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Biogeography
SCIENCES
Ecosystems and Environment
Field Directors – Françoise Gaill and Dominique Joly
Biodiversity, Subject Head – Fabienne Aujard
Biogeography
An Integrative Approach of the
Evolution of Living
Coordinated by
Eric Guilbert
First published 2021 in Great Britain and the United States by ISTE Ltd and John Wiley & Sons, Inc.
Apart from any fair dealing for the purposes of research or private study, or criticism or review, as
permitted under the Copyright, Designs and Patents Act 1988, this publication may only be reproduced,
stored or transmitted, in any form or by any means, with the prior permission in writing of the publishers,
or in the case of reprographic reproduction in accordance with the terms and licenses issued by the
CLA. Enquiries concerning reproduction outside these terms should be sent to the publishers at the
undermentioned address:
www.iste.co.uk www.wiley.com
ERC code:
PE10 Earth System Science
PE10_13 Physical geography
LS8 Ecology, Evolution and Environmental Biology
LS8_1 Ecosystem and community ecology, macroecology
Contents
Preface . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . xi
Eric GUILBERT
2.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 27
2.2. From narrative dispersal accounts to event-based methods (EBM) . . . . . . . 27
2.2.1. Parsimony-based tree fitting . . . . . . . . . . . . . . . . . . . . . . . . . . 29
2.2.2. Dispersal–vicariance analysis . . . . . . . . . . . . . . . . . . . . . . . . . 31
2.3. From parsimony-based to semiparametric approaches . . . . . . . . . . . . . . 34
2.4. A new revolution: parametric approaches in biogeography . . . . . . . . . . . 38
2.4.1. Ancestral range versus single state models: DEC and BIB . . . . . . . . . 41
2.4.2. Extending the DEC and BIB models . . . . . . . . . . . . . . . . . . . . . 47
vi Biogeography
Chapter 3. Phylogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
Inessa VOET and Violaine NICOLAS
3.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 59
3.2. The early days of phylogeography: cytoplasmic genomes and qualitative
post hoc explanations of historical processes . . . . . . . . . . . . . . . . . . . . . . 61
3.3. Statistical phylogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 63
3.4. Comparative phylogeography . . . . . . . . . . . . . . . . . . . . . . . . . . . 67
3.5. Integrative studies . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 69
3.5.1. Integration of ecological niche modeling in phylogeographic studies . . . 69
3.5.2. Integration of life-history traits in phylogeographic studies . . . . . . . . . 73
3.6. Conclusion . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
3.7. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 76
4.1. Introduction . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 81
4.2. Geophysical biogeography at large. . . . . . . . . . . . . . . . . . . . . . . . . 82
4.2.1. Present day . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 82
4.2.2. The dynamic Earth: continental drift . . . . . . . . . . . . . . . . . . . . . 84
4.2.3. Continental drift and climate . . . . . . . . . . . . . . . . . . . . . . . . . . 87
4.2.4. The fast pace of mass extinctions . . . . . . . . . . . . . . . . . . . . . . . 90
4.3. Geophysical biogeography at regional scale. . . . . . . . . . . . . . . . . . . . 92
4.3.1. Mountain belts and rifts . . . . . . . . . . . . . . . . . . . . . . . . . . . . 95
4.3.2. Epeirogenies, dynamic topography . . . . . . . . . . . . . . . . . . . . . . 99
4.3.3. Glacial cycles . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 100
4.4. Conclusions . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 104
4.5. References . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 105
Index. . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . . 337
Preface
Eric GUILBERT
UMR7179 MECADEV, National Museum of Natural History, Paris, France
I am pretty sure that most of the scientists working on evolution or the ecology
of living organisms did not start as biogeographers … However, when interested in
understanding how living organisms have evolved, and how they are organized in
relation to their environment, linked to biotic and abiotic variables, biologists
naturally arrive at biogeography. Biogeography is the main approach when
embracing the history of living species as a whole. It is a huge world, and it is only
growing. After a quick search online (Lavoisier.fr), I found 1,291 books on
Biogeography written since 1985. Another search, on the Web of Science
(wcs.webofknowledge.com), shows 36,567 papers (written between 1957 and 2021)
with “biogeography” in the title (Figure P.1). Among these, 25% are associated with
“ecology” and 19% with “evolutionary biology”. However, not only ecology and
evolutionary biology are linked to biogeography, but a wide range of disciplines,
from geography (9%) to genetics (8%) or conservation (7%).
