.com/product/justice-crime-and-ethics-ebook-pdf/

Organized Crime 11th Edition

https://ebookmass.com/product/organized-crime-11th-edition/

(Original PDF) Crime Control in America What Works 4th

Edition by John L. Worrall

https://ebookmass.com/product/original-pdf-crime-control-in-
america-what-works-4th-edition-by-john-l-worrall/

More Money, More Crime: Prosperity and Rising Crime in

Latin America Marcelo Bergman

https://ebookmass.com/product/more-money-more-crime-prosperity-
and-rising-crime-in-latin-america-marcelo-bergman/
Economics of Crime and Enforcement – Ebook PDF Version

https://ebookmass.com/product/economics-of-crime-and-enforcement-
ebook-pdf-version/

(eTextbook PDF) for Multiculturalism, Crime, and

Criminal Justice

https://ebookmass.com/product/etextbook-pdf-for-multiculturalism-
crime-and-criminal-justice/

Crime Control in America: What Works? 3rd Edition,

(Ebook PDF)

https://ebookmass.com/product/crime-control-in-america-what-
works-3rd-edition-ebook-pdf/

Crime and Justice: Learning through Cases 3rd Edition,

(Ebook PDF)

https://ebookmass.com/product/crime-and-justice-learning-through-
cases-3rd-edition-ebook-pdf/

An Introduction to Crime Scene Investigation 3rd

Edition, (Ebook PDF)

https://ebookmass.com/product/an-introduction-to-crime-scene-
investigation-3rd-edition-ebook-pdf/
Detailed Contents
Preface
About the Author
Part I: Foundations of Crime Analysis
Chapter 1: Crime Analysis and the Profession
Definition of Crime Analysis
Foundation of the Definition
Crime Analysis Definition
Definitions of GIS and Crime Mapping
History of Crime Analysis
Beginnings of Crime Analysis
United States: 1900 to 1970
United States: 1970 to 2000
History of Crime Mapping
Beginnings of Crime Mapping
United States: 1900 to 1970
United States: 1970 to 2000
Research on Crime Analysis and Crime Mapping: 2000 to Present
Crime Analysis as a Career Track
Crime Analyst Qualifications and Job Descriptions
Interns/Volunteers
Crime Analysis Assistant/Technician
Entry-Level Crime Analyst
Experienced Crime Analyst
Specialty Crime Analyst
Crime Analysis Supervisor
Embedded Criminologist
CAU Organizational Chart
Developing and Improving a Crime Analysis Unit
Crime Analysis Profiles
Challenges and the Future of Crime Analysis
Summary Points
Exercises
Chapter 2: Theoretical Foundations of Crime Analysis
Environmental Criminology
Problem Analysis Triangle
Rational Choice Theory
Crime Pattern Theory

8
 Routine Activities Theory
Law of Crime Concentration
Repeat Victimization, Near Repeats, and the 80/20 Rule
Situational Crime Prevention
Displacement and Diffusion of Benefits
Displacement of Crime
Diffusion of Benefits
Opportunity
Summary Points
Exercises
Chapter 3: Evidence-Based Policing and the Role of Crime Analysis
Research on Crime Analysis and Crime Reduction
Standard Model of Policing
Community Policing
Problem-Oriented Policing
Hot Spots Policing
Focused Deterrence
Disorder Policing
Compstat
Intelligence-Led Policing
Predictive Policing
Conclusions About Police Effectiveness and Crime Analysis
Stratified Policing: Framework for Crime Analysis
Problem Stratification
Immediate Problems
Short-Term Problems
Long-Term Problems
Stratification of Police Responsibility
Stratification of Accountability and Evaluation
Conclusion
Summary Points
Exercises
Part II: Crime Analysis Process, Data, and Purpose
Chapter 4: Crime Analysis Process and Application
The Crime Analysis Process
Collection
Collation
Analysis
Data Modification Subcycle
Dissemination

9
 Feedback
Summary
Types of Crime Analysis
Crime Intelligence Analysis
Tactical Crime Analysis
Strategic Crime Analysis
Administrative Crime Analysis
Not Types of Crime Analysis
Crime Mapping by Type of Analysis
Summary Points
Exercises
Chapter 5: Crime Analysis Data and Technology
Key Terms
Data Matrix
Database
Geographic Data
Tabular Data
Secondary Data
Primary Data
Computer-Aided Dispatch System
Records Management System
Agency Management System
Information-Sharing Platforms
Jail Management System (JMS)
Criminal Justice Information Services (CJIS)
National Crime Information Center (NCIC)
Geographic Data System
Databases Used in Crime Analysis
Crime Incidents
Arrests
Calls for Service
Traffic Crashes
Other Databases
Primary Data Collection
Data Considerations
“Reported” Activity
Local and Federal Crime Data Standards
Using Call for Service Data to Study Crime Problems
Using Arrest Data to Study Crime Problems and Offenders
Data Integrity

10
 Hardware and Software Considerations
Summary Points
Exercises
Chapter 6: Geographic Data and Crime Mapping
Geographic Data
Vector Data
Raster Data
Projections
Coordinate Systems
Scale
Getting Events on a Map
Crime Mapping Techniques
Types of Descriptive Crime Mapping
Single-Symbol Mapping
Buffers
Graduated Mapping
Chart Mapping
Interactive Crime Mapping
Methods for Descriptive Crime Mapping
By Category
By Statistical Classification
By Manual Method
Classification Guidelines and Summary
Analytical Mapping: Density Mapping
Summary Points
Exercises
Chapter 7: Crime Analysis Purpose and Audience
Crime Analysis Purpose
Situational Awareness
Crime Reduction
Contrasting the Purposes of Crime Analysis Results
Crime Analysis Audiences
Crime Analysis Typology
Situational Awareness Examples
Immediate
Operational
Management
Command
Short-Term
Operational

11
 Management
Command
Long-Term
Operational
Management
Command
Crime Reduction Examples
Immediate
Operational
Management
Command
Short-Term
Operational
Management
Command
Long-Term
Operational
Management
Command
Summary Points
Exercises
Part III: Tactical Crime Analysis
Chapter 8: Repeat Incidents and Tactical Data Collection
Repeat Incidents
Repeat Incident Data
Repeat Incident Location Report
Summary
Collection and Collation of Tactical Crime Analysis Data
Crime Report Data
Modus Operandi
Persons Involved in the Crime
Vehicle Involved in the Crime
Field Information
Summary Points
Exercises
Chapter 9: Pattern Identification Process
Pattern Identification Methodology
Types of Patterns
Deduction and Induction
Initial Pattern Identification

12
 Pattern Finalization
Identifying the Principal Case
Identifying Other Key Cases in the Pattern
Identifying Additional Related Cases
Identifying Investigative Leads
Clearing Cases
Conclusion
Summary Points
Exercises
Chapter 10: Identifying Meaningful and Useful Patterns
Persons Crime and Property Crime
Potential Responses to Patterns by Police
Persons Crime
Property Crime
Identifying Meaningful Patterns
Persons Crime Pattern Key Characteristics and Examples
Robbery Patterns
Sexual Crime–Related Patterns
Property Crime Pattern Key Characteristics and Examples
Theft from Vehicle Patterns
Vehicle Theft Patterns
Residential Burglary Patterns
Commercial Burglary Patterns
Hot Product Patterns
Conclusion
Summary Points
Exercises
Chapter 11: Describing and Disseminating Known Patterns
Describing Known Patterns
Modus Operandi Summary
Suspect and Vehicle Descriptions
Time Series Analysis
Exact Time
Exact Day of Week
Exact Time of Day and Day of Week
Exact Date: Intervals Between Events
Weighted Time Span Analysis
Spatial Analysis of Patterns
Pattern Bulletins
Summary Points

