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 Essentials of
Microeconomics

Essentials of Microeconomics is an excellent introduction to microeconomics. It

presents the basic tools of microeconomics clearly and concisely. It presents a vigorous
treatment of all relevant introductory microeconomic concepts, and emphasises mod-
ern economics – game theory and imperfect markets. Each chapter is self-contained
and includes the required key mathematical skills at the start.
Now in its second edition, this updated textbook includes:
• Expanded lecturer resources, including detailed lecture slides, sample exam ques-
tions and updated test bank multiple choice questions
• An additional section on Economics in Practice, focused on policy, econometrics
and behavioural economics
This book is ideal not only for introductory microeconomics courses, but its level of
analysis also makes the book appropriate for introductory level economics taught at
postgraduate level. With the emphasis on strategy, this text is also well suited for use
in business economics courses.
Bonnie Nguyen is an Honours graduate from the University of Sydney in both Law and
Economics. Bonnie’s research interests include an economic analysis of litigation and
the internal organisation and ownership structure of firms. She has published research
in the Journal of Institutional and Theoretical Economics and the Australian Economic
Review. Bonnie has previously taught microeconomics at the University of Sydney.
Andrew Wait is a Professor in the School of Economics at the University of Sydney.
He has a PhD from the Australian National University and a Bachelor of Economics
(Honours) from the University of Adelaide. Andrew’s research interests include indus-
trial organisation and organisational economics. Andrew has published in the RAND
Journal of Economics, the International Journal of Industrial Organization and the
Journal of Law, Economics and Organization. He is the co-convenor of the Annual
Organizational Economics Workshop.
“One of the standout features of this textbook is its clear and concise writing style,
which effectively conveys complex economic principles in an accessible manner. The
authors have succeeded in presenting the material in a way that is engaging and easy
to understand for students with varying levels of prior knowledge.”
Kadir Atalay, Associate Professor,
University of Sydney
“Without the clutter that sometimes chocks introductory texts, this book allows stu-
dents to focus on the core intuition and analytical tools. This book presents the
essentials of microeconomics. It provides an excellent platform for further study in
economics. This textbook will stand the test of time. I recommend it to instructors
teaching any principles or introductory microeconomics course.”
Alexander Matros, Professor of Economics,
Moore School of Business, University of South Carolina
“Good economic analysis requires understanding of how to apply models to the world.
This book strikes the right balance between technique and economic relevance. The
authors lead students through just enough analytical tools to be able to understand and
then apply economic models to real economic problems. By focusing on the essentials
this book makes economics accessible to students from all backgrounds. Its strength
lies in the brevity and clarity of its exposition. At no point is there anything extraneous,
yet all is covered.”
Guillaume Roger, Associate Professor (Research),
Monash University
“The second edition improves on what was already an outstanding introductory micro-
economics textbook. A particular strength of the book is how the presentation of the
material highlights the power of economic models for helping us understand the world
around us. The new chapters on policy and applications show students how to apply
what they have learnt. There is nothing extraneous in this book. A mastery of this
content will provide students with almost all of the microeconomics tools of analysis
they will need for the rest of their academic and professional careers. I recommend this
for any introductory microeconomics or business economics course.”
Kieron Meagher, Professor,
Australian National University
“Economic thought is vital to understanding individuals and their interaction within
societies. This textbook provides a set of tools for students to understand markets and
analyze the decisions made by consumers, businesses and governments. The exposition
is concise and precise, presenting fundamental techniques and insights in a way that
is accessible to students new to the discipline, giving a good balance of mathematical
rigour and verbal explanation. I used the first edition to great success and will use this
new edition too. I highly recommend it.”
Jonathan Newton, Professor,
Kyoto University, Japan
 Essentials of
Microeconomics

Second Edition

Bonnie Nguyen and

Andrew Wait
Designed cover image: © Getty Images
Second edition published 2024
by Routledge
4 Park Square, Milton Park, Abingdon, Oxon, OX14 4RN
and by Routledge
605 Third Avenue, New York, NY 10158
Routledge is an imprint of the Taylor & Francis Group, an informa business
© 2024 Bonnie Nguyen and Andrew Wait
The right of Bonnie Nguyen and Andrew Wait to be identified as authors of this
work has been asserted in accordance with sections 77 and 78 of the Copyright,
Designs and Patents Act 1988.
All rights reserved. No part of this book may be reprinted or reproduced or utilised
in any form or by any electronic, mechanical, or other means, now known or
hereafter invented, including photocopying and recording, or in any information
storage or retrieval system, without permission in writing from the publishers.
Trademark notice: Product or corporate names may be trademarks or registered trademarks,
and are used only for identification and explanation without intent to infringe.
First edition published by Routledge 2016
British Library Cataloguing-in-Publication Data
A catalogue record for this book is available from the British Library
Library of Congress Cataloging-in-Publication Data
Names: Nguyen, Bonnie, author. | Wait, Andrew, author.
Title: Essentials of microeconomics / Bonnie Nguyen and Andrew Wait.
Description: Second edition. | Abingdon, Oxon ; New York, NY : Routledge, 2024. |
Includes bibliographical references and index. | Identifiers: LCCN 2023034859 (print) |
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ISBN 9781032453668 (paperback) | ISBN 9781003376644 (ebook) |
ISBN 9781032647289 (ebook other) | Subjects: LCSH: Microeconomics.
Classification: LCC HB172 .N446 2024 (print) | LCC HB172 (ebook) |
DDC 338.5–dc23/eng/20230727
LC record available at https://lccn.loc.gov/2023034859
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ISBN: 9781032453675 (hbk)
ISBN: 9781032453668 (pbk)
ISBN: 9781003376644 (ebk)
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DOI: 10.4324/9781003376644
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by Newgen Publishing UK
Access the Support Material: www.routledge.com/9781032453668
 Contents

List of illustrations xi

Part I Key concepts and tools 1

1 Key economic concepts 3

1.1 Introduction 3
1.2 Scarcity and opportunity cost 4
1.3 Marginal analysis 5
1.4 Ceteris paribus 5
1.5 Correlation and causation 6
1.6 Concluding comments 6
2 Key mathematical tools 9
2.1 Introduction 9
2.2 Equations 9
2.3 Differentiation 11
2.4 Elasticity 12
2.5 Simultaneous equations 12
2.6 Concluding comments 13
3 Key strategic tools 15
3.1 Introduction 15
3.2 The essentials of game theory 15
3.3 Simultaneous-move games 16
3.4 Some types of simultaneous-move games 20
3.5 Sequential games 22
3.6 Concluding comments 26

Part II Gains from trade 27

4 Trade and the PPF 29

4.1 Introduction 29
4.2 Gains from exchange 29
4.3 Gains from specialisation 30
4.4 Concluding comments 36
vi 0 Contents

5 Bargaining 39
5.1 Introduction 39
5.2 Bargaining and surplus 39
5.3 Take-it-or-leave-it negotiations 40
5.4 Multiple-offer bargaining 42
5.5 Some caveats 43
5.6 Concluding comments 44