Biogeography,
coordinated by Eric GUILBERT.
© ISTE Ltd 2021.
xii Biogeography
of topics. The future of species is one of the major interests, with the history of
species distribution.
Not only do approaches differ according to the biology of organisms but they
also differ according to the environment. Dealing with the biogeography of
freshwater fish is not the same as dealing with the biogeography of marine fish. If
freshwater habitats can be considered as islands of water in the middle of the land
(see Chapter 8), oceans are much less fragmented and more stable environments (see
Preface xiii
Chapter 9). How should bacterial distribution in the soil be considered? Drivers of
community assemblages of bacteria are specific (Fierer et al. 2007). In addition,
taxonomic recognition of bacteria involves molecular tools (see Chapter 7). Is the
biogeography of water beetles studied in the same way as the biogeography of cave
beetles? The drivers of their distribution may not be the same (see, for example,
Arribas et al. 2012; Faille et al. 2014), even if both can be considered as in an
insular environment.
A lot of very good books on biogeography have been edited; see, for example,
the fifth edition of Biogeography (Lomolino et al. 2017), or Conservation
Biogeography (Ladle et al. 2011). Most provide the very bases of biogeography,
theories and methods, a wide range of approaches, historical and original cases with
nice illustrations. And yet, new studies and methodological novelties are coming out
every year, and the number and variety make biogeography so attractive and
exciting!
water in the land, while the other is open areas, where terrestrial approaches are not
always applicable. Finally, we will focus on particular approaches that are
challenging today and may be of greater importance in the future, such as the
biogeography of diseases (Chapter 11), a very current field of research, climate
change (Chapter 12) and conservation (Chapter 13), two fields that are also closely
linked to the human impact on the distribution of species in space and time.
Biogeography is not only a discipline that has been questioning the evolution of
species and ecology since naturalists started exploring the world. Understanding
patterns and processes of the distribution of species in space and time may provide
solutions to the challenges humanity has faced since the era called the Anthropocene
and its consequences, such as the biodiversity crisis and global warming.
July 2021
P.1. References
Arribas, P., Velasco, J., Abellan, P., Sanchez-Fernandez, D., Andujar, C., Calosi, P.,
Millan, A., Ribera, I., Bilton, D.T. (2012). Dispersal ability rather than ecological
tolerance drives differences in range size between lentic and lotic water beetles
(Coleoptera: Hydrophilidae). Journal of Biogeography, 39, 984–994.
Cook, L.G. and Crisp, M.D. (2005). Not so ancient: The extant crown group of Nothofagus
represents a post-Gondwanan radiation. Proceedings of the Royal Society B: Biological
Sciences, 272(1580), 2535–2544.
Dawson, M.N., Axmacher, J.C., Beierkuhnlein, C., Blois, J., Bradley, B.A., Cord, A.F.,
Dengler, J., He, K.A., Heaney, L.R., Jansson, R., Mahecha, M.D., Myers, C.,
Nogués-Bravo, D., Papadopoulou, A., Reu, B., Rodríguez-Sánchez, F., Steinbauer, M.J.,
Stigall, A., Tuanmu, M.-N., Gavin, D.G. (2016). A second horizon scan of biogeography:
Golden Ages, Midas touches, and the Red Queen. Frontiers of Biogeography, 8(4), 1–30.
Faille, A., Andújar, C., Fadrique, F., Ribera, I. (2014). Late Miocene origin of a
Ibero-Maghrebian clade of ground beetles with multiple colonisations of the subterranean
environment. Journal of Biogeography, 41, 1979–1990.