13
 Exercises
Part IV: Strategic Crime Analysis
Chapter 12: Analyzing Problems, Process and Statistics
Methodology: SARA Process
Scanning and Identifying a Problem
Analysis
Research Related to the Problem
Local Context of the Problem: Agency and Community
Developing and Testing Hypotheses
Response: Role of Analysis
Assessment
Basic Statistics Used in Crime Analysis
Frequency
Cross-Tabulation
Percent
Cross-Tabulation Percent
Percentile
Rate
Mean and Standard Deviation
Conclusion
Summary Points
Exercises
Chapter 13: Analyzing Problems, Application of Techniques, Part I
What Is the Nature of the Problem?
How Frequently Is the Problem Occurring?
Frequency and Percent
Rate
Mean and Standard Deviation
Has the Problem Increased or Decreased?
By Year
By Month or Quarter
By Week
Percent Change
Anticipatory Benefit
When Is the Problem Happening?
Seasonality
Time of Day and Day of Week
Summary Points
Exercises
Chapter 14: Analyzing Problems, Application of Techniques, Part II

14
 Where Is the Problem Occurring?
Identifying and Analyzing Problem Locations
Identifying Problem Areas
Manual Method
Graduated-Color Mapping
Ellipses
Density Mapping
Summary of Problem Area Identification Methods
Who Are the Victims and Does Repeat Victimization Exist?
Who Are the Offenders and Does Repeat Offending Exist?
Why Is the Problem Occurring?
Summary Points
Exercises
Chapter 15: Strategic Crime Analysis Results and Dissemination
Choosing Analysis Information to Disseminate
Initial Analysis Results
Problem Analysis Results
Strategic Crime Analysis Products
Types of Products
Using Email for Dissemination
Guidelines for Product Contents
Guidelines for Table and Chart Contents
Guidelines for Map Contents
Practical Examples
Summary Points
Exercises
Part V: Administrative Crime Analysis
Chapter 16: Administrative Crime Analysis: Crime Analysis for Accountability
Crime Reduction Accountability
Crime Reduction Goal Development
Crime Analysis for Weekly Action-Oriented Accountability Meetings
Crime Analysis for Monthly Evaluation-Oriented Accountability Meetings
Crime and Disorder Monthly Trend Charts
Problem Comparison Chart
Crime Pattern Trend Map
Crime Analysis for Assessment of Agency Goals
Performance Indicator (Output) Products
Success Indicator (Outcome) Products
Crime and Disorder Long-Term Trend Charts
Crime Trend Comparison Charts

15
 Yearly Comparison Maps
Summary Points
Exercises
Glossary
References
Index

16
Preface

Crime analysis is a field of study and practice in criminal justice that uses systematic research methods and
data, supports the mission of police agencies, and provides information to a range of audiences. Crime
mapping is a subset of crime analysis that focuses on understanding the geographic nature of crime and other
activity. Crime analysis is a relatively new topic in criminal justice education, and this book is one of the first
to bring crime analysis and crime mapping to an undergraduate audience. A class in crime analysis provides
students with opportunities to apply theory, research methods, and statistics learned in other courses, as well
as presents information on a viable career path for criminal justice majors.

My purpose in this book is to provide an introduction to crime analysis with crime mapping through
discussion of the concepts, theories, practices, data, analysis techniques, and the role of crime analysis in
policing associated with this field of study. My purpose is not to cover current general or specific crime
analysis software products or technology. This is because the rate of change of software products and
technology is high, even though the foundations and fundamentals of crime analysis practice have remained
the same over time.

In this fourth edition, I have updated and added to the content, so the book reflects current crime analysis
practice in the United States and internationally. New to this edition are perspectives from crime analysts
from countries outside the United States, from North America, Europe, and South America. These
international crime analysts provide insight into crime analysis practices as they are conducted across the
world. This book will not serve all purposes for the growing field of undergraduate education in crime
analysis, but it is necessary for classes in which an overview of the field and fundamental techniques are
taught. The book’s website [http://www.sagepub.com/bobasantos4e] provides students with a plethora of
practical examples contributed by working crime analysts, as well as opportunities to conduct crime analyses
themselves through a variety of exercises.

The book is divided into five parts. Part I covers the foundations of crime analysis, including key definitions, a
description of the crime analysis profession and its future, theoretical foundations of crime analysis, and the
role of crime analysis in evidence-based policing strategies. Part II addresses the data and processes used in
crime analysis, geographic data and crime mapping techniques, and the purpose of crime analysis products.
Part III covers the methods and techniques of tactical crime analysis. Part IV looks at the methods and
techniques of strategic crime analysis. Part V includes a chapter on crime analysis for crime reduction
accountability–an important topic within administrative crime analysis.

The chapters in Part I lay the foundation for the rest of the book. Chapter 1 presents definitions of crime
analysis and discusses the history and future of crime analysis and crime mapping; it also includes information
on crime analysis career opportunities. In 2014, the International Association of Crime Analysts (IACA)
developed a standardized definition of crime analysis as well as its types, so those are new to this edition.
Chapter 2 provides an overview of the criminological theories that help to guide the practice of crime analysis.

17
The illustration in Chapter 3 of the policing context in which crime analysis is conducted and the discussion
of the role of crime analysis in effective policing strategies has been updated with current research in this
edition.

The four chapters that make up Part II are devoted to the topics of the data and processes used in crime
analysis, geographic data and crime mapping techniques, and a typology for crime analysis results. Chapter 4
discusses the crime analysis process and the different types of crime analysis (which were standardized by
IACA in 2014). Chapter 5 provides a review of key terms, a discussion of the kinds of data commonly used
and databases commonly accessed in crime analysis (e.g., crime, arrests, calls for service, traffic crashes, and
primary data), information on what analysts must consider when using different kinds of data for analysis, and
a brief overview of some of the hardware and software commonly used in crime analysis. Chapter 6 covers, in
more detail, geographic data, types of geographic features, geocoding, descriptive crime mapping methods,
and density mapping. Chapter 7 outlines a typology that categorizes crime analysis results by type of problem
examined, purpose of the analysis, and type of audience for which the analysis results are produced.

The chapters in Part III describe the data, methodologies, techniques, and products of tactical crime analysis.
Chapter 8 contains details of data and analysis of repeat incidents as well as data collected specifically for
tactical crime analysis and pattern identification. Chapter 9 covers the methodologies analysts employ in
identifying and finalizing patterns. Chapter 10 discusses how police respond to patterns and provides current
examples of commonly identified patterns of persons and property crime. Chapter 11 highlights specific
analytic, temporal, and spatial techniques that analysts use to identify and understand crime patterns. The
chapter closes with guidelines for creating pattern bulletins and a bulletin template example.

The chapters in Part IV concentrate on the techniques that analysts use in analyzing long-term crime and
disorder problems and provide case examples of how the techniques have been used in practice. Chapter 12 is
an overview of the problem-solving process and covers the key statistics used in strategic crime analysis.
Chapters 13 and 14 illustrate the strategic analysis of problems by demonstrating various techniques that
answer key analysis questions. Chapter 15 discusses the types of strategic crime analysis products and provides
guidelines for the substantive and formative development of such products.