Part III Market fundamentals 47

6 Demand 49
6.1 Introduction 49
6.2 Benefit and willingness to pay 49
6.3 Individual demand 50
6.4 Market demand 52
6.5 Concluding comments 53
7 Production and costs 55
7.1 Introduction 55
7.2 The short run and long run 55
7.3 Production 56
7.4 Short-run costs 59
7.5 Long-run costs 62
7.6 Total revenue, total cost and economic profit 63
7.7 Concluding comments 64
8 Supply 67
8.1 Introduction 67
8.2 Firm supply 67
8.3 Market supply 69
8.4 Concluding comments 70
9 Equilibrium and welfare 71
9.1 Introduction 71
9.2 Market equilibrium 71
9.3 Comparative static analysis 72
9.4 Welfare 74
9.5 Pareto efficiency 78
9.6 Concluding comments 79
10 Elasticity 81
10.1 Introduction 81
10.2 Measuring elasticity 81
Contents vii

10.3 Applications 83
10.4 Concluding comments 89

Part IV Types of markets 91

11 Introduction to markets 93
11.1 Introduction to the four types of markets 93
12 Perfect competition 95
12.1 Introduction 95
12.2 Characteristics of perfect competition 95
12.3 Supply in the short run 96
12.4 Supply in the long run 99
12.5 Market supply in the long run 102
12.6 Concluding comments 105
13 Monopoly 107
13.1 Introduction 107
13.2 Characteristics of a monopoly 107
13.3 The single-price monopolist 108
13.4 Welfare under the single-price monopolist 111
13.5 Price discrimination 114
13.6 Natural monopoly 120
13.7 Regulating a natural monopoly 121
13.8 Concluding comments 123
14 Monopolistic competition 125
14.1 Introduction 125
14.2 Characteristics of monopolistic competition 125
14.3 The short run 126
14.4 The long run 127
14.5 Welfare under monopolistic competition 129
14.6 Concluding comments 129
15 Oligopoly 131
15.1 Introduction 131
15.2 Characteristics of an oligopoly 132
15.3 Simultaneous-move games 132
15.4 Product choice: an application of a coordination game 135
15.5 Sequential games 137
15.6 Concluding comments 142
viii 0 Contents

Part V Market failures 143

16 Price regulation, taxes and subsidies 145

16.1 Introduction 145
16.2 Price regulation 145
16.3 Taxes and subsidies 149
16.4 Concluding comments 159
17 Externalities 161
17.1 Introduction 161
17.2 External costs and benefits 161
17.3 The problem with externalities 164
17.4 Solutions to externalities 167
17.5 Government solutions to externalities 169
17.6 Concluding comments 174
18 Public goods and common resources 175
18.1 Introduction 175
18.2 Public goods 175
18.3 Common resources and the Tragedy of the Commons 177
18.4 Concluding comments 178

Part VI International trade 179

19 International trade 181

19.1 Introduction 181
19.2 The welfare effects of international trade 181
19.3 Barriers to trade 185
19.4 Arguments against free trade 190
19.5 Concluding comments 191

Part VII Economics in practice 193

20 Some other markets 195

20.1 Introduction 195
20.2 The housing market 195
20.3 The labour market 198
20.4 The capital market 201
20.5 Concluding comments 204
Contents ix

21 Economic policy 205

21.1 Introduction 205
21.2 Economic policy in practice: some examples 205
21.3 The Theory of Second Best 207
21.4 The policy cycle 208
21.5 Concluding comments 209
22 Economics and evidence 211
22.1 Introduction 211
22.2 Econometrics and statistics 211
22.3 Experimental economics 212
22.4 Behavioural economics 213
22.5 Concluding comments 214

Part VIII Review 215

23 Questions & answers 217

Index 253
 Illustrations

Figures
3.1 The normal form of a game 17
3.2 A game with a dominant strategy equilibrium 17
3.3 A game with two Nash equilibria 18
3.4 The prisoner’s dilemma 20
3.5 The coordination game 21
3.6 Battle of the sexes 22
3.7 Stag hunt 22
3.8 Extensive form of a game 23
3.9 The normal form of the game depicted in Figure 3.8 23
3.10 Extensive form of a game in which Jay and Daisy organise a date 25
4.1 The production possibility frontier (PPF) traces out combinations
of the quantity of two goods (X and Y) that can be produced if all
resources are used 31
4.2 A production possibility frontier for goods X and Y. If there is a shock
that boosts the production of both goods, the PPF will shift outwards
from origin along both axes 32
4.3 A production possibility frontier for goods X and Y. If there is a shock
that boosts the production of X only, the PPF will shift outwards from
origin along the x-axis only 32
4.4 A production possibility frontier for goods X and Y. A one-unit
increase in the production of X is less costly when the output of X is
low (e.g. from A to A′ ) than when the output of X is high (e.g. from B
to B′ ) 33
4.5 Michelle and Rodney’s PPFs. When Michelle and Rodney cannot
trade, their consumption points are given by the point A. When
Michelle and Rodney can trade with each other, their consumption
points are given by the point B. Note that point B lies outside each
person’s individual PPF 36
5.1 Negotiation in which A makes a single offer of p 40
5.2 Negotiation in which B makes a single offer of p 41
5.3 Negotiation in which A makes an offer of p1 and B makes a
counter-offer of p2 42
6.1 A typical marginal benefit curve 50
xii 0 Illustrations

6.2 An individual consumer’s demand curve is given by his or her marginal

benefit curve. A movement along the demand curve is known as a
‘change in the quantity demanded’ 51
6.3 A movement of the demand curve itself is called a ‘change in demand’ 52
6.4 The market demand curve (DM ) can be derived by summing
horizontally the individual demand curves (D1 and D2 ). In the first
graph, both individual demand curves have the same P-intercept. If
this is the case, then for every price p, the quantity demanded by
the market will be the sum of individual consumer demand (that is,
q1 + q2 ). In the second graph, the individual demand curves have
different P-intercepts. Above the price p̄, the quantity demanded by
individual 2 will be zero; therefore, in this range, the market demand
curve follows the individual demand curve for consumer 1. Below
the price p̄, the quantity demanded by both consumers is positive; in
this range, the quantity demanded by the market will be the sum of
individual consumer demand (that is, q1 + q2 ) 53
7.1 A typical short-run production function, where L represents the
amount of labour employed and q(L) represents the level of output 56
7.2 A typical total cost function, where q represents the level of output
and TC represents total cost 59
7.3 The typical shape of the average total cost curve, the average fixed
cost curve, the average variable cost curve and the marginal cost curve 61
7.4 The long-run average cost curve can be obtained by taking the
lower envelope of all short-run average cost curves. In this diagram,
the short-run average cost curves are represented by the unbroken
short-run average cost (SRAC) curves. The long-run average cost
curve is the dashed line that runs beneath them. As more short-run
average cost curves are drawn in, the long-run average cost curve will
become smoother 62
7.5 When the LRAC curve is downward sloping, the firm is experiencing
economies of scale; when the long-run average cost (LRAC) curve
is upward-sloping, there are diseconomies of scale. When the LRAC
curve is flat, there are constant average costs 63
8.1 An individual firm’s supply curve is given by its marginal cost curve.
A shift along the supply curve is known as a ‘change in the quantity
supplied’ 68
8.2 A movement of a firm’s supply curve itself is called a ‘change in
supply’ 69
8.3 The market supply curve (SM ) can be derived by summing horizontally
the individual supply curves (S1 and S2 ). In the first graph, both
individual supply curves have the same P-intercept. If this is the case,
then for every price p, the quantity supplied by the market will be the
sum of individual firm supply (that is, q1 + q2 ). In the second graph,
the individual supply curves have different P-intercepts. Below the
price p̄, the quantity supplied by firm 2 will be zero; therefore, in this
range, the market supply curve follows the individual supply curve
for firm 1. Above the price p̄, the quantity supplied by both firms is
positive; in this range, the quantity supplied by the market will be the
sum of individual firm supply (that is, q1 + q2 ) 70
Illustrations xiii

