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 ZUTTER
SMART
PRINCIPLES of

MyLab ™
MANAGERIAL

MANAGERIAL FINANCE
PRINCIPLES of
FINANCE
Improving Results
A proven way to help individual students achieve
the goals that educators set for their course.

Engaging Experiences BRIEF

Dynamic, engaging experiences that personalize
and activate learning for each student. 8th edition

An Experienced Partner
From Pearson, a long-term partner
with a true grasp of the subject,
excellent content, and an eye on the
future of education.

BRIEF
www.pearson.com

8th
edition ZUTTER SMART
 Dedicated to our good friend and mentor,
Dr. Lawrence J. Gitman,
who trusted us as coauthors and successors
of Principles of Managerial Finance, Brief Edition.
CJZ
SBS

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 Brief Contents

Contents ix
About the Authors xxv
Preface xxvii
Acknowledgments xl

PART 1 Introduction to Managerial PART 6 Long-Term Financial

Finance 1 Decisions 505
1 The Role of Managerial Finance 2 12 Leverage and Capital Structure 506
2 The Financial Market Environment 39 13 Payout Policy 558

PART 2 Financial Tools 71 PART 7 Short-Term Financial

3 Financial Statements and Ratio Analysis 72
Decisions 593
4 Long- and Short-Term Financial 14 Working Capital and Current Assets
Planning 137 Management 594
5 Time Value of Money 183 15 Current Liabilities Management 636

PART 3 Valuation of Securities 243

6 Interest Rates and Bond Valuation 244

7 Stock Valuation 291
Appendix A-1
Glossary G-1
PART 4 Risk and the Required Rate of
Index I-1
Return 327
8 Risk and Return 328
9 The Cost of Capital 376

PART 5 Long-Term Investment

Decisions 405
10 Capital Budgeting Techniques 406
11 Capital Budgeting Cash Flows and Risk
Refinements 445

vii

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 Contents

About the Authors xxv

Preface xxvii
Acknowledgments xl

PART 1 Introduction to Managerial Finance 1

1 1.1 Finance and the Firm 3 1.4 Developing Skills for Your
The Role of What Is Finance? 3 Career 30
Managerial What Is a Firm? 4 Critical Thinking 30
Finance 2 What Is the Goal of the Firm? 4 Communication and Collaboration 31
The Role of Business Ethics 8 Financial Computing Skills 31
in practice FOCUS ON PRACTICE:
Summary 31
Must Search Engines Screen Out Fake
News? 10 Self-Test Problem 33
➔ REVIEW QUESTIONS 10 Warm-Up Exercises 33
Problems 35
1.2 Managing the Firm 10 Spreadsheet Exercise 38
The Managerial Finance Function 11
➔ REVIEW QUESTIONS 19

1.3 Organizational Forms,

­ axation, and the Principal-Agent
T
­Relationship 19
Legal Forms of Business Organization 19
Agency Problems and Agency Costs 26
Corporate Governance 26
➔ REVIEW QUESTIONS 30

ix

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 x Contents

2 2.1 Financial Institutions 40 2.4 The Securities Issuing

The Financial Commercial Banks, Investment Banks, Process 53
Market and the Shadow Banking System 40 Issuing Common Stock 53
Environment 39 ➔ REVIEW QUESTIONS 42 ➔ REVIEW QUESTIONS 61

2.2 Financial Markets 42 2.5 Financial Markets in Crisis 61

The Relationship Between Institutions Financial Institutions and Real Estate
and Markets 42 Finance 62
The Money Market 43 Spillover Effects and Recovery from the
The Capital Market 44 Great Recession 64
The Role of Capital Markets 48 ➔ REVIEW QUESTIONS 65
in practice FOCUS ON ETHICS:
Should Insider Trading Be Legal? 50 Summary 65
Self-Test Problem 67
➔ REVIEW QUESTIONS 51
Warm-Up Exercises 68
2.3 Regulation of Financial Markets Problems 68
and Institutions 51 Spreadsheet Exercise 70
Regulations Governing Financial
Institutions 51
Regulations Governing Financial
Markets 52
➔ REVIEW QUESTIONS 53

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PART 2 Financial Tools 71
3 3.1 The Stockholders’ Report 73
Financial The Letter to Stockholders 73
Statements and The Four Key Financial Statements 73
Ratio Analysis 72 in practice FOCUS ON ETHICS:
Earnings Shenanigans 74
Notes to the Financial Statements 80
Consolidating International Financial
Statements 80
➔ REVIEW QUESTIONS 81
3.2 Using Financial Ratios 82
Interested Parties 82
Types of Ratio Comparisons 82
Cautions About Using Ratio Analysis 85
➔ REVIEW QUESTIONS 86
3.3 Liquidity Ratios 87
Current Ratio 87
Quick (Acid-Test) Ratio 89
➔ REVIEW QUESTIONS 90
3.4 Activity Ratios 90
Inventory Turnover 90
Average Collection Period 91
Average Payment Period 93
Total Asset Turnover 93
➔ REVIEW QUESTION 94
3.5
Debt Ratios 94
Debt Ratio 96
Debt-to-Equity Ratio 96
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Times Interest Earned Ratio 97
Fixed-Payment Coverage Ratio 98
➔ REVIEW QUESTIONS 98
3.6 Profitability Ratios 98
Common-Size Income Statements 98
Gross Profit Margin 99
Operating Profit Margin 101
Net Profit Margin 101
Earnings Per Share (EPS) 102
Return on Total Assets (ROA) 103
Return on Equity (ROE) 103
➔ REVIEW QUESTIONS 105
3.7 Market Ratios 105
Price/Earnings (P/E) Ratio 106
Market/Book (M/B) Ratio 108
➔ REVIEW QUESTION 109
3.8 A Complete Ratio Analysis 109
Summary of Whole Foods’ Financial
­Condition 109
DuPont System of Analysis 113
➔ REVIEW QUESTIONS 116
Summary 116
Self-Test Problems 118
Warm-Up Exercises 119
Problems 120
Spreadsheet Exercise 135
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 xii Contents