Finally, Part V has been significantly changed in this edition. It contains one new chapter that covers one
aspect of administrative crime analysis: crime analysis for crime reduction accountability. Chapter 16 focuses
on the foundation of and products that support a police department’s accountability structure, which ensures
that crime reduction activities are taking place and are effective.

By no means does this book cover all facets of crime analysis; however, it does lay a solid foundation for
students’ understanding of the conceptual nature and practice of crime analysis that assists police in preventing
and reducing crime and disorder. It provides an in-depth description of this emerging field and guidelines for
the practice of crime analysis that are based on research, practice, and recent innovations, as well as previously
available and new information. It also provides opportunities for students to explore possible future careers
that support and enhance the effectiveness of modern policing.

18
19
Student Study Site
This free student study site provides additional support to students using Crime Analysis With Crime Mapping,
4th Edition. Practical crime analysis products, exercises, suggested web resources, and SAGE journal articles
with discussion questions are included on this site to provide students with additional information and
support and to get students into original research. Visit the study site at
http://www.sagepub.com/bobasantos4e.

20
Instructor Teaching Site
This set of instructor’s resources provides a number of helpful teaching aids for professors to use Crime
Analysis With Crime Mapping, 4th Edition. Included on this site are PowerPoint slides, chapter outlines, test
questions and answers, a sample syllabus, and suggested web resources for each part of the text.

21
Acknowledgments
I would like to thank all the reviewers who have helped me make improvements in this fourth edition. Thank
you to SAGE reviewers Dr. Stephanie J. Bennett, University of Portsmouth; Don Gardiner, Governors State
University; Charles J. Kocher, Cumberland County College, New Jersey; and Jonathan Allen Kringen,
University of New Haven. I would also like to show appreciation to the following police agencies and their
crime analysts who contributed products and examples used in this edition and/or in the book’s resource
materials. They include the following:

Detective Dan Benz, Seattle (Washington) Police Department

Mark Bridge, Fredrick (Maryland) Police Department
Rachel Carson, Inspired Acts, Ltd., United Kingdom
Michelle Chitolie, Port St. Lucie (Florida) Police Department
Dawn Clausius, Olathe (Kansas) Police Department
Prof. Dr. João Apolinário da Silva, Presidente da Agência Brasileira de Análise Criminal
Cheryl Davis, Port St. Lucie (Florida) Police Department
Chisen Goto, Royal Canadian Mounted Police
Kendahl Hearn, Salisbury (Maryland) Police Department
Katrine Holt, Oslo (Norway) Police District
Brandon Inscore, Greensboro (North Carolina) Police Department
Ericka Jackson, Gainesville (Florida) Police Department
Carola Jersonsky, Metropolitan Police of Buenos Aires (Argentina)
Jessica LaBlanc, Fairfax County (Virginia) Police Department
April Lee, Fort Pierce (Florida) Police Department
Brian McGrew, Adventos
Mattis Michaelsen, Oslo (Norway) Police District
Tamara Otley, Fullerton, California, Police Department
Daniel Polans, Milwaukee (Wisconsin) Police Department
Karin Schmerler, Chula Vista (California) Police Department
Alex Schneider, Arlington (Texas) Police Department
Tyr Steffensen, Oslo (Norway) Police Department
Dr. Shefali Tripathi, Gainesville (Florida) Police Department
Julie Wartell, on behalf of San Diego (California) District Attorney’s office
Michelle Wentz, Port St. Lucie (Florida) Police Department
Damien Williams, Rockhill (South Carolina) Police Department
Alisha Wilson, Roanoke (Virginia) Police Department
Alyson Yaraskovitch, Ottawa (Ontario) Police Service

I would like to extend my appreciation to members of the SAGE publishing team for their support and
assistance–Jerry Westby for his faith in my first edition and encouragement for the next three editions, Amy

22
Harris, Libby Larson, Jessica Miller, and Laura Kirkhuff.

23
About the Author

Rachel Boba Santos

is a professor at Radford University in the Department of Criminal Justice. She works with police
departments around the world assisting them in improving their crime reduction efforts, data and crime
analysis, accountability, and community partnerships. Her research focuses on environmental
criminology, the effectiveness of crime reduction efforts by police, police accountability, and crime
analysis.

24
Part I Foundations of Crime Analysis

Part I contains three chapters that provide a practical and theoretical foundation for the field of crime analysis.
Chapter 1 defines crime analysis as well as crime mapping and geographic information systems (GIS), describes
the history of the crime analysis profession, and ends with specifics about crime analysis as a career track.
Chapter 2 outlines the theoretical concepts that are most relevant for crime analysis by focusing on
understanding the opportunities for crime in immediate crime settings. Chapter 3 provides the policing
context in which crime analysis is conducted by reviewing the research on effective strategies of policing for
preventing and controlling crime, discussing the role of crime analysis in each strategy, and providing a
stratified structure for implementing crime analysis, problem solving, and accountability in police
departments.

25
Chapter 1 Crime Analysis and the Profession

This chapter serves as a foundation for the discipline of crime analysis by providing definitions of crime
analysis and crime mapping, along with an overview of the crime analysis profession in the United States. The
overview includes the profession’s history, the current research findings about crime analysis, descriptions of
potential career paths for crime analysts, and ways to develop and improve a crime analysis unit. Finally, the
chapter ends with crime analysis profiles and discussion of the future of crime analysis.

26
Another random document with
no related content on Scribd:
 Family 1. Syllidae see p. 306 Family 8. Amphinomidae see p. 318
" 2. Hesionidae " 308 " 9. Eunicidae " 318
" 3. Aphroditidae " 309 " 10. Glyceridae " 320
" 4. Phyllodocidae " 313 " 11. Sphaerodoridae " 320
" 5. Tomopteridae " 315 " 12. Ariciidae " 321
" 6. Nereidae " 315 " 13. Typhloscolecidae " 321
" 7. Nephthydidae " 317

Sub-Order 2. Spioniformia.
Family 1. Spionidae see p. 321 Family 4. Magelonidae see p. 325
" 2. Polydoridae " 323 " 5. Ammocharidae " 325
" 3. Chaetopteridae " 323

Sub-Order 3. Terebelliformia.
Family 1. Cirratulidae see p. 325 Family 3. Ampharetidae see p. 330
" 2. Terebellidae " 327 " 4. Amphictenidae " 330

Sub-Order 4. Capitelliformia.
Family. Capitellidae, see p. 331.

Sub-Order 5. Scoleciformia.
Family 1. Opheliidae see p. 331 Family 4. Scalibregmidae see p. 334
" 2. Maldanidae " 332 " 5. Chlorhaemidae " 334
" 3. Arenicolidae " 333 " 6. Sternaspidae " 335

Branch B. Cryptocephala.
Sub-Order 1. Sabelliformia.
Family 1. Sabellidae see p. 336 Family 3. Amphicorinidae see p. 339
" 2. Eriographidae " 338 " 4. Serpulidae " 339

Sub-Order 2. Hermelliformia.
Family. Hermellidae, see p. 341.

Comparative Anatomy of the Polychaeta.

General Shape of the Body.—The majority of the Polychaeta have an elongated

and very mobile body, like that of Nereis, consisting of an indefinite and usually of
a considerable number of segments; a few, however, have a shorter body, with
fewer segments, definite in number, for instance Aphrodite and Polynoë, which
have thirty to forty segments; and some Hesionids, with only some seventeen to
twenty segments.

In Aphroditidae and certain Amphinomidae the body is more or less oval in shape.
In Lipobranchius and Sternaspis it is grub-like, short, and cylindrical, with rounded
ends; in the former it is difficult to distinguish head and tail, or dorsal and ventral
surfaces.