9.1 A market in equilibrium. The equilibrium price is P* and the

equilibrium quantity traded is Q* 72
9.2 When market price is above the equilibrium price, there is an excess
of supply in the market 72
9.3 When market price is below the equilibrium price, there is an excess
of demand in the market 73
9.4 The market for corn chips when there is an increase in the price of
potato chips 74
9.5 The market for cars when there is an increase in the price of steel 74
9.6 The area of consumer surplus in this market is denoted by the
grey-shaded area 75
9.7 When the market price falls from P1 to P2 , the area of consumer
surplus increases from A to A+B+C. The area B represents the
increase in consumer surplus that arises from an increase in the
net benefit of previously consumed units. The area C represents
the increase in consumer surplus arising from the consumption of
additional units 76
9.8 The area of producer surplus in this market is denoted by the
grey-shaded area 77
9.9 When the market price increases from P1 to P2 , the area of producer
surplus increases from A to A+B+C. The area B represents the
increase in producer surplus that arises from an increase in the net
benefit of selling units that would have been sold previously. The area
C represents the increase in producer surplus arising from the sale of
additional units 77
9.10 The area of total surplus in this market is denoted by the grey-shaded
area 78
10.1 When demand is perfectly inelastic, the demand curve is vertical (as
seen in the left panel). When demand is perfectly elastic, the demand
curve is horizontal (as seen in the right panel) 84
10.2 When the demand curve is linear, elasticity changes as we move along
the curve. The curve is most elastic near the P-axis and most inelastic
near the Q-axis 85
10.3 Total revenue can be calculated by multiplying price and quantity. In
this diagram, the size of total revenue is denoted by the grey-shaded
area 86
10.4 When supply is perfectly inelastic, the supply curve is vertical (as seen
in the left panel). When supply is perfectly elastic, the supply curve is
horizontal (as seen in the right panel) 87
12.1 The short-run supply curve of a firm is traced out by the part of the
MC curve that lies above AVC. In this diagram, it is denoted by the
black dashed line 97
12.2 A firm in a perfectly competitive market, making a profit. The
grey-shaded area represents the size of that profit 98
12.3 A firm in a perfectly competitive market, making a loss. The
grey-shaded area represents the size of that loss 98
xiv 0 Illustrations

12.4 The long-run supply curve of a firm is traced out by the part of the
MC curve that lies above ATC. In this diagram, it is denoted by the
black dashed line 100
12.5 When the market price is above average total cost, firms in the market
are making profits. This will encourage entry into the market, shifting
the supply curve right from S to S′ . In turn, this will put downward
pressure on market prices 101
12.6 When the market price is below average total cost, firms in the market
are making losses. This will encourage exit from the market, shifting
the supply curve left from S to S′ . In turn, this will put upward pressure
on market prices 101
12.7 In the long run, there are zero profits in a perfectly competitive market.
This requires P = ATC min . Because there are zero profits, there is no
incentive for any firm in the market to exit the market and there is no
incentive for any additional firms to enter the market. In this diagram,
the long-run equilibrium price is P* , the quantity traded in the market
is Q* and the output of the firm is q* 101
12.8 In the long run, free entry and exit means that the price in a
constant-cost industry will always be driven back to ATCmin . The
long-run market supply curve LRS is perfectly elastic at P = ATCmin 102
12.9 Following an unanticipated increase in demand, in the short run
price rises and firms in the industry make positive economic profits.
However, in the long run, entry forces prices back down to the
P* = ATCmin . Each firm sells q* units and economic profits are zero 103
12.10 In an increasing-cost industry, the long-run industry supply curve is
upward sloping 104
12.11 In a decreasing-cost industry, the long-run industry supply curve is
downward sloping 105
13.1 When the demand curve is linear, the marginal revenue curve has the
same vertical intercept and twice the slope of the demand curve 109
13.2 A profit-maximising monopolist sets MR = MC, and thus produces
quantity Qm . At this quantity, the market price will be Pm 110
13.3 The monopolist’s profit is given by π = Q(P − ATC ). The area
corresponding to the monopolist’s profit is shaded grey in this diagram 111
13.4 The quantity traded in the market is lower under monopolist than in a
competitive market 112
13.5 Consumer surplus and producer surplus under a monopoly 112
13.6 Total surplus and deadweight loss under a monopoly 113
13.7 When the monopolist engages in first-degree price discrimination, the
efficient quantity is traded in the market. However, all surplus in the
market is producer surplus 115
13.8 When the monopolist uses a two-part tariff, it charges a fixed fee (F)
equal to the size of the lighter grey-shaded triangle. It then charges a
per-unit fee (p) for each unit consumed equal to MC. The total revenue
from the per-unit fee is represented by the darker grey-shaded area.
The total revenue of the monopolist overall is represented by all of the
shaded areas 116
Illustrations xv

13.9 When a monopolist engages in third-degree price discrimination (with

constant MCs), it maximises profit where MRA = MRB = MC. The
price is higher in the market where demand is relatively inelastic; here
PA > PB 118
13.10 When a firm has a fixed cost and a constant marginal cost, the average
total cost curve will be downward sloping for all values of Q; this
industry will be a natural monopoly 120
13.11 Under marginal-cost price regulation, the government sets the
monopoly price at P = MC (assuming constant MC for simplicity).
However, this means that the monopolist makes a loss equal to the
grey-shaded area (that is, its fixed costs). The government will need to
subsidise the monopolist that amount to prevent them from leaving the
market 122
13.12 Under average-cost price regulation, the government sets the monopoly
price at P = ATC. However, the monopolist will produce less than the
efficient quantity, so there is still some dead weight loss equal to the
size of the grey-shaded area. However, note that regulation decreases
the area of deadweight loss, relative to no regulation at all 122
14.1 A firm in a monopolistically competitive market in the short run. The
firm maximises its profits by setting MR = MC. Thus, the firm sells
qn units of its product, at a price of Pn 127
14.2 A firm in a monopolistically competitive market in the long run.
Because of the entry and/or exit of other firms in the market, this
firm’s demand curve (and marginal revenue curve) has shifted such
that, at the quantity associated with MR = MC, the firm is making
zero profits 128
15.1 Price-setting game in an oligopoly 132
15.2 Advertising game in an oligopoly. In this figure, ‘A’ represents the
choice to advertise and ‘NA’ represents the choice to not advertise 133
15.3 Location-choice game in an oligopoly 135
15.4 Platform choice for game developers 136
15.5 Entry game in an oligopoly 138
15.6 Simultaneous technological choice 139
15.7 Technological choice game with a first-mover advantage 139
15.8 Natural monopoly entry game 140
15.9 Free-riding game 141
15.10 A ‘matching pennies’ game 142
16.1 A price floor set at P̄ 146
16.2 The welfare effects of a price floor set at P̄ 147
16.3 A price ceiling set at P̄ 147
16.4 The welfare effects of a price ceiling set at P̄ 148
16.5 A tax on consumers causes the demand curve to shift downwards by
the size of the tax 150
16.6 A tax on consumers creates a new market price and quantity at
(Qt , Pt ). However, the total amount paid by consumers is Pt + t 150
16.7 A tax on producers causes the supply curve to shift upwards by the
size of the tax 151
xvi 0 Illustrations