4 4.1 The Financial Planning 4.4 Profit Planning: Pro Forma

Long- and Short- Process 138 Statements 159
Term Financial Long-Term (Strategic) Financial Plans 138 Preceding Year’s Financial Statements 160
Planning 137 Short-Term (Operating) Financial Sales Forecast 160
Plans 139
➔ REVIEW QUESTION 161
➔ REVIEW QUESTIONS 140
4.5 Preparing the Pro Forma
4.2 Measuring the Firm’s Cash Income Statement 161
Flow 140 Considering Types of Costs and
Depreciation 140 Expenses 162
Depreciation Methods 141 ➔ REVIEW QUESTIONS 163
Developing the Statement of Cash
Flows 143 4.6 Preparing the Pro Forma
Free Cash Flow 148 Balance Sheet 163
in practice FOCUS ON ETHICS: ➔ REVIEW QUESTIONS 165
Is Excess Cash Always a Good
Thing? 149 4.7 Evaluation of Pro Forma
➔ REVIEW QUESTIONS 150 Statements 165
➔ REVIEW QUESTIONS 166
4.3 Cash Planning: Cash
Budgets 151 Summary 166
The Sales Forecast 151 Self-Test Problems 168
Preparing the Cash Budget 151 Warm-Up Exercises 170
Evaluating the Cash Budget 156 Problems 170
Coping with Uncertainty in the Cash Spreadsheet Exercise 181
Budget 157
Cash Flow within the Month 159
➔ REVIEW QUESTIONS 159

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5 5.1 The Role of Time Value in
Time Value of Finance 184
Money 183 Future Value Versus Present Value 184
Computational Tools 185
Basic Patterns of Cash Flow 187
➔ REVIEW QUESTIONS 187
5.2
Single Amounts 188
Future Value of a Single Amount 188
Present Value of a Single Amount 192
➔ REVIEW QUESTIONS 195
➔ EXCEL REVIEW QUESTIONS 195
5.3 Annuities 196
Types of Annuities 196
Finding the Future Value of an Ordinary
Annuity 197
Finding the Present Value of an Ordinary
Annuity 198
Finding the Future Value of an Annuity
Due 200
Finding the Present Value of an Annuity
Due 201
Finding the Present Value of a
Perpetuity 203
➔ REVIEW QUESTIONS 204
➔ EXCEL REVIEW QUESTIONS 204
5.4 Mixed Streams 205
Future Value of a Mixed Stream 205
Present Value of a Mixed Stream 207
➔ REVIEW QUESTION 208
➔ EXCEL REVIEW QUESTION 208
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5.5 Compounding Interest More
Frequently Than Annually 208
Semiannual Compounding 208
Quarterly Compounding 209
A General Equation for
Compounding 210
Using Computational Tools for
Compounding 210
Continuous Compounding 211
Nominal and Effective Annual Rates of
Interest 212
➔ REVIEW QUESTIONS 214
in practice FOCUS ON ETHICS:
Was the Deal for Manhattan a
Swindle? 214
➔ EXCEL REVIEW QUESTIONS 215
5.6 Special Applications of
Time Value 215
Determining Deposits Needed to
Accumulate a Future Sum 215
Loan Amortization 216
Finding Interest or Growth Rates 218
Finding an Unknown Number of
Periods 220
➔ REVIEW QUESTIONS 222
➔ EXCEL REVIEW QUESTIONS 222
Summary 223
Self-Test Problems 224
Warm-Up Exercises 225
Problems 226
Spreadsheet Exercise 242
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 xiv Contents

PART 3 Valuation of Securities 243

6 6.1 Interest Rates and Required 6.3 Valuation Fundamentals 265

Interest Rates Returns 245
and Bond Interest Rate Fundamentals 245
Valuation 244 Term Structure of Interest Rates 250
Risk Premiums: Issuer and Issue
Characteristics 255
➔ REVIEW QUESTIONS 256
6.2 Government and Corporate
Bonds 257
Legal Aspects of Corporate Bonds 257
Cost of Bonds to the Issuer 259
General Features of a Bond Issue 259
Bond Yields 260
Bond Prices 260
Key Inputs 265
Basic Valuation Model 266
➔ REVIEW QUESTIONS 267
6.4 Bond Valuation 267
Bond Fundamentals 268
Bond Valuation 268
Semiannual Interest Rates and Bond
Values 270
Changes in Bond Values 272
Yield to Maturity (YTM) 275
➔ REVIEW QUESTIONS 277
➔ EXCEL REVIEW QUESTIONS 277

Bond Ratings 261

Summary 277
Common Types of Bonds 261
Self-Test Problems 279
International Bond Issues 263
Warm-Up Exercises 280
in practice FOCUS ON ETHICS:
Problems 281
“Can Bond Ratings Be Trusted?” 263
Spreadsheet Exercise 290
➔ REVIEW QUESTIONS 264

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 xv
Contents

7 7.1 Differences Between Debt 7.4 Decision Making and Common

Stock Valuation 291 and Equity 292 Stock Value 312
Voice in Management 292 Changes in Expected Dividends 313
Claims on Income and Assets 292 Changes in Risk 313
Maturity 293 Combined Effect 314
Tax Treatment 293 ➔ REVIEW QUESTIONS 314
➔ REVIEW QUESTION 293
Summary 315
7.2 Common and Preferred Self-Test Problems 317
Stock 293 Warm-Up Exercises 317
Common Stock 294 Problems 318
Preferred Stock 297 Spreadsheet Exercise 326
➔ REVIEW QUESTIONS 299

7.3 Common Stock Valuation 299

Market Efficiency and Stock Valuation 299
Common Stock Dividend Valuation
Model 301
in practice FOCUS ON PRACTICE:
Understanding Human Behavior Helps
Us Understand Investor Behavior 302
Free Cash Flow Stock Valuation
­Model 306
Other Approaches to Common Stock
Valuation 309
➔ REVIEW QUESTIONS 312

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 xvi Contents

PART 4 Risk and the Required Rate of Return 327

8 8.1 Risk and Return 8.4 Risk and Return: The Capital
Risk and Fundamentals 329 Asset Pricing Model (CAPM) 348
Return 328 What Is Risk? 329 Types of Risk 348
What Is Return? 329 The Model: CAPM 349
Risk Preferences 331 ➔ REVIEW QUESTIONS 358
➔ REVIEW QUESTIONS 332
Summary 358
8.2 Risk of a Single Asset 332 Self-Test Problems 360
Risk Assessment 332 Warm-Up Exercises 361
Risk Measurement 335 Problems 362
➔ REVIEW QUESTIONS 340 Spreadsheet Exercise 374

8.3 Risk of a Portfolio 340

Portfolio Return and Standard
Deviation 340
Correlation 342
Diversification 343
Correlation, Diversification, Risk, and
Return 345
International Diversification 346
in practice GLOBAL FOCUS:
An International Flavor to Risk
Reduction 347
➔ REVIEW QUESTIONS 348