The segments composing the trunk may be all alike, or may constitute two more or
less sharply marked regions, the thorax and abdomen, differing in the character of
the chaetae, or in their arrangement, or in some other way, as in the Sabelliformia
and the Capitelliformia.

As peculiar cuticular structures, the curious shields of Sternaspis, and of certain of

the Maldanidae may be mentioned.

The posterior extremity is generally more or less narrowed, and most of the
Nereidiformia are provided with special elongated cirri, borne by the anal segment.
In the Maldanidae and others the body terminates in a funnel, at the bottom of
which is placed the anus. Only in a few cases is the anus not terminal; in
Notopygos and other Amphinomidae, as well as in some species of Polynoë, it is
dorsal.[313] In Sabellaria and Pectinaria the hinder end of the body undergoes
great degeneration; in the former it is achaetous, but cylindrical and bent forwards
alongside the body (Fig. 131). In Pectinaria (Fig. 177), this region, which is called
the "scapha," is leaf-like, and serves to close the narrower end of the tube in which
the worm lives. Arenicola marina, and some Terebellids have no chaetae in the
hinder, narrower part of the body.

Fig. 131.—Sabellaria alveolata L. × 3. (After Malmgren.) a, Anus.

The Head.—The prostomium is, in the majority of cases, rounded or conical,

though it may be square (Nephthys) or elongated and jointed (Glycera), or even
hammer-shaped (Tomopteris); or it may be fused with the peristomium, and
apparently absent (Arenicola). In the great group Cryptocephala, the peristomium
grows forwards so as to hide the prostomium entirely.

In a few of the Nereidiformia the prostomium is compressed, and in the

Amphinomidae it is provided with a dorsal ridge or "caruncle," which is a leaf-like
process overlapping three or more segments. In many Aphroditidae (as well as in
Polydora) there is a peculiar "frontal" ridge passing forwards from the prostomial
tentacle, and downwards into the mouth (Figs. 132, c, and 133, A, x).

Fig. 132.—Aphrodite aculeata L. Ventral view of anterior region, × 6. a, Prostomium;

c, frontal ridge on prostomium; d, neuropodial cirrus; l, lower lip; m, mouth; p,
palp; s, intersegmental groove; t, tentacle; I, foot of peristomium, which has
shifted forwards so as to lie in front of the mouth; II to V, successive feet.

In all the Nereidiformia, as well as in Sabelliformia and Chlorhaemidae, the

prostomium bears sensory processes of two kinds, viz. dorsal tentacles and
ventral palps. The latter are invariably two in number, and are particularly well
developed in Aphroditidae, Nereidae, Syllidae, some of the Eunicidae, and in
Chlorhaemidae. Even when they are apparently absent, as in Nephthys, it is
possible that they are represented by certain lobes at the sides of the mouth, for in
many Syllidae they are so fused with the prostomium as to be scarcely
distinguishable. In the Chlorhaemids the palps[314] are grooved, and in the
Sabelliformia they become considerably branched, and extend round the
prostomium so as to nearly meet dorsally and ventrally. Each palp is, in this sub-
Order, represented by a greater or smaller number of long, mobile filaments,
arising from a common base; they are grooved along the inner side, ciliated, and
provided with secondary processes. The crown of "gills," in fact, is nothing more
than the greatly subdivided and enormously elongated palps, as both Pruvot[315]
and Meyer[316] have shown. In such forms as Haplobranchus and Amphicorine
the process of subdivision (branching) has only gone a short way. In all the
Sabelliformia each filament, in addition to its sensory function, aids in conveying
food to the mouth by the action of the cilia, and has a blood-vessel within, thus
acting as a respiratory organ. The filament may carry compound eyes (Fig. 143)
either at its apex (Branchiomma) or at intervals along its course (Dasychone).
 Fig. 133.—A, Anterior end of Polydora enlarged. a, Prostomium; x, frontal ridge; I,
peristomium; c', its long cirrus; II, III, etc., the following segments; c, gill; B,
head of Sabellid; P, palps (branchial crown); t, position of tentacles; l,
processes of upper lip membrane; I, peristomium raised into a collar; II, III, IV,
following segments.

In the family Serpulidae one (rarely two) of the most dorsally placed gill filaments
is enlarged terminally, and acts as a stopper or "operculum," which closes the
mouth of the tube when the animal withdraws into it. Further, in Spirorbis this
operculum is grooved on one side, and serves as a brood pouch in which the eggs
undergo development (Fig. 184, p. 341). It will be seen, therefore, that the palps
may be very important organs for the life of the worm, and they are no less
interesting to the comparative anatomist, serving as they do as an excellent
illustration of the various uses which Nature finds for one and the same organ.

In the other sub-Orders the prostomium carries neither palps nor tentacles.

Fig. 134.—Heads of various Polychaeta (diagrammatic). A, Polynoid; B, Syllid; C,

Nephthys; D, Eunice; E, Phyllodoce; F, Trophonia: a, prostomium; c, normal
cirrus; c1, peristomial cirri; c2, cirrus of second segment; c3, cirrus of third
segment; el1 point of attachment of elytron; p, palp: s, nuchal organ (ciliated
pit); t, tentacle; I, peristomium; II, III, IV, segments.

The tentacles in the Nereidiformia present a wide variation in number; probably

the typical number is three, one of which is median and two lateral—as in
Polynoids, Syllidae, and some Eunicidae. Further, there is a certain amount of
evidence in the nerve supply of the median tentacle to show that it was originally
double. The presence of four tentacles, then, as in Nephthys, Phyllodoce, and
Glycera, may be a primitive condition. By the disappearance of the paired lateral
tentacles the worm possesses a single median one, as in Aphrodite and
Amphinomids;[317] whilst a duplication of these lateral ones leads to the condition
of Eunice and Hyalinoecia, which have five tentacles. In the Chlorhaemidae the
number is further increased to five or more on each side,[318] and in the
Terebellidae these prostomial processes become very numerous.

In the Cryptocephala there is never more than a single pair of tentacles, and these
are generally reduced to a group of sensory cells, though in Sabellaria they retain
a considerable size.

In a few genera, such as Aphrodite, Nephthys, Capitella, the first postoral segment
is distinguished from the succeeding segments only by its position with regard to
the mouth (Fig. 132) and by its smaller size. But in the remainder of the
Polychaeta, with here and there an exception, the peristomium is achaetous in the
adult.[319]

Except in the Nereidiformia, peristomial or tentacular cirri are rare, being

represented in the Spioniformia by the very long "tentacles." In the Nereidiformia
one or more of the following segments may be added to the peristomium, and
share in the "cephalisation," which is so characteristic a feature in this group. In
Amphinomids the first three or four chaetigerous segments are incomplete
ventrally, owing to the shifting of the mouth backwards; these segments form
lateral lips, but they are not otherwise modified. In Phyllodoce, however, there are
four cirri on each side of the mouth, and from the arrangement in the Alciopids we
are justified in concluding that the segment which carries the four pairs of cirri is
really made up of three segments (Fig. 134, E). Among the Hesionids there are
four such "cephalised" achaetous segments with long cirri.

Fig. 135.—Sabellaria alveolata L. Ventral view of anterior region, × 10. a, Notopodial

cirrus; b, notopodium; c, neuropodium; ch, peristomial chaetae; d, neuropodial
cirrus; m, mouth; P, multifid palp (gill filaments); P', ridges after removal of gill
 filaments; s, ventral (tubiparous) gland shield; T, tentacle; I, hood formed by
peristomium; II to VI, following segments.