16.8 A tax on producers creates a new market price and quantity at (Qt , Pt ).
However, the net amount received by producers is Pt − t 151
16.9 The welfare effects of a tax 152
16.10 A tax on consumers in the market for tomatoes. The legal incidence
of the tax is borne by consumers, but the economic incidence of the
tax is shared. The price that consumers pay increases from P* to Pt + t,
whereas producers receive Pt following the introduction of the tax,
rather than P* 153
16.11 If demand is perfectly inelastic, when a tax of t per unit is instituted,
consumers pay for all of the tax; following the introduction of the tax
consumers pay P* + t whereas suppliers continue to receive P* 154
16.12 When a tax of t per unit is implemented and supply is perfectly elastic,
consumers pay for all of the tax; following the introduction of the tax,
consumers pay P* + t whereas suppliers continue to receive P* 155
16.13 The deadweight loss generated by a tax depends on the elasticity
of supply and demand. In each panel the demand curve and the tax
implemented are the same. There is a larger DWL generated in the
market with relative elastic supply (the left-hand panel) as compared
with the DWL in the market in which supply is relatively inelastic (the
right-hand panel) 156
16.14 A subsidy for consumption creates a new market price and quantity at
(Qs , Ps ). The total amount paid by consumers is Ps − s 157
16.15 A subsidy for production creates a new market price and quantity at
(Qs , Ps ). The total amount received by producers is Ps + s 158
16.16 Consumer surplus and producer surplus as a result of a subsidy 159
16.17 Government revenue and deadweight loss as a result of a subsidy 159
17.1 The relationship between MPB and MSB in the presence of a positive
externality 162
17.2 The relationship between MPC and MSC in the presence of a negative
externality 163
17.3 The market equilibrium and the socially optimal outcome in the
presence of a positive externality. The area representing deadweight
loss is shaded in grey 165
17.4 The market equilibrium and the socially optimal outcome in the
presence of a negative externality. The area representing deadweight
loss is shaded in grey 165
17.5 The market equilibrium and the socially optimal outcome in the
presence of a negative consumption externality. The area representing
deadweight loss is shaded in grey 166
17.6 The market equilibrium and the socially optimal outcome in the
presence of a positive production externality. The area representing
deadweight loss is shaded in grey 167
17.7 A subsidy granted to the consumer and to the producer, used to address
a positive externality 170
17.8 A tax imposed on the consumer and on the producer, used to address a
negative externality 171
10 2 Key concepts and tools

When graphing this equation, we might want to know where the line intersects the
axes. Along the y-axis, x takes the value of zero, so we can solve for the y-intercept
by setting x = 0 (in equation 2.1, the y-intercept is y = c). Similarly, y takes the value
of zero along the x-axis, so setting y = 0 yields the x-intercept (in equation 2.1, the
x-intercept is x = − mc ).

Example. Suppose we are interested in the relationship between the price of a

good (P) and the quantity demanded of that good (Q). We might represent this
relationship using the equation P = 100 − 2Q. Here, Q and P take the places of x
and y, respectively. The slope of the line is −2 (see the next section). The line cuts
the P-axis at 100 and the Q-axis at 50.

2.2.2 The slope of a straight line

Note that, in equation 2.1, if we increase x by 1 unit, y must increase by m units. Thus,
the parameter m determines how a change in x affects a change in y (recall that this
relationship is the focus of marginal analysis – see Chapter 1). The parameter m is also
known as the slope or the gradient of the line; it tells us how steep the line is. That is,
if we graph equation 2.1 with x and y on the horizontal and vertical axes, respectively,
for every 1 unit we move horizontally (along the x-axis) we must also move m units
vertically (along the y-axis). Note that if an increase in x results in an increase in y, the
slope is positive. Conversely, if the increase in x results in a fall in y, the slope of the
line (by definition) is negative. It is the fact that the slope is constant that makes this
the equation of a straight line.

2.2.3 Determining the equation of a straight line

When we are not explicitly given the equation of a straight line, we can determine that
equation from any two points on the line. First, we need to determine the slope, m.
Recall that the slope is simply the change in y for a one-unit change in x. Therefore, if
we have two points on a straight line, (x1 , y1 ) and (x2 , y2 ), the formula for the slope m
is2 :
∆y y2 − y1
m= = (2.2)
∆x x2 − x1
In other words, the slope of the line m is the change in y (∆y) given a change in x (∆x).
As we have calculated the slope m, it can be substituted into the general equation of a
straight line, y = mx + c, so as to find c. To do this, we use the fact that the equation
must pass through the point (x1 , y1 ), so we can substitute these values into our equation
to determine c.

Example. Suppose a line passes through (4, 3) and (5, 1). Applying equation 2.2,
we can find the slope, m = 34−
−5
1
= −2. Recall that the general equation of a straight
line is y = mx + c. Substituting in m = −2 and (4, 3) yields 3 = −2 × 4 + c, so
c = 11. Therefore, the equation of the line is y = −2x + 11.
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Otoconcha is peculiar. The fresh-water Mollusca, besides the Isidora
characteristic of the sub-region, are partly related to New Caledonia
through the occurrence of Melanopsis, partly to Tasmania through
Potamopyrgus, while the peculiar Latia is possibly akin to
Gundlachia (Tasmania). The land operculates number only 5 genera
and 14 species in all, excluding a doubtful Diplommatina.[374]
Land and Fresh-water Mollusca of the Neozealanian Province
Schizoglossa 1
Paryphanta 5
Rhytida 6
Rhenea 2
Helicarion 1
Otoconcha 1
Microcystis 1
Trochonanina 1
Phacussa 3
Thalassohelix 5
Gerontia 2
Allodiscus 10
Pyrrha 1
Therasia 7
Phenacohelix 3
Suteria 1
Flammulina 13
Laoma 23
Endodonta 10
Charopa 28
Placostylus 1
Carthaea 1
Tornatellina 1
Janella 3
Latia 2
Ancylus 2
Limnaea 5
Amphipeplea 2
Planorbis 1
Isidora 7
Melanopsis 2
Potamopyrgus 4
Paxillus 1
 Lagochilus 7
Omphalotropis 1
Realia 4
Hydrocena 1
Unio 9
Sphaerium 1
Pisidium 2
Lord Howe’s I. is remarkable as containing a Placostylus, which
thus links the island with this province. The remainder of the fauna is
Polynesian, with the exception of a species (common to the Fijis) of
Parmella, a slug akin to Helicarion, Parmacochlea, and Cystopelta.
(3) The Polynesian Sub-region includes all the island groups of
the central and southern Pacific (except those classified in the
Papuan and Australian sub-regions), from the Pelews and Carolines
in the west to the Marquesas and Paumotus in the east, and from
the Tonga group in the south to the Sandwich Is. in the north. It may
be subdivided into (a) the Polynesian province proper, and (b) the
Hawaiian province, which includes the Sandwich Is. only.