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Contents

9 9.1 Overview of the Cost of 9.4 Cost of Common Stock 386

The Cost of Capital 377 Finding the Cost of Common Stock
Capital 376 in practice FOCUS ON ETHICS: Equity 387
The Cost of Capital Also Rises 377 Cost of Retained Earnings 390
The Basic Concept 378 ➔ REVIEW QUESTIONS 391
Sources of Long-Term Capital 380
➔ REVIEW QUESTIONS 381 9.5 Weighted Average Cost
of Capital 391
9.2 Cost of Long-Term Debt 381 Calculating the Weighted Average Cost of
Net Proceeds 381 Capital (WACC) 391
Before-Tax Cost of Debt 382 Capital Structure Weights 393
After-Tax Cost of Debt 384 ➔ REVIEW QUESTIONS 394

➔ REVIEW QUESTIONS 385

Summary 394
➔ EXCEL REVIEW QUESTION 385 Self-Test Problem 395
Warm-Up Exercises 396
9.3 Cost of Preferred Stock 385
Problems 397
Preferred Stock Dividends 386
Spreadsheet Exercise 404
Calculating the Cost of Preferred
Stock 386
➔ REVIEW QUESTION 386

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 xviii Contents

PART 5 Long-Term Investment Decisions 405

10 10.1 Overview of Capital 10.5 Comparing NPV and IRR

Capital Budgeting Budgeting 407
Techniques 406 Motives for Capital Expenditure 407
Steps in the Process 407
Basic Terminology 408
Capital Budgeting Techniques 409
➔ REVIEW QUESTION 410
10.2 Payback Period 410
Decision Criteria 411
Pros and Cons of Payback Analysis 411
➔ REVIEW QUESTIONS 414
10.3 Net Present Value (NPV) 414
Decision Criteria 415
NPV and the Profitability Index 416
NPV and Economic Value Added 417
➔ REVIEW QUESTIONS 418
➔ EXCEL REVIEW QUESTION 418
Techniques 422
Net Present Value Profiles 422
Conflicting Rankings 424
Which Approach Is Better? 427
in practice FOCUS ON ETHICS:
Baby You Can Drive My Car—Just Not
a VW Diesel 429
➔ REVIEW QUESTIONS 429
Summary 430
Self-Test Problem 431
Warm-Up Exercises 432
Problems 433
Spreadsheet Exercise 443

10.4 Internal Rate of Return

(IRR) 419
Decision Criteria 419
Calculating the IRR 419
➔ REVIEW QUESTIONS 422
➔ EXCEL REVIEW QUESTION 422

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11 11.1 Project Cash Flows 446
Capital Budgeting Major Cash Flow Types 446
Cash Flows Replacement Versus Expansion
and Risk Decisions 447
Refinements 445 Sunk Costs and Opportunity Costs 448
in practice FOCUS ON ETHICS:
Fumbling Sunk Costs 449
➔ REVIEW QUESTIONS 450
11.2 Finding the Initial
Investment 450
Installed Cost of the New Asset 451
After-Tax Proceeds from the Sale of the
Old Asset 451
Change in Net Working Capital 454
Calculating the Initial Investment 455
➔ REVIEW QUESTIONS 456
11.3 Finding the Operating Cash
Flows 456
Interpreting the Term Cash Flows 456
Interpreting the Term After-Tax 457
Interpreting the Term Incremental 459
➔ REVIEW QUESTIONS 461
11.4 Finding the Terminal Cash
Flow 461
After-Tax Proceeds from the Sale of New
and Old Assets 461
Change in Net Working Capital 462
➔ REVIEW QUESTION 463
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11.5 Risk in Capital Budgeting
­(Behavioral Approaches) 463
Breakeven Analysis 464
Scenario Analysis 466
Simulation 467
➔ REVIEW QUESTIONS 468
➔ EXCEL REVIEW QUESTION 468
11.6 Risk-Adjusted Discount
Rates 468
Determining Risk-Adjusted Discount Rates
(RADRs) 469
Applying RADRs 471
Portfolio Effects 474
RADRs in Practice 474
➔ REVIEW QUESTIONS 476
11.7 Capital Budgeting
Refinements 476
Comparing Projects with Unequal
Lives 476
Recognizing Real Options 479
Capital Rationing 480
➔ REVIEW QUESTIONS 483
➔ EXCEL REVIEW QUESTION 483
Summary 483
Self-Test Problems 485
Warm-Up Exercises 487
Problems 489
Spreadsheet Exercise 504
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 xx Contents

PART 6 Long-Term Financial Decisions 505

12 12.1 Leverage 507 12.4 Choosing the Optimal Capital

Leverage Breakeven Analysis 508 Structure 539
and Capital Operating Leverage 511 Linkage 539
Structure 506 in practice FOCUS ON PRACTICE: Estimating Value 540
Qualcomm’s Leverage 514 Maximizing Value Versus Maximizing
Financial Leverage 515 EPS 540
Total Leverage 519 Some Other Important
Considerations 542
➔ REVIEW QUESTIONS 521
➔ REVIEW QUESTIONS 543
12.2 The Firm’s Capital
Summary 543
Structure 521
Self-Test Problems 544
Types of Capital 522
Warm-Up Exercises 546
External Assessment of Capital
Structure 522 Problems 546
Capital Structure of Non–U.S. Firms 524 Spreadsheet Exercise 557
Capital Structure Theory 525
Optimal Capital Structure 533
➔ REVIEW QUESTIONS 535

12.3 EBIT–EPS Approach to

Capital Structure 536
Presenting a Financing Plan
Graphically 536
Comparing Alternative Capital
Structures 537
Considering Risk in EBIT–EPS
Analysis 538
Basic Shortcoming of EBIT–EPS
Analysis 539
➔ REVIEW QUESTION 539

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 xxi
Contents

13 13.1 The Basics of Payout Policy 559
Payout Policy 558 Elements of Payout Policy 559
Trends in Earnings and Dividends 559
Trends in Dividends and Share
Repurchases 560
➔ REVIEW QUESTIONS 562
in practice
FOCUS ON ETHICS:
Buyback Mountain 563
13.2 The Mechanics of Payout
Policy 563
Cash Dividend Payment Procedures 564
Share Repurchase Procedures 566
Tax Treatment of Dividends and
Repurchases 567
Dividend Reinvestment Plans 568
Stock Price Reactions to Corporate
Payouts 568
➔ REVIEW QUESTIONS 569
13.5 Types of Dividend Policies 575
Constant-Payout-Ratio Dividend
Policy 575
Regular Dividend Policy 576
Low-Regular-and-Extra Dividend
Policy 577
➔ REVIEW QUESTION 577
13.6 Other Forms of Dividends 577
Stock Dividends 578
Stock Splits 579
➔ REVIEW QUESTIONS 581
Summary 581
Self-Test Problem 583
Warm-Up Exercises 583
Problems 584
Spreadsheet Exercise 591