Fierer, N., Bradford, M.A., Jackson, R.B. (2007). Toward an ecological classification of soil
bacteria. Ecology, 88(6), 1354–1364.
Hill, R.S., Jordan, G.J., Macphail, M.K. (2015). Why we should retain Nothofagus sensu lato.
Australian Systematic Botany, 28(3), 190.
von Humboldt, A. and Bonpland, A. (1805). Essai sur la géographie des plantes ;
accompagné d’un tableau physique des régions équinocoxiales. Levrault, Schoell & Co.,
Paris.
Preface xv
Ladle, R.J. and Whittaker, R.J. (2011). Conservation Biogeography. Wiley-Blackwell Press,
Oxford.
Lomolino, M.V., Riddle, B.R., Whittaker, R.J. (2017). Biogeography, 5th edition.
Oxford University Press, Sunderland, MA.
MacArthur, R.H. and Wilson, E.O. (1967). The Theory of Island Biogeography. Princeton
University Press, Princeton, NJ.
Pourrut, X., Kumulungui, B., Wittmann, T., Moussavou, G., Délicat, A., Yaba, P., Nkoghe, D.,
González, J.-P., Leroy, E.M. (2005). The natural history of Ebola virus in Africa.
Microbes and Invection. 7, 1005–1014.
Wallace, A.R. (1876). The Geographical Distribution of Animals: With a Study of the
Relations of Living and Extinct Faunas as Elucidating the Past Changes of the Earth’s
Surface. Harper & Brothers, New York, NY.
1
Origins of Biogeography:
A Personal Perspective
Malte C. EBACH
University of New South Wales and The Australian Museum, Sydney, Australia
In the 1810 Preface of his Theory of Colours, Johann Wolfgang von Goethe
wrote, “The history of an individual displays his [or her] character, so it may here be
well affirmed that the history of science is science itself” (Goethe, cited in Duck and
Petry 2016, p. xxxv). What Goethe meant is that the past practices of scientists are
instances of scientific practice regardless of age. In other words, scientists do not
need historians of science to interpret past scientific practice. Scientific practice, no
matter how old, will always remain a part of science.
Where, then, does history fit in? Many historians and philosophers of science
discuss scientific ideas rather than practice. The reason is that historians and
philosophers of science do not engage in scientific practice and are therefore not
always able to interpret what we do due to a lack of training or experience or both.
British biologist Peter Medawar discussed this in 1968: “What scientists do has
never been the subject of a scientific, that is, ethological inquiry… It is no use
looking to scientific ‘papers’, for they not merely conceal but actively misrepresent
the reasoning that goes into the work they describe” (Medawar 1968, p. 151).
Can historians and philosophers of science trust what scientists say as opposed to
what they do? Understanding what scientists do is perhaps a much better way to
Biogeography,
coordinated by Eric GUILBERT.
© ISTE Ltd 2021.
Biogeography: An Integrative Approach of the Evolution of Living,
First Edition. Eric Guilbert.
© ISTE Ltd 2021. Published by ISTE Ltd and John Wiley & Sons, Inc.
2 Biogeography
understand the scientific process, rather than believing what they say. Take the
science of systematics or taxonomy as an example. Edgar Anderson noted that it is
“difficult to write about the taxonomic method [because] in its broadest aspects it
has never been described. Taxonomists are more like artists than like art critics; they
practice their trade and don’t discuss it” (Anderson, cited in Haas 1954, p. 65).
Indeed. What of biogeographical practice? In the case of taxonomy, the result is a
list of diagnostic characteristics, names and photographic plates. In biogeography,
the results are usually maps. Maps are representative of classifications and are an
ideal starting point to understand the early biogeographic method. Before we look at
the 18th and 19th biogeographical practices, it is important to understand what
biogeography is and how it may be defined1.