Fig. 216.—Characteristic
Polynesian Mollusca: A,
Achatinella vulpina Fér.,
Sandwich Is.; B, Partula
planilabrum Pease, Society
Is.
(a) The general features of the Polynesian province are very
similar throughout, although the Mollusca of each island group are in
the main peculiar. The species are mostly small and obscure. Helix
scarcely occurs, its place being taken by small Zonitidae
(Microcystis, Charopa, Trochomorpha, etc.), and by groups of so-
called Patula (Endodonta, Pitys, etc.), the exact position of which is
not yet settled. Libera, remarkable for its method of ovipositing (p.
128), is peculiar to the Society and Hervey Is.; Partula is almost
universal, attaining its maximum (40 sp.) in the Society Is.;
Tornatellina, Pupa, and Vertigo occur throughout.
The land operculates consist chiefly of Omphalotropis, Pupina,
Realia, and Helicina. Diplommatina and Palaina are abundant on the
Pelews, and a Moussonia occurs in the Samoa Is. Ostodes, a small
form of Cyclophorus, is found in some of the southern groups. The
fresh-water operculates are Melania, Neritina (including Clithon, a
sub-genus furnished with spines), and Navicella; there are no
Unionidae, while fresh-water Pulmonata are very scarce.
(b) The land Mollusca of the Hawaiian province are distinguished
by the possession of four entirely peculiar genera—Achatinella,
Leptachatina, Carelia, and Auriculella. More than 300 of the two
former genera have been described, every mountain valley of some
of the islands having its own peculiar species. The destruction of the
indigenous herbage by goats is rapidly extinguishing many forms.
Partula, and the small land operculates, so characteristic of the other
groups, are, with the exception of Helicina, entirely wanting. The
occurrence of one of the Merope group of Helix (Solomon Is.) is
remarkable, and there is a rich development of Succinea. “Patula,”
Microcystis, Tornatellina, and the other small Polynesian land
Pulmonata are well represented. The presence of Isidora, absent
from the central Pacific groups, is remarkable, and Erinna is a
peculiar genus belonging to the Limnaeidae.
 CHAPTER XI
GEOGRAPHICAL DISTRIBUTION OF LAND MOLLUSCA (continued)—THE
ETHIOPIAN, NEARCTIC, AND NEOTROPICAL REGIONS

D. The Ethiopian Region

The Ethiopian region includes the whole of Africa south of the
Great Desert, and Southern Arabia, together with the outlying
islands, excepting those of the Atlantidean province (p. 297).
Regarded as a whole, the Ethiopian is poorest in land Mollusca of
all the tropical regions. And yet its characteristics are very
remarkable. The entire Achatina group is peculiar, and takes,
especially in W. Africa, some curious forms (Columna, Perideris,
Pseudachatina). Carnivorous Mollusca (Ennea, Gibbus, etc.) are
highly developed, especially in the south and east, the largest known
helicoid form (Aerope) being from Natal. In the possession of these
types of the Agnatha, Africa is more closely related to the
Australasian than to the Oriental region. The true Cyclostoma are
entirely peculiar to the region, but are absent from West Africa.
Fresh-water Mollusca are abundant and characteristic, especially
in and near the Great Lakes. Lanistes, Cleopatra, and Meladomus,
among the operculates, together with Mutela and Aetheria
(Unionidae), Galatea and Fischeria (Cyrenidae), are peculiar.
In its negative, as well as its positive features, the Ethiopian
region is markedly isolated. Helicidae and Naninidae are equally
deficient, the former, indeed, attaining some numerical
predominance in the extreme south, but the species are nearly all
insignificant in size and colouring. It is only in Madagascar that Helix
asserts itself. Arion, Limax, Hyalinia, Clausilia, and a number of other
genera abundant along the Mediterranean, are either altogether
absent, or are very scantily represented. Land operculates, so
characteristic of other tropical countries, are almost entirely wanting.
If we disregard the Malagasy sub-region, there are scarcely forty
species of land operculates on the whole African continent.
 The Ethiopian region may be divided into three sub-regions: (1)
the Central African; (2) the South African; (3) the Malagasy.
(1) The Central African Sub-region is bounded on the north by
the Great Desert, on the east and west by the ocean, and on the
south by a line roughly drawn between the mouth of the Orange
River and Delagoa Bay; it also includes S. Arabia. No natural
features exist which tend to break up this vast district into areas of
independent zoological development. The absence of long and lofty
mountain ranges, the enormous size of the great river basins, and
the general uniformity of climate, equalise the conditions of life
throughout. It will be convenient to break the sub-region up into
provinces, but in most cases no precise line of demarcation can be
laid down.
(a) The Senegambian Province may be regarded as extending
from the mouth of the Senegal River to Cape Palmas. Only 8 genera
of land Mollusca are known, including 4 Limicolaria and 3 Thapsia,
with 1 small Cyclophorus. Fresh-water genera are abundant, and
include most of the characteristic Ethiopian forms.
(b) The West African Province extends from Cape Palmas to the
mouth of the Congo, and is rich in Mollusca. The great Achatina,
largest of land snails, whose shell sometimes attains a length of 6½
in., Limicolaria, Perideris, and Pseudachatina are the characteristic
forms. The Agnatha are represented by Ennea, Streptaxis, and
Streptostele. Rachis and Pachnodus, sub-genera of Buliminus, occur
also on the east coast. A special feature is the development of
several peculiar slug-like genera, e.g. Oopelta, perhaps a form of
Arion; Estria, a slug with an external shell, akin to Parmacella; and
Aspidelus, a form intermediate between Helicarion and Limax.
Claviger, a handsome group akin to Cerithium, is peculiar to the
estuaries of West African rivers.
About sixteen species are known from the Cameroons District, but
no peculiar genera occur. The French Congo District has not yet
been well explored. Tomostele, a genus allied to Streptostele, is
peculiar, and Pseudachatina attains its maximum.
 Fig. 217.—Columna
flammea Müll., Princes I.
St. Thomas and Princes Is., in the Gulf of Guinea, are well known.
Princes I. has 22 species, 14 peculiar, and 2 common to St. Thomas
only, one of the latter being the great sinistral Achatina bicarinata
Chem. The remarkable genus Columna (Fig. 217) is peculiar, and
Streptostele (4 sp.) attains its maximum. Peculiar to St. Thomas are
Pyrgina, a turreted form of Stenogyra; Thyrophorella, a sinistral form
of Zonites; and Atopocochlis, a large bulimoid shell, whose true
relationships are not yet known. Homorus, a group of Achatina with
an elongated spire, occurring also in the Angola District and on the
east coast, has 4 species. No fresh-water species have as yet been
discovered in either of the islands.
The Angola and Benguela District, extending from the Congo to
the Cunene R., probably belongs to the West African Sub-region, but
until its fauna is better known it is advisable to consider it apart.
Achatina continues abundant, but the other characteristic West
African forms (Pseudachatina, Streptostele, Perideris) diminish or
are absent altogether. No Helix and only 1 Cyclophorus occur.
Ovampo, Damara, and Great Namaqualand, lying between the
Cumene and Orange rivers, seem to form a transition district
between the West and South African faunas. Helix reappears, while
the characteristic West African genera are almost entirely wanting.
 (c) The East African Province extends from about Delagoa Bay to
the Abyssinian shores of the Red Sea. In general out-line the
province consists of a flat marshy district, extending inland for many
miles from the sea; this is succeeded by rising ground, which
eventually becomes a high table-land, often desolate and arid,
whose line of slope lies parallel to the trend of the coast. The
Mollusca are little known, and have only been studied in isolated
districts, usually from the discoveries of exploring expeditions.
The Mozambique District, from Delagoa Bay to Cape Delgado,
includes no genus which does not occur on the west coast, except
Cyclostoma (2 sp.). Trochonanina (4 sp.), Urocyclus, a characteristic
African slug (2 sp.), Rachis (6 sp.), Pachnodus (2 sp.), and Achatina
(5 sp.), are the principal groups.