13.3 Relevance of Payout Policy 569

Residual Theory of Dividends 569
The Dividend Irrelevance Theory 570
Arguments for Dividend Relevance 571
➔ REVIEW QUESTIONS 572

13.4 Factors Affecting Dividend

Policy 572
Legal Constraints 573
Contractual Constraints 574
Growth Prospects 574
Owner Considerations 574
Market Considerations 575
➔ REVIEW QUESTION 575

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 xxii Contents
PART 7 Short-Term Financial Decisions 593
14 14.1 Net Working Capital
Working Fundamentals 595
Capital and Working Capital Management 595
Current Assets Net Working Capital 596
Management 594 Tradeoff between Profitability and
Risk 596
➔ REVIEW QUESTIONS 598
14.2 Cash Conversion Cycle 598
Calculating the Cash Conversion
Cycle 599
Funding Requirements of the Cash
Conversion Cycle 600
Strategies for Managing the Cash
Conversion Cycle 604
➔ REVIEW QUESTIONS 604
14.3 Inventory Management 604
Differing Viewpoints about Inventory
Level 605
Common Techniques for Managing
Inventory 605
International Inventory Management 610
➔ REVIEW QUESTIONS 610
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14.4 Accounts Receivable
Management 610
Credit Selection and Standards 611
in practice FOCUS ON ETHICS:
If You Can Bilk It, They Will Come 612
Credit Terms 616
Credit Monitoring 618
➔ REVIEW QUESTIONS 620
14.5 Management of Receipts and
Disbursements 620
Float 621
Speeding Up Collections 621
Slowing Down Payments 622
Cash Concentration 622
Zero-Balance Accounts 623
Investing in Marketable Securities 624
➔ REVIEW QUESTIONS 625
Summary 626
Self-Test Problems 628
Warm-Up Exercises 628
Problems 629
Spreadsheet Exercise 634
02/01/18 5:45 PM
Another random document with
no related content on Scribd:
The nervous system consists of a central ganglion situated in the
proboscis sheath; it is oval and flattened in shape. The ganglion
gives off nerves to the proboscis, and two main trunks which pierce
the proboscis-sheath and run backward surrounded by a cluster of
muscle-fibres, the whole being termed the retinaculum; in the male
they are in connexion with a special genital ganglion which lies near
the ductus ejaculatorius.

With the exception of certain sensory papillae in the neighbourhood

of the male genital orifice, and of three similar papillae mentioned by
Kaiser on the proboscis, the Acanthocephala are devoid of sense
organs.

The Acanthocephala are dioecious; their generative organs are

developed in connexion with the ligament, a cord-like structure which
arises between the inner and outer layer of the hinder end of the
proboscis sheath and traverses the body-cavity, ending posteriorly in
connexion with the genital ducts. The testes lie in this ligament; they
are paired oval bodies which open each into a vas deferens. The
vasa deferentia each bear three lateral diverticula, the vesiculae
seminales; and three pairs of cement glands pour their secretion into
a duct which opens into the vasa deferentia; the latter unite and
open by a penis which is withdrawn into a genital bursa, but is
capable of being extruded.
 Fig. 96.—An optical section through a male Neorhynchus clavaeceps
Zed. (From Hamann.) a, Proboscis; b, proboscis sheath; c,
retractor of the proboscis; d, cerebral ganglion; f, f, retractors of
the proboscis sheath; g, g, lemnisci, each with two giant nuclei; h,
space in sub-cuticular layer of the skin; l, ligament; m, m, testes;
o, glands on vas deferens; p, giant nucleus in skin; q, opening of
vas deferens.

The two ovaries are formed in the ligament of the female in a

corresponding position to that occupied by the testes in the male, but
at an early stage they break down into packets of cells, of which
those of the peripheral layer develop into ova at the cost of the
central cells, which serve them as a food supply. As these masses
grow and increase in number they rupture the walls of the ligament,
and escape into the body-cavity, in which they float. The ova are
fertilised whilst floating in the fluid of the body-cavity. The eggs
segment and the embryo is formed whilst still in the body of the
mother.

The embryos escape by means of a complicated apparatus the

details of which vary in the different species, but which, like many of
the organs in these animals, consists of very few cells with very large
nuclei. This apparatus consists of three parts: the bell, the uterus,
and the oviduct. The bell is a large funnel-shaped structure, which
opens into the body-cavity, and is connected with the end of the
ligament; near its lower end, where it is continuous with the uterus, is
a second smaller opening situated dorsally. By the contraction and
expansion of its lips the oval embryos are swallowed and pass on
through the uterus to the oviduct, which opens at the posterior end of
the body. If the bell takes in any of the less mature eggs which are
spherical in shape, they are passed back into the body-cavity
through the above-mentioned dorsal opening, and the same orifice
permits the passage of the spermatozoa even when the bell is full of
embryos.
 Fig. 97.—An egg of Echinorhynchus acus Rud. surrounded by three
egg-shells. Highly magnified. The egg has segmented, and the
cells are differentiated into a, the entoblast, and b, the ectoblast;
c, spines. (From Hamann.)

Embryology.—After fertilisation the egg surrounds itself with several

egg-shells, three of which are usually distinguished; the embryo is
already far advanced in its development by the time it leaves the
body of the mother and passes out into the alimentary canal of the
Vertebrate host. It leaves the body of this second host with the
faeces, and is eaten by the first or larval host, usually a small
Crustacean or water-insect, but in some cases a fish, within whose
alimentary canal it casts its membranes and becomes actively
mobile. By means of a ring of hooks developed round the anterior
end it bores its way through the wall of the alimentary canal, and
after some time—three weeks in E. proteus—comes to rest in the
body-cavity of its host. By this time most of the organs of the adult,
with the exception of the reproductive glands, are already well
established; the latter only attain maturity when the first host is eaten
by the second, and the larvae find themselves in the intestine of a
Vertebrate.
 Fig. 98.—A, A larval Echinorhynchus proteus Westrumb. further
developed than in Fig. 97. Highly magnified. The entoblast has
developed inside it the proboscis a; b, b, the giant nuclei of the
ectoblast. B, The entoblast at a more advanced stage, the
ectoblast is not shown. The outermost layer of cells will form the
muscles of the body-wall; the body-cavity has appeared; a,
proboscis; b, cerebral ganglion; c, body-cavity; d, d, the testes
beginning to appear in the ligament; e, cells which will form the
generative ducts.