In a few cases, such as the Chlorhaemids and Sternaspis, and to a slight degree
in Arenicola, the "head" and even the anterior part of the worm is capable of being
withdrawn into the body.

The Parapodia and Chaetae.—The typical parts of a parapodium have been

described in the preceding chapter; here it is only necessary to refer to the series
of diagrams (Figs. 136, 137) representing the parapodia of the more common
Polychaetes, and to add a few remarks about them.

Fig. 136.—Parapodia. A, Nephthys; B, Amphinome; C, Glycera (the unlettered lobe

above g is the notopodial cirrus); D, Syllis; E, Eunice; F, Phyllodoce. a,
Notopodial cirrus; b, notopodium; c, neuropodium; d, neuropodial cirrus; g,
special gill; n, aciculum (omitted in B); x, cirriform lip of chaetigerous sac.

In most Annelids the chaetae are in two bundles on each side, but there are
certain families in which the dorsal bundle, and even the notopodium itself, is
absent, as in the Eunicidae, Syllidae, and Phyllodocidae; or the dorsal bundle may
be absent only in certain regions of the body, as in the hind-body of Terebellids. In
some Amphinomidae and Aphroditidae the notopodium is scarcely distinct as a
separate lobe, being a slight tubercle on the upper surface of the neuropodium;
but the notopodial chaetae are present, and indeed particularly well developed in
many cases.

But whilst, in the Nereidiformia, the parapodia, whether consisting of two lobes or
only one, are always well developed, and project to a more or less pronounced
degree from the sides of the body, it is otherwise in the rest of the group, where
the chaetigerous lobes are usually reduced to mere tubercles or ridges, no doubt
in relation to their burrowing or tubicolous habits. In Sternaspis the chaetae issue
directly from the body-wall.
Amongst the Nereidiformia we find examples in which the parapodia, instead of
being more or less conical "legs," are flattened fore and aft so as to serve as
efficient "fins," as in the active swimmers, Nereis virens and Nephthys caeca, and
in the pelagic Phyllodocids, Alciopids, Typhloscolecids, and Tomopteris.

Fig. 137.—Parapodia. A, Polynoë; B, Scoloplos; C, Euphrosyne. (Transverse

section of body.) a', Accessory cirrus; y, doubtful branchiae; D, Sabella
(thoracic). a, Notopodial cirrus ("elytron" in A, "gill" in B); b, notopodium; c,
neuropodium; d, neuropodial cirrus; n, aciculum (accidentally omitted in C).

Of the typical dorsal and ventral cirri, the ventral is only absent in some
Amphinomids amongst the Nereidiformia; the dorsal is absent in Nephthys and
degenerate in Glycera, whilst in a very large number of families of the other sub-
Orders neither cirrus is present. These cirri, though originally filamentous and
sensory, may, by virtue of special blood supply, become "gills," and this occurs in
several families of different sub-Orders. Thus in Eunice this gill is comb-like; in
Amphinome and in Arenicola (on certain segments) it is arborescent, as it is also
in one to three segments in Terebellids; whilst in Ariciidae, Spioniformia,
Cirratulidae, Opheliidae, and Sabellaria it remains more or less finger-shaped or
filamentous. In the family Serpulidae the thoracic cirri, both dorsal and ventral,
become flattened and extended antero-posteriorly, and unite with one another to
form the "thoracic membrane."[320] In Phyllodocidae the cirri are foliaceous and
natatory, and they contain a great quantity of glands of a peculiar character. The
Aphroditidae are distinguished from other Annelids by the possession of "elytra" or
dorsal scales, which appear to be the dorso-ventrally flattened cirri, retaining their
sensory nature, but adding to this function several others.[321]

The chaetae or bristles are mainly used in locomotion, but it is not unreasonable
to believe that some of the stronger, serrated kinds may be used as weapons of
offence and defence; certainly the Polynoids, bristling as they do with stiff chaetae
along each side, must be rather unpleasant to their smaller enemies.

The various bristles may be placed in three chief groups, viz. (1) simple; (2)
jointed; (3) uncini (see Fig. 138).
(1) The simple chaetae may be smooth and hair-shaped, i.e. "capillary," such as
are present in nearly all families: or they may be forked (Amphinomidae), comb-
shaped (Eunice), notched or serrated, or provided with a series of frills at right
angles to their length, as in Aphroditidae; or fringed along one or both sides with a
membranous expansion, as in Terebellids and Sabellids. The simple chaetae may
also be short and spine-like, as in the ventral bundles of Arenicola; or they may be
slightly curved at the end and notched, forming what are generally termed
"crotchets," such as are common amongst Oligochaeta. These "crotchets" may be
simple, or have numerous denticulations at the end (Maldanidae), or be provided
with a membranous hood (Spioniformia, Capitelliformia). In Hermione peculiar
sheathed, spear-like bristles occur (Fig. 138, N).

(2) Jointed chaetae have already been described (p. 246); they are confined to the
sub-Order Nereidiformia, and occur only in certain families.

(3) The uncini are very short chaetae, which are simply embedded in the skin, and
do not extend beyond the body-wall into the body-cavity. An uncinus is a sharply
curved hook, which may have more or less numerous secondary teeth on it. They
are characteristic of the Sabelliformia and the Terebelliformia.

The chaetae appear as solid, usually fibrillated structures, of a yellow or golden

tint, transparent and refringent. Chemically they consist of chitin, and each chaeta
is the product of a single cell. The chaetae of Euphrosyne are hollow and
calcareous, being peculiar in both characters.
 Fig. 138.—Chaetae of various Polychaetes (the magnification is not the same in all
cases). A, Doubly-fringed capillary, from Terebellid; B, hooded crotchet, from
Polydora; C, a fork, from Euphrosyne; D, jointed chaeta, from Phyllodoce; E,
simple chaeta, with serrated ridges or frills, from a Polynoid; F, jointed chaeta,
from Eunice; G, uncinus, from Pomatoceros (Serpulid); H, one of the outer
series of paleae from the hood of Sabellaria spinulosa; I, jointed chaeta, from a
Syllid; J, multidenticulate crotchet, from a Maldanid; K, comb-shaped chaeta,
from Eunice; L, uncinus of a Sabellid; M, uncinus of Terebellid (Amphitrite
Johnstoni); a, edgewise; b, side view; m, attachments of muscles into ba, basal
plate; x, accessory teeth. N, Sheathed spear of Hermione; a, the spear-shaped
capillary removed from its sheath; b, the same, with sheath.

Certain modifications of the chaetae presented by various worms deserve

mention. In Polydora (Fig. 133, A) and in Chaetopterus (Fig. 173, p. 324) those of
one segment are especially strong, but their significance is uncertain. In Capitella
those of the notopodium of the eighth and ninth segments are specially modified;
they are analogous to the copulatory chaetae of Oligochaeta. In Aphrodite, in
addition to the ordinary locomotor chaetae, there are brilliant, iridescent bristles
and peculiar felting threads arising from the indistinct notopodium; these latter,
however, are not true "chaetae," but are separate chitinous filaments similar to the
constituent fibres of an ordinary chaeta.[322]

While the chaetae in the Nereidiformia and others are grouped in bundles, those
of many other families are in vertical, transverse rows, as in Maldanidae and in
Arenicola. The uncini are always embedded in such rows, usually slightly raised
from the general level of the body surface, each being termed a "torus
uncinigerus." These tori are usually limited to the sides of the body, but in Myxicola
and in Notomastus they encroach upon the dorsal surface, and in Chaetozone,
also upon the ventral, so as nearly to encircle the body, recalling the
"perichaetous" condition of some earth-worms.