Fig. 218.—Urocyclus comorensis Fisch., Comoro Is.:

G, Generative orifice; M, mucous gland; O,
orifice leading to internal shell; P, pulmonary
orifice; T, tentacles. (After Fischer.)
The Zanzibar District, from Cape Delgado to the Somali country,
has the same general features. Meladomus, a large sinistral
Ampullaria, is characteristic, while Cyclostoma (5 sp.) becomes more
abundant. Helix is still absent, but the carnivorous forms (Streptaxis
2 sp., Ennea 7 sp.) are rather numerous.
The Somali District is characterised by operculate groups of the
Otopoma type (Georgia, Rochebrunia, Revoilia) whose generic value
is rather doubtful. Petraeus, in an Arabian type, supplants Rachis
and Pachnodus. Achatina is nearly wanting, but Limicolaria has 9
species. A few Helix, said to be of the Pisana group, occur.
 The District between the Great Lakes and the coast region is fairly
well known through recent explorations, especially those associated
with Emin Pasha. Streptaxis (6 sp.) and Ennea (24 sp.) are
numerous, Helix is wanting, and the Naninidae are represented by
Trochonanina (7 sp.), and other forms at present grouped under
Nanina or Hyalinia. On the high ground Buliminus, Cerastus, and
Hapalus replace, to some extent, the Achatina and Limicolaria of the
marshy plains. Land operculates (Cyclophorus 1, Cyclostoma 8) are
more numerous; among fresh-water genera we have Lanistes (5
sp.), Cleopatra (3 sp.), Meladomus (1 sp.), and Leroya, a sinistral
form with the facies of a Littorina. The characteristic African bivalves
(Mutela, Spatha, etc.) are few in number.
(d) Province of the Great Lakes.—The Mollusca of the four great
lakes of Eastern Central Africa—Lakes Albert Nyanza (Luta Nzige,
2720 ft.), Victoria Nyanza (Oukéréwé, 3700 ft.), Nyassa (1520 ft.),
and Tanganyika (2800 ft.)—are well known, and supply an
interesting problem in distribution. Those of the three first mentioned
lakes differ in no way from the rest of tropical Africa, but the Mollusca
of Tanganyika include, in addition to the ordinary African element, a
number of peculiar operculate genera, belonging principally to the
Melaniidae and Hydrobiidae. Several of these possess a solidity of
form and compactness of structure which is unusual in fresh-water
genera, and has led to the belief, among some authorities, that they
are the direct descendants of marine species, and that Tanganyika
represents an ancient marine area. This view appears untenable.
The Victoria Nyanza and Nyassa are part of the same system as
Tanganyika, and it is not easy to see how, if Tanganyika were once
an arm of the sea, they were not equally so, especially as they are
several hundred miles nearer the Indian Ocean as at present
defined. Nor, as will be seen from the figures given above, is there
anything in the altitudes which would make us expect anything
exceptional in Tanganyika. The similar case of L. Baikal must be
compared (p. 290), where again a number of specialised forms of
Hydrobia occur.
Of the genera concerned, Paramelania and Nassopsis are forms
of Melaniidae; Tiphobia (Fig. 219), which is allied to Paludomus, is a
compact shell with angulated spinose whorls; Lacunopsis,
Ponsonbya, Limnotrochus, and Tanganyicia are probably forms of
Lithoglyphus, some, as their names denote, being of decidedly
marine facies; Syrnolopsis and Turbonilla (?) look like
Pyramidellidae, Horea and Reymondia like Rissoina; Bourguignatia
appears to belong to Vivipara, with which has now been merged the
genus Neothauma. Recently discovered forms from the adjacent L.
Mweru are evidently of kindred origin.
(e) The Afro-Arabian Province includes Abyssinia, with S. Arabia,
the African shores of the Gulf of Aden, and Socotra. The province
contains a singular mixture of types. The high ground of Abyssinia
stands like a lofty European island in the midst of a tropical plain,
with Palaearctic genera flourishing like hardy northern plants on a
mountain in low latitudes. Helix, Vitrina, and Pupa abound, with a
few Clausilia and even a Limax. On the lower levels occur
Limicolaria (3 sp.), Subulina (7 sp.), Helicarion, and Homorus, but
land operculates are entirely wanting. Characteristic of the province
as a whole are various forms of Buliminus, which in Socotra are
represented by two peculiar sub-genera, Achatinelloides and
Passamaiella. In S. Arabia the mixture of types produces curious
results: the Helix, Clausilia, and Vitrina being Palaearctic, the
Limicolaria and all the operculates Ethiopian, while the single
Trochomorpha is Indian. Indian influence, indeed, comes out
unmistakably throughout the province. Thus in Socotra there are two
Cyclotopsis, in Abyssinia two Africarion (closely related to the Indian
Girasia), two Microcystis, and a Glessula, and in the Scioa district
there is a Sitala. The fresh-water Mollusca of Socotra are Indian
forms.
 Fig. 219.—Tiphobia Horei E.
A. Smith, L. Tanganyika.