Some of the details of the development are very remarkable, and a

short account of them may be given. The segmentation of the egg is
unequal; it results in the formation of a central biscuit-shaped mass
of small cells and a peripheral mass of larger cells; the former is
called by Hamann[218] the entoblast, the latter the ectoblast. From
the entoblast arise all the organs of the body but the sub-cuticle and
the associated lemnisci, which are formed from the ectoblast. The
latter has a remarkable history; the cells begin to break down and
lose their outlines, whilst their nuclei fuse together and form a small
number of giant nuclei, which lie scattered throughout the syncytium
thus formed. The syncytium surrounds the entoblast on all sides; by
this time the anteriorly-placed hooks have appeared; in E. proteus
there are ten of these, but the number is not the same in all species.
The syncytium is in a fluid state, with a few gigantic nuclei floating in
it; these now lose their spherical shape, and throwing out processes
become amoeboid; in this way they bud off small portions of their
substance, and from these the oval nuclei of the sub-cuticle and the
lemnisci arise. The rest of the syncytium hardens into the fibrillar
matrix of the sub-cuticle, leaving, however, scattered spaces which
form the sub-cuticular sinuses of the adult. An interesting feature of
N. clavaeceps and Arhynchus hemignathi is that the skin of the adult
retains the larval features, and it and the lemnisci consist of a
syncytium with a very few giant nuclei scattered through it. Hamann
counted only eight in the skin and two in each lemniscus in the
example figured on p. 178.

Fig. 99.—A, The larva of Echinorhynchus proteus from the body-cavity

of Phoxinus laevis, with the proboscis retracted and the whole still
enclosed in a capsule. B, A section through the same; a, the
invaginated proboscis; b, proboscis sheath; c, beginning of the
neck; d, lemniscus. Highly magnified. (Both from Hamann.)

The whole of the rest of the body is formed by the entoblast. Within
the latter a circular split arises which separates a single layer of
outermost cells from an axial strand of many cells (Fig. 98, B). The
split is the future body-cavity; the axial strand forms the proboscis, its
sheath, the cerebral ganglion, muscles, etc., and the ligament with
the contained generative organs; the outermost layer of cells forms
the muscular lining to the skin. It is interesting to note that these cells
destined to become muscle-fibres are at first arranged as a single
layer of cubical epithelial cells lining the body-cavity; most of them
become circular muscle-fibres, but a few are pushed inwards so as
to lie next the body-cavity, and these become the longitudinal fibres.

Classification.—Until recently the Acanthocephala were supposed

to include but one genus, Echinorhynchus, with several hundred
species, but Hamann[219] has pointed out that these species present
differences which enabled him to divide the group into three families,
each with a corresponding genus. To these I have ventured to add a
fourth family, to include a remarkable species, Arhynchus
hemignathi, described below. The characters of the first three
families in the account given below are taken from Hamann's paper.
 Fig. 100.—Fully formed larva of Echinorhynchus proteus from the
body-cavity of Phoxinus laevis. (From Hamann.) Highly magnified.
a, Proboscis; b, bulla; c, neck; d, trunk; e, e, lemnisci.

Family I. Echinorhynchidae.—The body is elongated and smooth.

The proboscis-sheath has a double wall, and the proboscis is
invaginated into it. The central nerve-ganglion lies in the middle line,
as a rule on the posterior blind end of the proboscis-sheath. The
papillae which bear the hooks are only covered with a chitinous cap
at their apex, and the hooks have a process below. This family is by
far the largest; a few species only can be mentioned.
Echinorhynchus proteus lives in its mature form in fishes; the young
forms, up to a centimetre in length, are found living freely in the
intestine of numerous fresh-water fishes. Those found in Gobio
fluviatilis, the gudgeon; Leuciscus virgo; Lota vulgaris, the burbot or
eel-pout; young trout; Thymallus vulgaris, the grayling, seldom
surpass this size, but those found in Acerina cernua, the pope fish; in
Abramis bipunctatus; in Esox lucius,the pike, and in older trout,
attain or surpass double the length. As the parasites grow older they
bury their proboscis and neck in the wall of the intestine, the inner
surface of which is studded with the orange-coloured bodies of the
parasites. The proboscis is so deeply sunk in the wall of the
alimentary canal as to form a papilla on its outer surface (Fig. 92).
The larvae of E. proteus are found in the body-cavity of Gammarus
pulex, one of the Amphipod Crustacea, and also in the same position
in numerous fresh-water fishes; they must have passed into this first
host by the mouth and alimentary canal. If the liver of an infested
minnow, Leuciscus phoxinus, be examined, it will be found to contain
on its surface numerous spherical or egg-shaped capsules of an
orange colour, 2 to 2.5 mm. in length; these contain the larval forms
of the parasite. They develop into the adult form when the first host
is eaten by a carnivorous fish, but a complication may take place
when the larval form is found in Gammarus, as the latter, the first
host, may be eaten by a fish (intermediate host) in which the larva
does not become mature, and only develops sexual organs when
eaten by a carnivorous fish (second host). The larval form is also
found in Nemachilus barbatulus, Gobio fluviatilis, and the
sticklebacks Gasterosteus aculeatus and G. pungitius.

E. clavula Duj. is found in Salmo fario, Abramis brama, Cyprinus

carpio, Gobius niger, Lepadogaster gouanii, etc.; E. linstowi Ham. in
Leuciscus idus, Abramis ballerus, Abramis bipunctatus, and
Acipenser huso; E. lutzii Ham. was found by Dr. Lutz in Brazil in the
intestine of Bufo agua; E. angustatus Rud. occurs in such numbers
in the perch, Perca fluviatilis, as to almost occlude the lumen of the
intestine, and one out of every three or four fish in certain districts is
infested by it. It is also found in the pike, Esox lucius, and the barbel,
Barbus vulgaris. The first or larval host of this species is the Isopod
Asellus aquaticus. E. moniliformis Brews. is stated to attain maturity
in the human intestine. Except for the fact that G. gigas has once
been observed in the same place, this is the only human parasite
amongst the Acanthocephala. Its normal second hosts are Mus
decumanus and Myoxus quercinus, and its first or larval host, the
larvae of the beetle Blaps mucronata. E. porrigens Rud. is found in
considerable numbers in the small intestine of a fin-whale
(Balaenoptera sibbaldii), and E. strumosus Rud., in the small
intestine of a seal (Phoca vitulina), and in the body-cavity of the
angler fish (Lophius piscatorius). E. acus is common in the whiting,
Gadus merlangus.