Fig. 139.—Aphrodite. Foot, × 2. a, Elytron; b, notopodium; c, neuropodium; d,

neuropodial cirrus; n, aciculum; 1, iridescent bristles; 2, stiff chaetae; 3, felt.

Gills.—We have already seen that several different organs, e.g. the palps in
Sabelliformia, the prostomial tentacles of Chlorhaemidae, and the notopodial cirri
of sundry other Polychaetes, may take on a respiratory function. There are,
however, certain "gills" developed either on the parapodium itself or elsewhere on
the body which it is difficult to homologise. Such are the retractile gills on the
parapodia of the Glyceridae (Fig. 136, C); those of Dasybranchus, near the
abdominal neuropodia; those of Mastobranchus, near the notopodia. Nephthys
has a sickle-shaped gill on the under surface of the notopodium. The long gill
filaments at the posterior end of Sternaspis, again, are only doubtfully interpreted
as the dorsal cirri of some of the posterior segments.

Since primitively the whole skin of the worm is respiratory, any part of the skin may
become more or less specialised for this function, and chiefly, of course, on the
more actively moving parapodia. The blood-vessels constituting the essential part
of the "gill" may make use of any already existing outgrowth (such as a cirrus or a
tentacle), or may push the body-wall out on their own account.

Internal Anatomy.

Probably those organs which have the greatest effect in modifying the shape of
the body are the septa, for we find in the long, free-swimming worms that these
are regularly present throughout the body, and external "segmentation" of the
body is well marked. In burrowing and tubicolous forms the septa are frequently
incompletely developed, or more or fewer may be absent; and the body becomes
less distinctly segmented externally, tends to vary greatly in diameter during
movement, or becomes plumper. With the disappearance of the septa there is also
a diminution in the number of nephridia, as in Arenicola, with only six pairs.
Further, there is frequently a dimorphism of these organs; instead of all of them
serving equally as excretory organs and as genital ducts, some of the most
anterior in the Sabelliformia and Terebelliformia become greatly enlarged, and
take on practically the whole of the former function; whilst more or fewer of the
posterior nephridia dwindle in size, and become genital ducts. The absence of
septa allows a free communication between the successive segments, and thus a
freer flow of coelomic fluid for the distension of the anterior end of the worm during
burrowing.

The alimentary system presents certain modifications of a systematic value. In

the Nereidiformia the muscular pharynx, which is always protrusible and is
preceded by an eversible buccal region, frequently encloses thickened cuticular
plates which serve as crushing and grasping organs. The form, number, and
arrangement of these "jaws" vary in the different families. They form valuable
fossil records of extinct worms.

In the Scoleciformia and Capitelliformia the buccal region exists, but there are no
jaws. In the Sabelliformia and Terebelliformia eversion does not take place and
jaws are absent.
Amongst the Nereidiformia the jaws are absent in the Phyllodocidae and
Hesionidae; when present they are usually set in the direct course of the food.
There may be one small tooth used for stabbing, as in some Syllids (Fig. 141, A);
or a circle of such denticles (Autolytus, Fig. 140, D). To these are added powerful
grasping jaws in Nereis (E); or the latter may alone be present, as in Glycera (F).
In Polynoë the four jaws are carried by hard pieces, to which the muscles are
attached (C and G). In Nephthys there is a dorsal and a ventral jaw.

Fig. 140.—Jaws of various Chaetopods. A, Transverse section of the anterior end of

Eunice; a, b, c, d, various parts of the upper series of denticles lying in a
special chamber; g, oesophagus; k, lower jaw: B, the denticles of Eunice
separated; U, upper series; a, grinder; b, forceps; c, rasping plates; d, grater; L,
lower series; j, tooth; k, base into which muscles are inserted: C, Polynoid; U,
upper, and L, lower jaws; j, tooth; k, base: D, Diagrammatic section across
pharynx of Autolytus; E, of Nereis; F, of Glycera; G, of Polynoë.

In the Eunicidae, however, the numerous denticles are carried in a special pouch
below the food tract, with which it communicates anteriorly.[323] They are arranged
in an upper and lower series. The lower series (L) consists of a pair of flat plates
(k) on each side partially embedded in and acted upon by muscles, with a harder
enamelled piece—the actual lower "tooth" (j)—at its anterior end. The upper series
(U) consists of several pieces, varying in shape and size in the various genera of
this family; but developmentally they result from modifications of two rows of small,
similar pieces.[324]

The intestine is generally straight and cylindrical, and is usually constricted by the
septa, if these are present. In the Polynoids the intervening sacculations become
so long as to receive the name of "caeca," which, in Aphrodite, become
enormously elongated (Fig. 142); there are eighteen pairs of them (c), each being
a slender tube bent upon itself, giving off short branches and dilated distally,
where it lies in the base of the parapodium.
 Fig. 141.—A, Alimentary canal of Syllid: B, transverse section of pharynx of the
same; b, buccal region; d, oesophageal outgrowth; g, salivary glands; i,
intestine; j, tooth; p, pharynx; s, gizzard: C, alimentary canal of Petta (after
Wirén); i, intestine; o, oesophagus; r, rectum; s, stomach.

Fig. 142.—Alimentary canal of Aphrodite. × 1. (From Gegenbaur.) a, Anus; b,

pharynx; c, caeca; o, mouth.

The intestine is looped in a few genera only, as in Trophonia, or coiled, as in

Sternaspis, Petta (Amphictenid, Fig. 141, C), and Ammotrypane. In the course of
the tube there may be a thick-walled muscular gizzard, with hard chitinous lining,
as in certain Terebellids, where it appears to replace, in function, the pharynx of
the Nereidiformia; in the Syllidae the gizzard is present in addition to the pharynx
(Fig. 141, A).

Glandular appendages of the oesophagus are present in many worms. Amongst

the Nereidiformia, the Syllidae and Hesionidae possess oesophageal diverticula
(Fig. 141, A, d), which are used, not for secreting a digestive fluid, but as
reservoirs for water and air swallowed by the worms; and are provided with
muscular walls, by which their contents can be driven out. They appear, in fact, to
be used like the swim-bladder of fishes.[325] Many Chaetopods take in water by
the anus—no doubt for respiratory purposes—and pass it forwards along the
intestine. In the Capitelliformia a special groove conducts the water for some
distance, then the groove becomes closed to form a canal, which, after a course
forwards as a free tube below the intestine, again enters the latter, constituting a
"siphonal apparatus," similar to that of the Echiuroids and the sea urchins.
Sense Organs.—In addition to the prostomial eyes, which are present in nearly all
the Nereidiformia and Spioniformia, eyes may exist elsewhere on the body: thus
Myxicola infundibulum and Fabricia possess a pair on the anal segment; in M.
aesthetica Clap. there is a pair to every segment; in Branchiomma there is a
compound eye near the tip of each gill filament (i.e. palp); whilst in Dasychone a
series occurs along each gill filament. All these examples belong to the
Cryptocephala, in which, owing to certain peculiar modes of life, these sense
organs are required in correspondingly peculiar positions. It is usually stated that
Polyophthalmus possesses, in addition to the usual prostomial eyes, twelve pairs
on as many successive segments; but the minute structure of these organs points
rather to their function as light-producing organs.