Fig. 220.—Mollusca
characteristic of L.
Tanganyika: A, Nassopsis
nassa Woodw.; B, Spekia
zonata Woodw.; C,
Syrnolopsis lacustris E. A.
Smith.
 Fig. 221.—Achatina zebra
Lam., S. Africa. × ½.
(2) The South African Sub-region.—The principal characteristic
of the Mollusca of S. Africa is the occurrence of numerous small
species of Helicidae, belonging chiefly to the groups Pella, Phasis,
Dorcasia, and Sculptaria, all of which are practically peculiar.
Carnivorous genera are also prominent, Ennea here attaining its
maximum. Rhytida (to which several species still regarded as Pella
belong) is common only to the S. Pacific and Australasia, and forms,
with Isidora among the fresh-water pulmonates, a remarkable link of
connexion. Aerope, the largest of all helicoid carnivorous genera,
and Chlamydephorus, a carnivorous slug with an internal shell, are
peculiar. Achatina is still abundant, but Limicolaria is wanting.
Livinhacea, a form with a continuous peristome, perhaps akin to
Bulimus; Apera, a form of slug; and Coeliaxis, a genus perhaps akin
to the Papuan and Queensland Perrieria, are all peculiar. The land
operculates, which are not numerous, are of the East African type.
Land Mollusca of the S. African Sub-region
Chlamydephorus 1
Ennea 31
Aerope 5
Rhytida 3
Helicarion 3
Trochonanina 1
Trochozonites 1
Limax 1
Apera 1
 Vitrina 7
Nanina 6
Conulus 2
Patula 2
Pella 44
Dorcasia 8
Phasis 1
Sculptaria 2
Helix (inc. sed.) 4
Rachis 1
Pachnodus 3
Buliminus (?) 4
Pupa 20
Vertigo 2
Achatina 18
Livinhacea 1
Stenogyra 4
Coeliaxis 1
Succinea 3
Vaginula 2
Cyclophorus 1
Cyclostoma 7
Cyclotus (?) 1
Blanfordia 1
St. Helena.—The Molluscan fauna of St. Helena is perhaps the
most puzzling, as regards its geographical affinities, of any in the
world. It consists of 29 peculiar species of land Mollusca (fresh-water
species being unknown), 19 of which are recently extinct, partly
owing to the destruction of the forest, but are found in considerable
abundance in a state of good preservation.[375] The genera are—
Hyalinia 1
Patula 4 (3 extinct)
Endodonta 10 (7 extinct)
Bulimulus 7 (5 extinct)
Pachyotus 1 (extinct)
Tomigerus (?) 1 (extinct)
Pupa 2 (extinct)
Succinea 3
The 5 genera which concentrate our attention are Patula,
Endodonta, Pachyotus (Fig. 222), Tomigerus, and Bulimulus, all of
which appear utterly strange to an oceanic island in the middle of the
S. Atlantic. Patula and Endodonta are essentially Polynesian forms,
occurring abundantly on all the island groups in the Central Pacific.
Pachyotus, Tomigerus (assuming its correct identification), and
Bulimulus are all S. American forms, the two former being especially
characteristic of Brazil. How this mixture of genera now confined to
regions so widely distant, not only from St. Helena itself, but from
one another, became associated here, is a problem obviously not
easy of solution. The fauna is probably a remnant of a very ancient
type, possibly at one time much more widely distributed. Endodonta
(an essentially insular form, like Omphalotropis) actually occurs on
Fernando Noronha, off the Brazil coast, and we shall see how an
Indian and even a Polynesian element is present off the eastern
coasts of Africa.
Ascension I.—One indigenous species, a so-called Limax, is all
that has ever been discovered.
(3) The Malagasy Sub-region includes Madagascar with its
attendant satellites Bourbon, Mauritius, and Rodriguez, and the
Seychelles and Comoro groups. No land Mollusca are known from
the Amirantes, the Chagos, or from Aldabra. The special
characteristics of the sub-region are the great development of the
carnivorous land Mollusca (Ennea, Gibbus), the occurrence of a
considerable number of true Helicidae of great size and beauty, and
the prominence of the genus Cyclostoma.
(a) The Madagascan Province.—The land Mollusca of
Madagascar, although as yet imperfectly known, possess a striking
individuality. Two of the chief characteristics of the Ethiopian region
are the paucity of its land operculate and of its Helix fauna;
Madagascar is especially distinguished by the rich development of
both these groups. For size, colouring, and beauty of shape, the
Helicidae of the two sub-genera Ampelita and Helicophanta rival, if
they do not surpass, any in the world. They are quite peculiar to this
sub-region, not a trace of them occurring on the Mascarenes,
Seychelles, or even on the Comoros. Helicophanta is distinguished
by the enormous size of its embryonic shell, which persists in the
adult (Fig. 223), and in this respect the group appears to be related
to Acavus (Ceylon, Fig. 204) and Panda (N.E. Australia). As is usual
when Helix is well developed, Nanina (about 12 sp.) is
proportionately scanty.
The African Bulimini (Pachnodus and Rachis) are represented by
two species, but Achatina, so abundant on the mainland, is scarce.
Two other groups of Buliminus, Leucotaenia and Clavator, are
peculiar. The presence of a single Kaliella, specifically identical with
a common Indian form, is very remarkable.
Cyclostoma proper, of which Madagascar is the metropolis, is
richly developed (54 sp.). Many of the species are of great size and
of striking beauty of ornamentation. Unlike its Helicidae, this genus is
not restricted to Madagascar; several species occur on the mainland,
6 on the Comoros, one on the Seychelles, and 16 in Mauritius. The
sub-genera Acroptychia and Hainesia are peculiar.

Fig. 222.—Pachyotus auris

vulpina Desh., St. Helena (sub-
fossil).
 Fig. 223.—Helix (Helicophanta)
Souverbiana Fisch.,
Madagascar, showing
embryonic shell. × ⅔.