Family II. Gigantorhynchidae.—Large forms with ringed, flattened,

and Taenia-like bodies. The hook-papillae are covered all over with
transparent chitinous sheaths with two root-like processes. The
proboscis-sheath is muscular and without a lumen. The central
nervous system is excentrically placed below the middle of the so-
called sheath. The lemnisci are long twisted tubes with a central
canal.

Hamann places three species in this family: Gigantorhynchus

echinodiscus, G. spira, and G. taenioides; but as he points out that
E. gigas resembles these in its more important structural features, it
seems advisable to include it here under the name G. gigas. The
members of the first family often present a transversely ringed
appearance after death, but the Gigantorhynchidae are ringed when
alive, and the circular canals in the skin show a certain regularity,
being arranged one between each two rings. There is no lumen in
the proboscis-sheath, which is not attached to the boundary between
the proboscis and the trunk, but to the inner surface of the proboscis,
and the whole can be retracted within the anterior portion of the
body, which is invaginable. There are always eight cement-glands,
and other differences exist in the musculature, hooks, and position of
the nervous system.

G. gigas occurs in the adult state in the small intestine of swine; in

Europe its first or larval host is believed to be the grubs of
Melolontha vulgaris and Cetonia aurata, but these beetles are
absent from America, though the parasite infests American hogs.
Stiles[220] has recently made some experiments which tend to show
that in the United States the source of infection is some species of
the beetle Lachnosterna, and he has succeeded in infecting the grub
of L. arcuata by feeding it on the eggs of the parasite; from one larva
he took 300 parasites six weeks after feeding it. L. arcuata is, like M.
vulgaris, phytophagous, but the grubs of both the beetles are fond of
frequenting manure heaps and patches of dung, and thus are much
exposed to the dangers of infection.

G. echinodiscus inhabits the intestine of ant-eaters, having been

found in Myrmecophaga jubata and Cycloturus didactylus. G. spira
lives in the king vulture Sarcorhampus papa, and G. taenioides in
Dicholophus cristatus, a species of Cariama.
Family III. Neorhynchidae.—Sexual maturity is reached in the larval
stage. The proboscis-sheath has a single wall. A few giant nuclei
only are found in the sub-cuticle and in the lemnisci. The circular
muscle layer is very simply developed. The longitudinal muscle-cells
are only present in certain places.

This family includes two species, Neorhynchus clavaeceps and N.

agilis, which afford interesting examples of paedogenesis. The sub-
cuticle and the lemnisci are dominated by a few giant nuclei, which
remain in the embryonic state and do not break up into numerous
nuclei as in other forms. The musculature is but little developed and
the longitudinal sheath hardly exists. The proboscis-sheath consists
of a simple muscular layer, and the short proboscis has few hooks
and presents an embryonic appearance.

The sexually-mature form lives in the carp, Cyprinus carpio; the

larval form is found, according to Villot,[221] encysted in the fat
bodies of the larva of Sialis lutaria, one of the Neuroptera, and in the
alimentary canal of the leech Nephelis octocula, and successful
experiments have been made in infecting some species of the water
snail Limnaea. N. agilis occurs in Mugil auratus and M. cephalus.

Family IV. Arhynchidae.—Short forms with the body divided into

three well-marked regions—head, collar, and trunk. The head is
pitted, the collar smooth, and the trunk wrinkled, not annulated, in
spirit specimens. There is no eversible introvert, and no introvert
sheath and no hooks. The sub-cuticle and the lemnisci have a few
giant nuclei, and the lemnisci are long and coiled.[222]

This family resembles the Gigantorhynchidae in the length and

curvature of its lemnisci, and the Neorhynchidae in the persistence
of the embryonic condition of the nuclei in the sub-cuticle and the
lemnisci; but in the shape of the body, its division into three well-
marked regions, the absence of eversible proboscis, proboscis
sheath, and hooks it stands alone, though it is nearer to the
Neorhynchidae than to either of the other families.

The single species Arhynchus hemignathi was found attached to the

skin around the anus of a Sandwich Island bird, Hemignathus
proceros. The bird is a member of a family Drepanididae, which is
entirely confined to the Sandwich Island group. Professor Newton
tells me that it is probable that the "food of Hemignathus consists
entirely of insects which it finds in or under the bark of trees," hence
it is probable that the second host of this parasite, if such exists,
must be looked for amongst the Insecta.

CHAPTER VII

CHAETOGNATHA

STRUCTURE—REPRODUCTION—HABITS—FOOD—CLASSIFICATION TABLE
OF IDENTIFICATION

At certain seasons and at certain times of the day the naturalist who
is investigating the fauna of the surface of the sea is apt to find his
tow-net crammed with innumerable transparent spindle-shaped
animals, which by their number and the way in which they become
entangled with rarer objects, often render useless the result of his
labours. These animals belong to the class Chaetognatha, which
includes three genera, Sagitta, Spadella, and Krohnia. Amongst
them are divided about twenty species, some of which, however, are
of doubtful value.

Anatomy.—The body of these animals is as transparent as crystal; it

is elongated, and bears a resemblance to certain torpedos, except
that the head forms a somewhat blunt termination to the spindle-
shaped body. The tail bears a caudal fin, and Spadella and Krohnia
have a single pair, and Sagitta two pairs, of lateral fins; all of which
are flattened horizontally.

The body is externally divisible into three regions—head, trunk, and

tail—and these correspond with the arrangement of the internal
organs.

Fig. 101.—Sagitta bipunctata. a, Vesicula seminalis. × 4. (After

Hertwig.)

The head is surrounded by a fold of skin, forming a hood, which is

most prominent at the sides (Fig. 102, g); within the hood the head
bears from two to four rows of short spines, and outside these a right
and left row of sickle-shaped hooks, the free ends of which in a state
of rest converge round the mouth, but when disturbed these hooks
can be widely divaricated.

The cavity of the body, or coelom, is divided into three distinct

chambers by the presence of two thin transverse walls or septa, one
situated between the head and the trunk, the other between the
trunk and the tail (Figs. 104, 105). In the head, this cavity is much
reduced by the presence of special muscles which move the spines,
hooks, etc.; and in the small species, such as Spadella
cephaloptera, the other two cavities are almost entirely occupied by
the digestive and reproductive organs[223]; but in the large species,
e.g. Sagitta hexaptera, a considerable space is left between the
internal organs and the skin, and this is occupied by a coelomic fluid.
If the skin of one of these larger species be punctured the fluid
escapes and the animal shrivels up. A longitudinal partition or
mesentery, with numerous pores in it, runs through these spaces,
dividing the body-cavity into a right and left half; in the region of the
trunk this mesentery supports the alimentary canal.