1.2.1. Terminology
The goal of finding a phyto- and zoogeographical classification united 18th and
19th century plant and animal geographies. It is important to note that the term
1. For a detailed account of the history of 18th and 19th century biogeographies, see Ebach (2015).
Origins of Biogeography: A Personal Perspective 3
biogeography only appeared in the latter half of the 19th century; Jordan (1883)
coined it in German, and Merriam (1892) in English; however, neither author
defined the term. The above OED definition of biogeography may have first
appeared in Nelson (1978), 90 years after it was first coined2. Regardless, it would
be historically incorrect to use the term biogeography for any theory, aims or
methods used by 18th and 19th century plant and animal geographers. The term
biogeography gained popularity in the latter half of the 20th century. I will use the
terms plant and animal geographies, botanical geography, and phytogeography and
zoogeography to suit the parlance of the time.
Earth
Southern Hemisphere
Australia
2. I avoid using the phrase “Father of ”. Coining a term does not justify ownership of the
whole field. Herman Jordan and Clinton Hart Merriam used the term only once. Jordan
casually refers to the term as though it was already in use, and Merriam uses it to describe his
“Bio-geographic map”.
4 Biogeography
There was also an understanding of a basic classification with the poison ivy:
Plant
Ivy
Green
Old World
Europe
Africa
Asia
New World
North America
South America
Australasia
Holarctic kingdom
Nearctic region
Palearctic region
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“Hullo, Mark! What ages you have been!” exclaimed his cousin.
“We can make room at this corner—come along, old man.”
Mark and his companion found themselves posted at the two
corners at the end of the table, and were for the moment the
cynosure of all eyes.
In a few seconds, as soon as the newcomers had been looked
after and given the scraps, the party continued their interrupted
conversation with redoubled animation. They all appeared to know
one another intimately. Captain Waring had evidently fallen among
old friends. They discussed people and places—to which the others
were strangers—and Mrs. Bellett was particularly animated, and
laughed incessantly—chiefly at her own remarks.
“And so Lalla Paske is going to her Aunt Ida? I thought Ida
Langrishe hated girls. I wonder if she will be able to manage her
niece, and what sort of a chaperon she will make?”
“A splendid one, I should say,” responded a man in a suit like a
five-barred gate—“on the principle of set a thief to catch a thief.”
“And old Mother Brande, up at Shirani, is expecting a niece too.
What fun it will be! What rivalry between her and Ida! What husband-
hunting, and scheming, and match-making! It will be as good as one
of Oscar Wilde’s plays. I am rather sorry that I shall not be there to
see. I shall get people to write to me—you for one, Captain Waring,”
and she nodded at him graciously.
Mark noticed his companion, who had been drinking water
(deluded girl—railway station water), put down her glass hastily, and
fix her eyes on Mrs. Bellett. No one could call her pale now.
“I wonder what Mrs. Brande’s niece will be like?” drawled her
sister. “I wonder if she, like her aunt, has been in domestic service.
He, he, he!” she giggled affectedly.
There was a general laugh, in the midst of which a clear treble
voice was heard—
“If you particularly wish to know, I can answer that question.” It
was the pale girl who was speaking.
Mrs. Coote simply glared, too astounded to utter a syllable.
“I was not aware that my aunt had ever been in domestic service;
but I can relieve you at once of all anxiety about myself. I have never
been in any situation, and this is the nearest approach I have ever
made to the servants’ hall!”
If the lamp in front of them had suddenly exploded, there could
scarcely have been more general consternation. Mrs. Bellett gasped
like a newly-landed fish; Captain Waring, purple with suppressed
laughter, was vainly cudgelling his brain for some suitable and
soothing remark, when the door was flung back by the guard,
bawling—
“Take your seats—take your seats, please, passengers by the
Cawnpore mail.”
Undoubtedly the train had never arrived at a more propitious
moment. The company rose with one consent, thrust back their
chairs, snatched up their parcels, and hurried precipitately out of the
room, leaving Honor and her escort vis-à-vis and all alone.
“If those are specimens of Englishwomen in India,” she exclaimed,
“give me the society of the natives; that dear old creature in the hut
was far more of a lady.”
