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Morphological Evolution, Aptations,
Homoplasies, Constraints
and
Evolutionary Trends
Catfishes as a Case S t u d y o n General Phylogeny
and Macroevolution
Morphological Evolution, Aptations,
Homoplasies, Constraints
and
Evolutionary Trends
Catfishes as a Case Study on General Phylogeny
and Macroevolu tion

Rui Diogo
University of Liege
Liege, Belgium

Science Publishers, Inc.

Enfield (NH), USA Plymouth, UK
SCIENCE PUBLISHERS, INC.
Post Office Box 699
Enfield, New Hampshire 03748
United States of America

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L i b r a r y of Congress Cataloging-in-Publication Data

Diogo, Rui.
Morphological evolution, aptations, homoplasies, constraints and
evolutionary trends: catfishes as a case study on general phylogeny
and macroevolution/Rui Diogo.
p. cm.
Includes bibliographical references and index.
ISBN 1-57808-291-9
1. Catfishes--Phylogeny. 2. Macroevolution. I. Title
QL637.9.S5D56 2004
597l.49--dc22

ISBN 1-57808-291-9

O 2005, Copyright Reserved

All rights reserved. No part of this publication may be reproduced, stored in

a retrieval system, or transmitted in any form or by any means, electronic,
mechanical, photocopying or otherwise, without the prior permission.

Published by Science Publishers, Inc., NH, USA

Printed in India.
 Preface

The catfishes, or Siluriformes, are included in the superorder Ostariophysi, a

major group of teleosts including, apart the Siluriformes, the
Gonorynchiformes, the Cypriniformes, the Characiformes and the
Gymnotiformes. With their 34 families, about 437 genera and more than 2700
species, catfishes represent about one third of all freshwater fishes and are
one of the most diverse and economically important groups of fresh and
brackish fishes in the world. They have a particularly wide and complex
geographical distribution, being found in North, Central and South America,
Africa, Eurasia, South-East Asia, Japan and Australia, with fossil catfishes
having even being reported in Antarctica.
The amazing diversity and complexity of order Siluriformes renders very
difficult the study of higher level phylogeny and thus general evolution of
this group as a whole. As highlighted by De Pinna (1998), in such a diverse
and complex group, historical processes such as phylogenetic convergences,
parallel adaptations to similar habitats or evolutionary reversions (e.g. in
subterranean catfishes) may expectedly be quite frequent. Such frequency of
alterations in historical processes raise serious problems in correctly inferring
the relationships among catfish groups from all continents, however, and
consequently from similar habitats within different regions of the globe. More-
over, such broad general studies are further complicated by another serious
drawback, related to the logistics of obtaining and including an appropriate
number of representatives of all major groups of catfishes in the phylogenetic
analysis. And not to be overlooked is the challenging and time-consuming
task of analysing and controlling, for each of these numerous terminal taxa,
the even more numerous characters necessarily obligatory to undertake such
a general phylogenetic analysis in a satisfactory manner.
The logistical difficulties probably explain why in spite of long-standing
scientific interest in the phylogeny of Siluriformes the only cladistic analyses
to date dealing with the interfamilial relationships of the order as a whole are
Mo's published work (1991) and De Pinna's unpublished thesis (1993).
vi Morphological Evolution, Aptations, Homoplasies, Constraints and Evolutionary Trends

Logistical difficulties might well account for neglect of muscular characters in

traditional studies of catfish interrelationships as these are often not readily
detectable nor readily documented and, as just mentioned, analysing and
controlling a large number of such features in each taxon constitutes a rather
difficult task. Moreover, museums are much more reluctant to loan the
specimens necessary for complete muscular dissection than specimens for
osteological observations.
Given all the obstacles mentioned above and the associated fact that the
only general cladistic studies on Sjluriformes are those of Mo (1991) and De
Pinna (1993), it is not surprising that the vast majority of catfish researchers
continue to view siluriform higher level phylogeny as a largely unresolved
issue (see Alves-Gomes, 2001; Chardon et al., 2003; Gayet and Meunier, 2003;
Ng, 2003; Teugels, 2003). One of the major purpose of this book is tc, analyse
the puzzling, intriguing, higher-level interrelationships of catfishes, and to
discuss the general evolution of these fishes. To achieve this purpose meant
devoting a large part of the work to a cladistic analysis of the higher level
phylogeny of the group. Some terminal taxa not included in previous analy-
ses, and particularly a large number of characters traditionally excluded from
those analyses, such as those concerning catfish myology, have been incorpo-
rated in this analysis, which lends particular importance to complex struc-
tures. The cladistic analysis of catfish general phylogeny and the focus on
their complex structures thus pave the way for a discussion on the evolution
of these complex structures within the whole order Siluriformes and on cat-
fish general evolution. Lastly, the focus on catfish higher level phylogeny and
evolution allows a broader, theoretical discussion on general phylogeny and
macroevolution.