Fig. 143.—A gill filament, A, of Branchiomma, B, of Dasychone. a, Axis; f,

secondary filaments; o, compound eye; x, lappets.

The Capitelliformia and Opheliidae possess a pair of peculiar "ciliated pits" or

"nuchal organs" at the upper side of the head, between the prostomium and
peristomium, and capable of eversion (Fig. 144). They are most characteristically
developed in the Capitelliformia, where each organ abuts upon a special lobe of
the brain. The function of these "ciliated organs," which bear a great resemblance
to those of the Nemertines, is a matter of speculation. Similar organs, in the form
of simple pits or grooves, occur in many of the Nereidiformia, Terebelliformia, and
others.[326]

Otocysts are rare. Arenicola possesses a pair at the base of the prostomium, each
of which in some species retains an opening to the exterior.[327] They probably
serve as "organs of direction" rather than of "hearing." Aricia and Polyophthalmus
likewise have such organs on the prostomium; whilst Fabricia, Myxicola, Terebella,
and a few others possess them in the peristomium, or in some other segment of
the body.
 Fig. 144.—Ammotrypane aulogaster Rathke, enlarged. (From Cuningham.) Anterior
end. a, Prostomium; b, everted buccal region; c, notopodial cirrus; X, ciliated
organ everted; I, II, III, first three segments.

Reproductive Phenomena.—With a few exceptions mentioned below, the

Polychaeta are unisexual. The sexual cells are developed in all cases from the
lining epithelium of the body-cavity. The exact spot at which this occurs varies in
different cases; it may be, though rarely, on the floor of the body-cavity; it is more
usually on the wall of some blood-vessel, either the ventral vessel or on branches
of it; or on the many blind blood-vessels of Aphrodite. The number of such genital
organs is very great in most worms, but in those presenting two regions of the
body they are confined to the posterior segments (Sabelliformia, Terebelliformia,
Capitelliformia). The number is very limited in Arenicola and other worms
presenting but few nephridia: in the former genus there being six pairs, in
Trophonia only one pair.

The following genera are hermaphrodite:—Amphiglena, Salmacina, Protula,

Spirorbis, belonging to the Sabelliformia, to which must be added some
Hesionidae. In this family ova and spermatozoa are developed around the same
blood-vessel. But in the former group of worms (as also in Ophryotrocha) the two
kinds of cells are produced in different regions of the body. Thus in Protula the
anterior abdominal segments are male, the posterior ones female, while in
Spirorbis the reverse arrangement holds; and in Syllis corruscans the anterior
segments of the body contain eggs, whilst the posterior region contains
spermatozoa, and this region separates and becomes a male worm.

The eggs and spermatozoa in the Polychaeta are discharged into the sea either
by rupture of the body-wall or through the nephridia; the male and female
elements unite, and the resulting fertilised eggs undergo development, either
floating separately in the water, or embedded in jelly, or attached to the body or to
the tube of the worm.

The result of the segmentation of the egg is a free-swimming larva known as a

"Trochosphere," similar to that of Polygordius. The larvae of different species
present various more or less marked departures from this type, for instead of the
two girdles of cilia there may be only the anterior girdle, or there may be several
complete or incomplete girdles between the two typical ones, or there may be
(Chaetopterids) only a single girdle of cilia about the middle of the body, the two
typical girdles being absent.[328] The postoral region, after elongation, generally
becomes marked out into three segments, and these segments develop chaetae,
which are usually temporary and specially long.

The little animal is thus equipped for an independent life: the provisional chaetae
help in keeping it balanced; and in some cases (Spionidae) serve to protect the
little soft creature, for when it is touched it curls up, and its chaetae stick out at the
sides, so that it looks like a hairy caterpillar. But the larva is quite at the mercy of
the sea, for it is carried hither and thither by currents, and in this way the species
is disseminated. The larvae of the Polychaetes, like those of other animals, occur
at certain periods of the year in large quantities at the surface of the sea, and
serve as food for various larger animals.

Fig. 145.—A, Trochosphere of Nephthys. × 65. a, Anus; b, apical plate (brain); c,

apical tuft of cilia; c', girdle of cilia; i, intestine; m, mouth; st, stomach. B, Larva
of Spio, with three segments, eight days old. × 100. c, Preoral girdle of cilia; c',
preanal girdle; ch, long provisional chaetae; pr, prostomium with eyes. (From
Claparède and Metschnikoff.)

These larvae are at first very different from the adult animal, and the necessary
changes to be passed through are more or less great according to the species. It
is not our intention to describe these changes in detail.[329] The larva increases in
size, the permanent chaetae make their appearance in regular order, and the body
exhibits segmentation, the new segments always appearing just in front of the anal
segment. The internal organs gradually develop, and the prostomial and
parapodial appendages grow out in their turn. In the Sabelliformia the
multifilamentous "gills" arise by the continued branching of an at first simple
process (the palp) arising from the latero-ventral surface of each side of the
preoral lobe.[330] These gradually encroach dorsally and ventrally till the
prostomium is more or less encircled; meanwhile the peristomium grows forwards
so as to conceal the prostomium, which no longer increases at the same rate as
does the rest of the body.

Although most worms appear to discharge their ova directly into the sea and take
no further care of them, some make provision for their offspring either by laying
the eggs in a jelly, which will serve as food for the young larvae—Aricia, Ophelia,
Protula, Phyllodoce—or by attaching them to their body. In certain Polynoids the
eggs are attached by means of a secretion to the back, under the elytra, where
they undergo development up to a certain stage. In Exogone and some other
Syllids they are attached to the ventral cirri, or in Grubea limbata, all over the
back. In the female Autolytus (Sacconereis) a ventrally-placed brood sac is formed
by the hardening of a secretion; the eggs develop into embryos inside the brood
sac, and then become free, with head appendages and three pairs of parapodia.
Enormous numbers of such embryos may occur; for instance, some 300 were
counted in a brood sac of Autolytus ebiensis. In the case of tubicolous worms, the
eggs are frequently attached to the tube, either inside or outside. In Spirorbis and
Salmacina the operculum serves as a brood pouch.

Only a very few species are known to be viviparous, viz. Syllis vivipara Kr.,
Cirratulus chrysoderma Clap., Marphysa sanguinea Mont., and Nereis diversicolor
Müll.

In most genera there is no external difference between a mature worm filled with
generative products and an immature one, except, it may be, in the colour; for the
yolk of the eggs is frequently tinted yellow, or pink, or bluish, while the
spermatozoa in mass are white; so that the normal colouring of the worm may be
modified when filled with these elements. But in a few instances striking
anatomical peculiarities are exhibited by the mature worm.[331] In many species of
Nereis, for instance, those segments containing the generative products undergo
more or less extensive changes, while the anterior ones remain unaltered. The
body of the ripe Nereis is then distinguishable into an anterior non-sexual region
and a posterior sexual region; and so great are these changes in certain species
that the mature worms were for a long time believed to belong to a different genus,
and received the name Heteronereis. But we now know their true relations, thanks
to the work of Claparède and others. The males in the Heteronereid phase have
fewer unaltered anterior segments than the females, so that there is a sexual
dimorphism.

Fig. 146.—Male "Heteronereis" of N. pelagica L. × 1. A, Non-sexual region; B,

sexual, modified region. (From Ehlers.)