“Oh, you must not judge by Mrs. Bellett! I am sure she must be
unique. I have never seen any one like her, so far,” he remarked
consolingly.
“I told you,” becoming calmer, and rising as she spoke, “that I
could not hold my tongue. I can not keep quiet. You see I have lost
no time—I have begun already. Of course, the proper thing for me to
have done would have been to sit still and make no remark, instead
of hurling a bombshell into the enemy’s camp. I have disgraced
myself and you; they will say, ‘Evil communications corrupt good
manners.’ I can easily find a carriage. Ah, here is my treasure of a
chuprassi. You have been extremely kind; but your friends are
waiting for you, and really you had better not be seen with me any
longer.”
She was very tall; and when she drew herself up their eyes were
nearly on a level. She looked straight at him, and held out her hand
with a somewhat forced smile.
He smiled also as he replied, “I consider it an honour to be seen
with you, under any circumstances, and I shall certainly see you off.
Our train is not leaving for five minutes. A ladies’ compartment, I
presume, and not with Mrs. Bellett?”
They walked slowly along the platform, past the carriage in which
Mrs. Bellett and her sister were arranging their animals and parcels
with much shrill hilarity.
Miss Gordon was so fortunate as to secure a compartment to
herself—the imbecile chuprassi gibbering and gesticulating, whilst
the sahib handed in her slender stock of belongings. As the train
moved away, she leant out of the window and nodded a smiling
farewell.
How good-looking he was as he stood under the lamp with his hat
off! How nice he had been to her—exactly like a brother! She drew
back with a long breath, that was almost a sigh, as she said to
herself, “Of course I shall never see him again.”
CHAPTER XIII.
TOBY JOY.
As the sun died down, the moon arose above the hills and lighted
the travellers along a path winding by the shores of an irregular
mountain lake, and overhung by a multitude of cherry trees in full
blossom.
“Look!” cried Mrs. Brande, joyfully, “there in front you see the lights
of the Dâk Bungalow at last. You will be glad of your dinner, and I’m
sure I shall.”
Two men, who sat in the verandah of the same rest-house, would
also have been most thankful for theirs. The straggling building
appeared full of soldiers and their wives, and there seemed no
immediate prospect of a meal. The kitchen had been taken
possession of by the majestic cook of a burra mem sahib, who was
shortly expected, and the appetites of a couple of insignificant
strangers must therefore be restrained.
These travellers were, of course, Captain Waring and Mark Jervis,
whom the former invariably alluded to as “his cousin.” It was a
convenient title, and accounted for their close companionship. At first
Mark had been disposed to correct this statement, and murmur, “Not
cousins, but connections,” but had been silenced by Clarence
petulantly exclaiming—
“Cousins and connections are the same thing. Who cares a straw
what we are? And what’s the good of bothering?”
“I’m nearly mad with hunger,” groaned Captain Waring. “I’ve eaten
nothing for ten hours but one hard-boiled egg.”
“Smoke, as the Indians do,” suggested his comrade unfeelingly,
“or draw in your belt a couple of holes. Anyway, a little starvation will
do you no harm—you are getting fat.”
“I wonder, if I went and sat upon the steps with a placard round my
neck, on which was written, ‘I am starving,’ if this good lady would
give us a dinner? Hunger is bad enough, but the exquisite smell of
her roast mutton aggravates my pangs.”
“You have only to show yourself, and she will invite you.”
“How do you know, and why do you cruelly raise my hopes?”
“Because I hear that she is the soul of hospitality, and that she has
the best cook on the hills.”
“May I ask how you discovered this really valuable piece of
information?”
“From the harum-scarum youth who passed this afternoon. He
forgot to mention her name.”
“Here she comes along by the weir,” interrupted Waring. “Mark the
excitement among the servants—her meal will be ready to the
minute. She must be truly a great woman, and has already earned
my respect. If she asks me to dinner, I shall love her. What do you
say, Mark?”
“Oh, I think, since you put it in that way, that I should find it easier
to love the young lady!”