In addition to certain muscles in the head, which move the hooks,

etc., there is a muscular lining to the body-wall. This is divided into
two dorsal and two ventral bands, much in the same way as in
Nematodes. The muscle fibres are striated.

The mouth, situated either terminally—Spadella marioni[224]—or

below the head, leads into a pharynx; this passes into an intestine
lined by a single layer of ciliated cells with a few glandular ones
intermingled. The intestine runs straight through the body without
loop or coil, and opens by an anus situated at the junction of the
trunk and the tail. In most cases the anus is ventral or on the lower
surface, but Gourret asserts that in Spadella marioni it is on the
upper surface.

There are no special respiratory, excretory, or circulatory organs,

unless a glandular structure described by Gourret in the head of
Spadella marioni be a real kidney.

The nervous system consists of a supra-oesophageal ganglion or

brain situated in the head, and of a ventral ganglion lying in the
trunk; both these nerve centres are embedded in the epidermis, and
are connected with one another by means of two stout peri-
oesophageal nerves (Figs. 102, 104). The brain also gives off a pair
of nerves to the eyes, another pair to the olfactory organ, and a pair
which ultimately meet one another and so form a ring; on this are
certain ganglia giving off nerves which supply the muscles of the
head. Both the chief ganglia give off numerous nerves, which divide
and split up into a network of fibres which permeate the whole skin.
The sense organs are comparatively simple. A pair of very small
eyes lie in the skin of the head; they are of complex structure, and to
some extent remind one of the simple eyes of certain Crustacea.
Behind the eyes and also on the upper surface of the animal is an
unpaired organ which is usually described as olfactory in function
(Figs. 103, 105). This is a ring-shaped modification of the epidermis
drawn out into different shapes in the various species. The modified
epidermal cells bear long cilia. The remaining sensory organs found
in the group consist of clumps of modified cells scattered in round
groups over the surface of the body and of the fins. The central cells
of each group bear long tactile hairs, and are surrounded by
supporting cells.

Fig. 102.—Head of Sagitta bipunctata. A, Dorsal view; B, ventral view.

× about 33. (From Hertwig.) A, a, spines; b, nerves to lateral
cephalic ganglia; c, hooks; d, cephalic ganglion; e, olfactory
nerve; f, optic nerve; g, hood; h, commissure to ventral ganglion; j,
olfactory organ: B, a, c, and g as in A; k, mouth.

The Chaetognatha are hermaphrodite, and carry the female organs

in the trunk, the male in the tail. In a mature specimen the two
ovaries occupy almost all the space in the trunk between the
alimentary canal and the skin, and each is supported by a narrow
lateral mesentery. The ovary is traversed by a oviduct which often
contains spermatozoa; it is not clear how the eggs make their way
into the oviduct, which seems to have no internal opening and to act
largely as a receptaculum seminis. The oviducts open externally on
the upper side at the base of the lateral fin, close to the junction of
the tail and the trunk.
The cavity of the tail is divided into two lateral chambers by the
extension backward of the median vertical mesentery. In each of
these a testis and a vas deferens are found. The testes are solid
ridges formed by the growth of the lining cells of this part of the
body-cavity; the cells mature into spermatozoa, which break off and
float freely in the coelomic fluid. At the breeding season the whole
tail may be crowded with masses of spermatozoa, which are kept in
a more or less regular circulation by the ciliated cells lining the body-
wall. The vas deferens opens internally into the space where the
spermatozoa lie, and at the other end into a vesicula seminis, which
opens to the exterior. The position of the latter structure varies, and
is of some systematic value.

The eggs are laid in the water and as a rule float at the surface of the
sea. Spadella cephaloptera is, however, an exception to this rule, as
it attaches its eggs by means of a gelatinous stalk to sea-weeds. The
segmentation of the ovum is regular, and gives rise to a two-layered
stage or gastrula, which opens by a pore, the blastopore. This does
not, however, become the mouth, but closes up and the mouth
arises at the opposite pole. Perhaps the most interesting feature of
the development of Sagitta is that the cells destined to form the
reproductive organs separate from the other cells of the embryo at a
very early date, whilst it is still in the gastrula stage. There is no
larval form, but the young hatch out from the egg in a state
resembling the adult in all respects but that of size.
 Fig. 103.—Spadella cephaloptera. Dorsal view. x 30. (From Hertwig.) a,
Cephalic ganglion; b, commissure to ventral ganglion; c, olfactory
organ; d, alimentary canal; e, ovary; f, oviduct; g, testis; h,
vesicula seminalis.

Habits.—The Chaetognatha are essentially pelagic, and resemble

many other creatures that dwell at the surface of the ocean in being
almost completely transparent. Most species have been taken far out
at sea, but some are perhaps rather more numerous near the coast,
and one species, Spadella cephaloptera, is littoral. They swim by
means of muscular movements of the whole body; the fins have no
movement of their own, and seem to serve as balancers, and not as
locomotory organs. Although usually found at the surface of the
water, many species have been taken at considerable depths.
Chun[225] states that they are found in countless numbers at depths
of from 100 metres to 1300 metres. The commonest species at
these depths are Sagitta hexaptera and Sagitta serratodentata.
Sagitta bipunctata, according to the same authority, confines itself to
the surface. Whether the change of depth is diurnal, or whether it
has any relation to sexual maturity, or to any other cause, has not
been satisfactorily determined.

The food of the Chaetognatha consists of floating diatoms, Infusoria,

small larvae, and such Copepods as Calanus finmarchicus, and
small Amphipods as Phoxus plumosus.[226] At times they also
devour small larval or post-larval fishes, and owing to their incredible
numbers, they doubtless do considerable damage to sea fisheries. It
is also recorded that they eat one another, and specimens have
been taken which have ingested the whole body of another Sagitta
except the head, which hangs out of the mouth of the eater, and
gives it the appearance of a double-headed monster.[227] It has been
said that they attack hydroid polypes, but here at any rate they do
not have it all their own way. Masterman[228] has figured the apical
group of five polypes of Obelia, three of which are engaged in
ingesting as many young Sagitta.
They exist in incredible numbers; Grassi describes the surface of the
sea at Messina on certain days as being literally covered with them,
and they must form the food supply of numerous animals which prey
upon the pelagic fauna. The immense number of individuals is
probably accounted for to some extent by the fact that they lay eggs
all the year round, and pass through a very short and rapid
development. They are not known to be phosphorescent.