Fig. 224.—Cyclostoma
campanulatum Pfr., Madagascar.
The fresh-water Mollusca of Madagascar contain further traces of
Indian relationship. Thus we find two species of Paludomus, a genus
whose metropolis is Ceylon, India, and Further India, and which is
barely represented on the Seychelles and in the Somali district.
Melanatria, which is peculiar to Madagascar, has its nearest affinities
in the Cingalese and East Indian faunas. Several of the Melania and
the two Bithynia are of a type entirely wanting in Africa, but common
in the Indo-Malay sub-region. Not a single one of the characteristic
African fresh-water bivalves (Mutela, Spatha, Aetheria, Galatea, etc.)
has been found in Madagascar. On the other hand, certain African
Gasteropoda, such as Cleopatra and Isidora, occur, indicating, in
common with the land Mollusca, that an ultimate land connexion with
Africa must have taken place, but at an immeasurably remote period.
Land and Fresh-water Mollusca of Madagascar
Ennea 9
Urocyclus 2
Helicarion (?) 1
Macrocyclis (?) 1
Kaliella 1
Nanina (inc. sed.) 9
Ampelita 35
Helicophanta 17
Pachnodus 2
Rachis 2
Leucotaenia 2
Clavator 2
Achatina 3
Opeas 2
Subulina 3
Vaginula 4
Limnea 2
Planorbis 3
Isidora 3
Melania 7
Melanatria 4
Paludomus 2
Vivipara 1
Bithynia 2
Cleopatra 2
Ampullaria 6
Cyclophorus 2
Cyclotus (?) 1
Cyclostoma 54
Otopoma 5
Lithidion 1
Acroptychia 3
Hainesia 3
Unio 1
Corbicula 2
Sphaerium 1
Pisidium 1
 The Comoro Islands.—This isolated group possesses about 100
species, almost all of which are peculiar. The principal feature is the
rich development of Ennea (30 sp.). On the whole the group shows
more relationship to Madagascar than to the mainland. Thus we
have six species of true Cyclostoma, and only one Achatina, while
among the fresh-water genera is Septaria, which is characteristic of
the whole Malagasy Sub-region, but is absent from the mainland.
The Helicidae are all of insignificant size. Peculiar to the group is the
remarkable genus Cyclosurus (Fig. 152, p. 247).
(b) The Mascarene Province (Mauritius, Bourbon, Rodriguez, and
the Seychelles).—The percentage of peculiar species, which is very
high, can only be paralleled in the case of some of the West Indian
islands, and sufficiently attests the extreme isolation of the group
from Madagascar. We have—
Total sp. Land sp. Fresh-water Peculiar Peculiar to
sp. group
Mauritius 113 104 9 78 102 (90 p.c.)
Bourbon 45 40 5 19 38 (84 p.c.)
Rodriguez 23 19 4 15 21 (95 p.c.)
Seychelles 34 27 7 24 30 (90 p.c.)
The Mollusca of the group exhibit three distinct elements, the
Indigenous, the Madagascan, and the Indian and Australasian.
The genus Pachystyla (Naninidae) is quite peculiar, forming the
main portion of the land snails proper. It attains its maximum in
Mauritius (17 sp.), with 5 sp. in Bourbon and one sub-fossil sp. in
Rodriguez, while in the Seychelles it is absent. But the principal
feature of the Mascarene group is the extraordinary development of
the carnivorous genus Gibbus, which has 27 sp. in Mauritius, 8 in
Bourbon, 4 in Rodriguez; in the Seychelles, it is replaced by
Edentulina and Streptostele. The principal link with Madagascar is
found in a part of the operculate land fauna. Cyclostoma is present
(with Otopoma) in several fine living forms, and the number of sub-
fossil species is a clear indication that this group was, not long ago,
much more abundant, for of the 16 Cyclostoma known from
Mauritius 10 are sub-fossil. The operculates form a decided feature
of the land fauna; thus in Mauritius there are 32 species, or more
than 28 per cent of the whole.

Fig. 225.—Characteristic Mauritian land

shells: A, Gibbus palanga Fér.; A´,
young of same; B, Gibbus
lyonetianus Pall.
Indian and Australasian affinities are unmistakably present. Thus
Omphalotropis, a genus characteristic of small islands, is profusely
represented, but it does not occur in Madagascar or Africa. Two
Helicina (Mauritius and Seychelles) and a single Leptopoma
(possibly a Leptopomoides) are also of eastern relationship.
Cyclotopsis, Cyathopoma, and Geostilbia are markedly Indian
genera. Microcystis, Patula, and Tornatellina are Polynesian.
Hyalimax—and this is a very striking fact—occurs nowhere else but
in the Andamans and Nicobars, and on the Aracan coast. The
nearest relation to the Seychelles Mariaella appears to be the
Cingalese Tennentia. Not a single representative of these eleven
genera has been found even in Madagascar.
The fresh-water Mollusca (omitting the Neritidae) are: Mauritius 9
species, Bourbon 5, Rodriguez 4, Seychelles 6, with only 15 species
in all. The one Planorbis and the Vivipara, the Paludomus and two of
the Melania are of Indian types. The Lantzia (peculiar to Bourbon) is
probably allied to the Indian Camptonyx. Owing to the paucity of
permanent streams, no fresh-water bivalves occur. Among the
Neritidae is a single Septaria, a genus which, though occurring in
Madagascar, is entirely strange to Africa, and is abundant in the
Oriental and Australasian regions.
It would seem probable that when the closer connexion which at
one time undoubtedly existed between India and Eastern Africa
began to be less continuous,[376] the Mascarene group was first
severed from what ultimately became Madagascar, while the
Seychelles, and perhaps the Comoros, still continued united to it.
The Comoros, which lack the great Helices, separated off from
Madagascar first, while the Seychelles continued in more or less
direct union with that island sufficiently long to receive the
progenitors of Stylodonta (a peculiar group of Helix), but became
disunited at an exceedingly remote period.

E. The Nearctic Region

The southern boundary of this region may be regarded as roughly

corresponding to that of the United States, i.e. Lower California and
Mexico are excluded. The southern portion of Florida belongs to the
Antillean sub-region.
The principal characteristic of the Nearctic Region is the
remarkable poverty of its land Mollusca. No district in the world of
equal extent is so poor in genera, while those which occur are
generally of small size, with scarcely anything remarkable either in
colouring or form. The elongated land shells (Clausilia, Buliminus),
so characteristic of Europe, are entirely wanting, but a few
Bulimulus, of Neotropical origin, penetrate Texas, and from the same
sources come a few species of Glandina (as far north as S.
Carolina), Holospira (Texas), and Helicina.
The region falls into two well-marked sub-regions, the N.
American and the Californian, with the Rocky Mountain district as a
sort of debatable ground between them. The Californian sub-region
consists of the narrow strip of country between the Sierra Nevada,
the Cascade Mountains and the coast-line, from San Diego to
Alaska; the N. American sub-region consists of the remainder of the
region.
(1) The N. American Sub-region.—The Carnivorous genera are
represented solely by the few Glandina mentioned above, and by the
indigenous genus Selenites, a form midway between Testacella and
Limax, whose metropolis is on the Pacific slope, but which spreads
eastward into the Antilles. Among the Limacidae, Limax is common
to both sub-regions, but Tebennophorus (4 sp., 3 of which belong to
the genus Pallifera), a genus found also in China and Siam, and
Vitrinozonites do not occur in the Californian. Hyalinia (Zonites) is
fairly abundant, especially in the groups Mesomphix and
Gastrodonta (peculiar to this sub-region), and Hyalinia proper. Patula
is well represented. The Helicidae belong principally to the groups
Mesodon, Stenotrema, Triodopsis, Polygyra, and Strobila, only 6 of
which, out of a total of 84, reach the Pacific slope. Land operculates
are conspicuous for their almost complete absence (see map,
frontispiece).

Fig. 226.—Characteristic North American Mollusca. A,

Helix (Mesodon) palliata Say, Ohio. B, Helix
(Polygyra) cereolus Mühlf., Texas. C, Patula
alternata Say, Tennessee.
The poverty of the land fauna is atoned for by the extraordinary
abundance and variety of the fresh-water genera. A family of
operculates, the Pleuroceridae, with 10 genera and about 450
species, is quite peculiar, a few stragglers only reaching Central
America and the Antilles. The nucleus of their distribution is the
Upper Tennessee River with its branches, and the Coosa River.
They appear to dislike the neighbourhood of the sea, and are never