“I thought you fought shy of young ladies; and you must have cat’s
eyes if you can see one at this distance.”
“I have the use of my ears, and I have had nothing to do, but
concentrate my attention on what is evidently to be the only meal of
the evening. I heard the cook telling the khitmatghar to lay a place
for the ‘Miss Sahib.’”
“What a thing it is to be observant!” cried Captain Waring. “And
here they are. By George! she is a heavy weight!” alluding to Mrs.
Brande, who was now let down with a dump, that spoke a whole
volume of relief.
The lady ascended the verandah with slow and solid steps, cast a
swift glance at the famishing pair, and went into her own well-
warmed room, where a table neatly laid, and adorned with cherry-
blossoms, awaited her.
“Lay two more places,” were her first commands to the salaaming
Khitmatghar; then to her niece, “I am going to ask those two men to
dinner.”
“But you don’t know them, Aunt Sara!” she expostulated rather
timidly.
“I know of them, and that is quite enough at a dâk bungalow. We
are not so stiff as you are in England; we are all, as it were, in the
same set out here; and I am sure Captain Waring will be thankful to
join us, unless he happens to be a born idiot. In this bungalow there
is nothing to be had but candles and jam. I know it of old. People
who pass up, are like a swarm of locusts, and leave nothing behind
them, but empty tins and bottles. Now I can give him club mutton
and champagne.”
Having carefully arranged her dress, put on her two best diamond
rings, and a blue cap (N.B.—Blue had always been her colour), Mrs.
Brande sailed out into the verandah, and thus accosted the
strangers—
“I shall be very happy if you two gentlemen will dine with me in my
rooms.”
“You are really too good,” returned Captain Waring, springing to
his feet and making a somewhat exaggerated bow. “We shall be
delighted, for there seems no prospect of our getting anything to eat
before to-morrow.”
“You shall have something to eat in less than five minutes,” was
Mrs. Brande’s reassuring answer, as she led the way to her own
apartment.
“This,” waving her hand towards Honor, “is my niece, Miss
Gordon, just out from England. I am Mrs. Brande—my husband is in
the Council.”
“We have had the pleasure of meeting Miss Gordon before,” said
Captain Waring; “this will not be the first time we have sat at the
same table,” and he glanced at her, with sly significance.
“Yes,” faltered Honor, with a heightened colour, as she bowed and
shook hands with Mark. “This is the gentleman of whom I told you,
Aunt Sara, who rescued me when I was left alone in the train.”
“Ah! indeed,” said Mrs. Brande, sitting down as she spoke, and
deliberately unfolding her serviette, “I’m sure I’m greatly obliged to
him,” but she secretly wished that on that occasion Honor had been
befriended by his rich associate.
“Let me introduce him to you, Mrs. Brande—his name is Jervis,”
said Captain Waring, with his most jovial air. “He is young, idle, and
unmarried. My name is Waring. I was in the Rutlands, but I chucked
the service some time ago.”
“Well, now we know all about each other” (oh, deluded lady!) “let
us begin our dinner,” said Mrs. Brande. “I am sure we are all
starving.”
Dinner proved to be excellent, and included mahseer from the
lake, wild duck from the marshes, and club mutton. No! Mrs.
Brande’s “chef” had not been over praised. At first every one
(especially the hostess and Clarence Waring) was too frankly hungry
to talk, but after a time they began to discuss the weather, the local
insects, and their journey—not in the formal manner common to
Britons on their mournful travels—but in a friendly, homely fashion,
suitable to a whitewashed apartment, with the hostess’s bed in one
corner.
Whilst the two men conversed with her niece, Mrs. Brande
critically surveyed them, “took stock” as she said to herself. Captain
Waring was a man of five or six and thirty, well set up, and soldierly
looking; he had dark cropped hair, bold merry eyes, and was
handsome, though sunburnt to a deep tan, and his face was deeply
lined—those in his forehead looking as if they had been ruled and
cut into the very bone—nevertheless, his habitual expression was as
gay and animated as that of Toby Joy himself. He had an extremely
well-to-do air (undoubtedly had never known a money care in his
life), he wore his clothes with ease, they fitted him admirably, his
watch, studs, and linen were of the finest quality; moreover, he
appreciated a good dinner, seemed to accept the best of everything
as a matter of course, and looked about intelligently for peppers and
sauces, which were fortunately forthcoming.