Classification.—The features of the Chaetognatha which have most

systematic value are the size of the adult, the relations of the length
to the breadth, and of the three divisions to one another; the size,
number, and position of the lateral fins, and of the hooks and spines
on the head; the thickness of the epidermis, and the structure of the
olfactory organ; and, finally, the form of the reproductive organs.

Strodtmann,[229] who gives the latest and most complete account of

the species of Chaetognatha, arranges them under three genera,
which he characterises as follows:—

(i.) Sagitta Slabber.—Two pairs of lateral fins, two rows of spines on

the head. The lateral thickening of the epidermis absent or
insignificant.

Under this genus are included nine definite species and five others—
S. gracilis Verrill, S. elegans Verrill, S. darwini Grassi, S. diptera
d'Orbigny, and S. triptera d'Orbigny—whose position, owing to the
inadequacy of their description, is of doubtful validity.
 Fig. 104.—Sagitta hexaptera. Ventral view. × 4. (From Hertwig.) a,
Mouth; b, hooks; c, anterior septum; d, alimentary canal; e,
commissure from the brain to the ventral ganglion; f, ventral
ganglion; g, ovary; h, oviduct; i, posterior septum; j, testis; k,
vesicula seminalis.

The distribution of the other species may be mentioned. S.

hexaptera is the largest Chaetognath known, and reaches in the
adult stage a length of 7 cm. It is very widely distributed, being found
in practically all the temperate and warm seas, usually at the surface
of the water, though at times it is found at a depth of one metre, or
even deeper. S. lyra, Mediterranean, very rare. S. tricuspidata,
widely distributed. S. magna, Mediterranean and Madeiran, living at
the surface. S. bipunctata, the most frequently described form,
smaller than the preceding species, 1-2 cm. in length, widely
distributed, and as a rule living near the coast line. S. serratodentata,
Mediterranean. S. enflata, on the surface of the sea, Mediterranean
and Madeiran. S. minima, a very small species, 1 cm. in length,
Mediterranean. S. falcidens, Atlantic, off the coast of New Jersey.

(ii.) Krohnia Langerhans.—A single lateral fin extending on to both

trunk and tail segment, no lateral epidermal extensions behind the
head, only one row of spines on the head. Trunk longer than the tail.

Krohnia has but two species: K. hamata Möbius, with a length of 3-4
cm., found in the North Atlantic and at considerable depths, 200 to
300 fathoms; and K. subtilis Grassi, 1.5 cm. long, with an
extraordinary slender body and a relatively large head, found at
Messina, but very rare; as a rule only one specimen has been found
at a time.

(iii.) Spadella Langerhans.—A single pair of lateral fins; these are

situated on the tail segment. Behind the head a thickening of the
epidermis extends down each side of the body to the fin, or even
farther. Two rows of spines on the head. Small animals, not longer
than 1 cm.

Fig. 105.—Spadella draco. Dorsal view. × 12. (From Hertwig.) a,

Cephalic ganglion; b, commissure between the cephalic ganglion
and the ventral; c, eye; d, olfactory organ; e, alimentary canal; f,
ovary; g, oviduct (the line goes a little beyond the duct); h, testis; j,
vesicula seminalis.

S. cephaloptera Busch is the smallest species of Chaetognatha,

attaining at most a length of .5 cm. The body is not so transparent as
in other species, and is of a yellowish colour. It has been found from
the Orkney Islands to the Mediterranean. Strodtmann is of the
opinion that the three species S. mariana Lewes, S. batziana Giard,
and S. gallica Pagenstecher differ from the above-named only in
size, or that their description is too indefinite to permit of accurate
characterisation. He recognises three other distinct species: S.
pontica Uljanin, from the Black Sea; S. marioni Gourret, from the
Gulf of Lyons; and S. draco Krohn, Mediterranean and Madeiran,
and from the Canaries.

Much confusion has been introduced into the classification of the

Chaetognatha by Grassi,[230] who calls some—but not all—of what
other writers term Sagitta, Spadella, and vice versâ. The following
table was compiled by Strodtmann,[231] but I have incorporated in it
two species recently described from Amboyna by Béraneck,[232] and
called by him Sagitta bedoti and Spadella vougai respectively:—

CHAETOGNATHA

I. Two pairs of lateral fins; two rows of spines on the head;

slender forms.

(i.) Number of spines in posterior row greater than in anterior.

a. Border of hooks smooth, their point not curved.

α. No interval between the two fins on each side. 3.5 cm.

long; 4-7 anterior spines, 8-11 posterior spines; olfactory
organ lying entirely on the trunk. The anterior nerves of the
ventral ganglion lie close to one another as far as the
head.—Sagitta lyra.

β. A distinct interval between the two fins on each side.

aa. Adult animals large; hooks 6-7; anterior spines 3-4;

posterior spines 5-7; tail ¼ or ⅕ of the total length; lateral
areas relatively larger.—Sagitta hexaptera.

bb. Greatest length 1-2 cm.

 αα. Thickening of the epidermis behind the head;
prominently projecting vesiculae seminales; olfactory
organ very long; hooks 8-10; anterior spines 4-6;
posterior spines 10-15.—Sagitta bipunctata.

ββ. No epidermal thickening; two caeca on the anterior

end of intestine; length 1 cm.; hooks 6-9; anterior
spines 3-4; posterior spines 7-8; point of the hooks
somewhat bent round.—Sagitta minima.

γγ. Epidermis thin; no caeca; hooks 8-9, their ends not

bent; anterior spines 3-4; posterior spines 7-8; length 2
cm.; small head; trunk proportionately thick.—Sagitta
enflata.

δδ. Hooks 11-14, usually 12; length 1.8 cm.; anterior

spines 6-7; posterior spines 18.—Sagitta falcidens.

εε. Hooks 7 on each side; length 1.3 cm.; anterior

spines 8-10, posterior spines 18-22; no olfactory organ.
—Sagitta bedoti.

b. Edge of hooks toothed and their point bent round; hooks

6-8; anterior spines 6-8; posterior spines 10-12; length 1.5
cm.; slender; conspicuously projecting vesiculae seminales.
—Sagitta serratodentata.

(ii.) Number of the spines in posterior row smaller than in

anterior.

a. Anterior spines 3; posterior spine 1; hooks 8; length 3.5

cm.—Sagitta tricuspidata.