The changes which Nereis undergoes in its transformation affect chiefly (a) the
shape of the parapodia, and (b) the form of the chaetae of these parapodia. Other
organs may also be affected, though less noticeably; thus the eyes become
enlarged, the intestine may become so compressed by the generative products as
to be functionless, and the tail develops special sensory papillae.[332]

In the parapodia an increase in size and a sharper delineation of the various parts
take place; then flattened foliaceous outgrowths (Fig. 147, x, y) arise from certain
lobes of the feet, in which, too, the blood supply becomes greatly increased. The
old chaetae are pushed out by the development of new ones of quite a different
shape; these are jointed like the old ones, but the appendix is, in many species at
least, flattened and oar-shaped (Fig. 123, C, p. 246); and the chaetae are
arranged in a fan-like manner. Both these modifications are in evident relation to
the free-swimming habit which the Heteronereid now adopts. The new foot serves
as a swimming organ, the old one was a walking appendage.

Fig. 147.—Parapodium of male "Heteronereis" of N. pelagica L. × 10. (From Ehlers.)

a, Notopodial cirrus; b, notopodium; c, neuropodium with new chaetae; c',
foliaceous outgrowth; d, neuropodial cirrus; x, y, foliaceous outgrowths.

Whilst some species, such as the common British N. diversicolor, undergo no

change, and others become modified as just described, others, again, are
polymorphic. Claparède was the first to show that N. dumerilii may occur in at
least five different mature forms; these differ from one another in size, colour,
mode of life, character of the eggs, etc. The immature forms may become ripe and
lay eggs while still retaining the "Nereid" characteristics, or these immature forms
may become "Heteronereids"[333] whilst the sexual elements are ripening. There
are then three different kinds of males and of females in this one species, some
being found at the bottom of the sea, as the large Heteronereid form, while the
small Heteronereid swims on the surface. The relations of these various forms to
one another, and the causes leading to the assumption of a Heteronereid
condition in some cases and not in others, are unknown.

A somewhat similar phenomenon is exhibited by members of the family Syllidae.

[334] In this family sexual reproduction is frequently accompanied by the asexual
modes of fission and gemmation. In some genera, such as Eusyllis, Odontosyllis,
and Exogone, there occur changes quite similar to those characterising
"Heteronereis"—that is, the posterior segments in which the genital organs exist
become altered, so that the worm consists of two distinct regions, and is termed a
"Heterosyllis." The most marked change is the appearance of a dorsal bundle of
long capilliform chaetae in each of the genital segments (Fig. 148, I).

But in other genera the hinder genital region of the body becomes separated, on
maturity, from the anterior non-sexual region. Various stages of this "schizogamy,"
or fission into a sexual and a non-sexual zooid, have been observed in different
genera. In the genus Syllis the first segment of the sexual zooid, after its
separation from the asexual zooid, proceeds to bud forth a head. The character of
the head is alike in both sexes, though different species present heads of different
shapes; and as the worms were originally described as distinct genera, the names
then given are retained as descriptive terms. Thus the "Chaetosyllis" form has only
two tentacles; the "Ioda" form has three tentacles and a pair of palps. One and the
same species (e.g. S. hyalina) may successively pass through these stages.

With regard to the asexual portion, there is a regeneration of the tail segments
after the sexual zooid has separated; and the number of segments so regenerated
is usually equal to those that have become sexual. After a time these newly
formed segments will produce generative organs, and take on the characteristic
natatory chaetae, and this region will in its turn separate.

But in other genera, such as Autolytus, the regeneration of segments may

commence before the separation of the sexual zooid; and the head of the sexual
zooid becomes budded out before separation from the asexual portion. So that the
animal now consists of two worms, each with its own head, separated by a region
or zone of proliferation (Fig. 148, IV). Moreover, in some species not only is the
hinder part of the body converted into a sexual zooid, but the zone of proliferation
becomes very active, and produces by gemmation a large number of segments,
which become marked out, by the appearance of heads at intervals, into a number
of zooids, in which genital organs will later make their appearance. A chain of as
many as sixteen zooids may be formed in Autolytus (Fig. 148, V)—the hindermost
by conversion of the hinder part of the body of the original "stock," the intervening
zooids by gemmation.
 Fig. 148.—Diagrams illustrating the various stages in the asexual formation of a
chain of zooids. (Modified from Malaquin.)
I, Heteronereid or Heterosyllid stage. A, Non-sexual; A', sexual region of the body,
with modified parapodia.
II, Syllis. The hinder sexual region, B, is similarly modified, and will separate from
the parent zooid, A, and become an independent zooid.
III, Autolytus. The hinder zooid, B, develops a head by budding before separation.
IV, Autolytus, etc. A zone of budding (z) makes its appearance in front of the head of
B, and by its growth will give rise to a series of new segments in the middle of
the body.
V, Myrianida, Autolytus, etc. From this zone of budding a very large number of
segments have been formed, which have, further, become grouped so as to
form three individuals, C, D, E; B is the hindmost zooid, which is either formed
from the hinder segments of the parent zooid or is produced by budding, like C,
D, E.

One original "stock," or asexual zooid, thus produces several sexual zooids, but
these are only of one sex for a given stock. The males differ in several important
characters from the females; so different, indeed, are the two sexes that before
their history was worked out by Agassiz[335] they were placed in different genera.
The male zooid has thus come to be known as Polybostrichus (Fig. 149, B). It has
three tentacles and two bifid palps; there are two pairs of peristomial cirri; the
testes are confined to the four anterior segments, which are without natatory
chaetae. The female is termed Sacconereis, owing to the possession of a great
ventral brood sac; its head possesses no separate palps; the peristomium carries
only one cirrus on each side; ova occur in every segment of the body, and may
even extend into the hinder segments of the asexual zooid (Fig. 149, C).
 Fig. 149.—Myrianida fasciata. (From Malaquin.) The bright red markings of the living
animal are here represented black. A, An asexual individual which has
produced by budding from the zone (z) a chain of twenty-nine zooids, the
oldest being labelled 1, the youngest 29. B, A ripe male zooid (Polybostrichus),
with three tentacles and a pair of forked palps (p). There are five unaltered
anterior segments. C, A ripe female zooid (Sacconereis) with the palps fused
with the prostomium; s, the ventral brood pouch projecting on each side; t,
tentacles.

A further development of this process of gemmiparity is exhibited by Myrianida.

Here, there is no conversion of the hinder segments, but the normal preanal zone
of proliferation gives rise to a large number of new segments. After a time the
most anterior of these becomes a head, and thus a new zooid is marked out. The
zone of proliferation immediately in front of the new head now proceeds to form
new segments, and a second zooid results. This process goes on till a
considerable number of new worms have been formed at the tail of the original
one, the oldest of these new ones being the most posterior, the youngest next the
original "stock." In each zooid there is a zone of activity which adds to its number
of segments, so that as we pass backwards the zooids increase in size. As many
as twenty-nine such zooids may be formed in this way entirely by gemmation; and
as each zooid becomes completed, genital organs make their appearance, and
when these are ripe the zooid separates from the "colony" and leads an
independent life. Here, as in Autolytus, the sexes are dimorphic, the male and
female resembling those of that genus.

The process of gemmation, as seen in Autolytus, closely resembles that exhibited

by certain Oligochaeta (Naididae), where there exists a definite alternation of
generations; the production of new individuals by gemmation occurring throughout
the greater part of the year, and sexual reproduction recurring only at certain
intervals. In the Polychaeta such alternation exists in Myrianida; but it is only the
terminal link of a series, which takes its starting-point in the process exhibited by
the majority of Annelids, where no sexual character marks maturity. The next
stage is presented by "epigamous" forms like Heteronereis and Heterosyllis; then