“The companion,” as Mrs. Brande mentally called him, was a
younger man, in fact a mere youth of about two and twenty, well set
up, squarely built, with good shoulders and a determined mouth and
chin. He wore a suit of flannels, a silver watch, with a leather chain,
and looked exactly what he was—an idle, poor hanger-on!
Mrs. Brande left him to talk to Honor, and indeed entirely
neglected him for his more important kinsman. Her niece was
secretly aware of (and resented) her aunt’s preference, and
redoubled her efforts to entertain her slighted fellow-traveller. She
had a fellow-feeling for him also. Were they not both dependents—
both poor relations?
“Well, Captain Waring, so you are coming up to see Shirani?” said
Mrs. Brande, with her most gracious air.
“Yes, and I rather want to recall old times out here, and have a
nice lazy summer in the hills.”
“Then you have been in India all the winter?” (The inspection of his
kit the crafty lady kept to herself.)
“Yes. We came out in October. Had a bit of a shoot in Travancore,
and had a couple of months in Calcutta.”
“Then perhaps you came across a Miss Paske, there? Though I
don’t suppose she was in the Government House set. Her uncle is a
nobody.”
“To be sure. We know Miss Paske, don’t we, Mark? She was very
much in the Government House set. All the A.D.C’s adored her. A
little bit of a thing, with tow-coloured, fluffy hair, and a nez retroussé.”
“I know nothing about her nose or hair, but she is at Shirani now.”
“You don’t say so! I am delighted to hear it. She is capital fun!”
Mrs. Brande’s face fell. She sat crumbling her bread for some
seconds, and then said absently, “Did you notice those monkeys on
the way up?”
She had a peculiar habit of suddenly jumping from one topic to
another, figuratively, at the opposite pole. She declared that her
ideas travelled at times faster than her speech. Possibly she had her
own consecutive, if rapid, train of thought, and may thus have
connected Miss Paske with apes.
“Yes, swarms of those old grey fellows with black faces. I suppose
they have a fair club at Shirani, and keep up the whist-room? Are
there many men who play?”
“Only too many. I don’t approve of cards—at least gambling. I do
love a game of whist—I play a half-anna stamp on the rubber, just to
give it a little interest.”
“Do they play high at Shirani?” he asked with a touch of
impatience.
“Yes, I believe they do; and that horrid old Colonel Sladen is the
worst of all.”
“What! is he still up here? he used to play a first-class rubber.”
“He will play anything—high or low stakes—at either night or day
—he pays—his wife pays,” concluded Mrs. Brande, looking quite
ferocious.
“Oh, is she out again? Nice little woman.”
“Out again! She has never been home yet,” and she proceeded to
detail that lady’s grievances, whilst her companion’s roving eyes
settled on his cousin and Miss Gordon.
She was a remarkable-looking, even fascinating girl, quite different
to his impression of her at first sight. She had a radiant smile,
wonderfully expressive eyes (those eyes alone made her beautiful,
and lifted her completely out of the commonplace), and a high-bred
air. Strange that she should be related to this vulgar old woman, and
little did the vulgar old woman guess how she had been championed
by her English niece. The moon shining full on the lake tempted the
whole party out of doors. Captain Waring made a basely ungrateful
(but wholly vain) attempt to exchange ladies with his friend. Mrs.
Brande, however, loudly called upon him to attend her, as she paced
slowly down to the road; and as he lit his cigar at his cousin’s, he
muttered angrily under his moustache—
“I call this beastly unfair. I had the old girl all dinner time. You’ve
got six to four the best of it!”
CHAPTER XV.
A PROUD MOMENT.