Liege, Belgium RUI DIOGO

February 2nd, 2004
 Acknowledgement

This project greatly benefited from the precious help of P. Vandewalle and
M. Chardon, to whom I owe a deep debt of gratitude. Since they accepted me
in the Laboratory of Functional and Evolutionary Morphology, in 1998, they
have introduced me to the anatomy, functional morphology, phylogeny and
systematics of Vertebrates in general and of Teleostei in particular. They have
shown me how interesting are the Siluriformes, not only in what concerns the
study of many different issues concerning the order itself, but also in what
refers to the ample and diverse implications that the analysis of these issues
has for a general discussion on theoretical Biology. They have not only spared
their time continuously to discuss various points concerning the project and
biological sciences in general, but also introduced me to Belgium and common
life in this beautiful country. I also want to thank very, very much my
laboratory's colleague, E. Parmentier. He is really one of the brightest young
scientists I have ever met, and his persistence, the remarkably ability that he
has to solve all the different type of problems, and the courage he has to enter
and to get deep involved in all type of scientific areas were really inspiring
for me.
A very special thanks to the late G. G. Teugels (Muske Royal de 1'Afrique
Centrale), for kindly providing several specimens studied in this work, for
participating in so many projects and for discussing catfish phylogeny and
systematics with me, and for allowing me to undertake bibliographical re-
search on his Museum, which revealed to be fundamental for this work. I am
also particularly thankful to R. Vari, as well as their colleagues S. Weitzman,
J. Williams and S. Jewett (National Museum of Natural History), for accept-
ing me in his amazing Museum during two academic years, for providing a
large part of the specimens analyzed in this work, and for reading and com-
menting several papers included in this work. I am thankful to I. Doadrio
(Museu Nacional de Ciencias Naturales) and F. Poyato-Ariza (Universidade
Autonoma de Madrid) for also receiving me in their laboratories and for
valuable scientific discussions.
viii Morphological Evolution, Aptations, Homoplasies, Constraints and Evolutio?~ay Trends

I acknowledge J. Cambray (Albany Museum of Grahamstown), P. Laleyk

(Universitk Nationale du Bknin), P. Duhamel (Muskum National D'Histoire
Naturelle), W. L. Fink, D. W. Nelson and H. H. Ng (Museum of Zoology,
University of Michigan), and M. Stiassny (American Museum of Natural
History) for providing numerous specimens studied in this work. Thanks to
R. Bills and P. Skelton (South African Institute for Aquatic Biodiversity) for
the kind donation of three austroglanidid specimens to the Laboratory of
Functional and Evolutionary Morphology, as well as to B. Hall, F. Galis,
T. Grande, T. Abreu, C. Ferraris, J. Lundberg, M. Brito, M.M. de Pinna,
P. Skelton, R. Reis, L. Soares-Porto, P. Bockmann, A. Zanata, E. Trajano,
B.G. Kapoor, F. Meunier, C. Oliveira, P. Peng, M. Hardman, S. He, D. Adriaens,
F. Wagemans, H. Gebhardt, M. Ebach, A. Wyss, J. Waters, B. Perez-Moreno,
G. Cuny, A. Choudhury, M. Vences, S.H. Weitzman, L. Cavin, F. Santini,
J.C.Briggs, L.M. Gahagan, M. Philiphe, J.G. Maisey, J. Alves-Gomes, T. Roberts,
M.J.L. Stiassny, R. Winterbottom and M. Gayet, P. Lecointre and L. Taverne
for their helpful criticism, advice and assistance.
I would like to thank very, very much G. Arratia. Being closely in contact
with her work while editing the book "Catfishes", seeing the seriousness and
the remarkable quality of her research, and hearing from her many and many
comments and, sometimes, I must say, hard critiques, has been extremely
important for me and for my personal formation as a scientist. Thank you so
much.
My special thanks to all my friends, particularly to Pedro Brito, Gregory
Piskula, Joao Malcato, Pedro Castro, Henry Evrard, Denoel Mathieu, Neo
Toumbos, Thomas Stokart and particularly Claudia Oliveira. Thank you very
much to you, Alejandrita Pelito Lindo. A very, very, very special thanks to
my parents, Valter and Fatima, to my brothers, Hugo and Luis, and to my
grandfather, Raul.
I really would also like to thank R. Primlani, who kindly invited me to
publish this work as a book of the prestigious company Science Publishers
Co. Thank you very much for the confidence in my work and for the several
projects we have together. I hope J will not disappoint you.
A particularly thorough review of the whole book was done by Margaret
Majithia, from Oxford & IBH Publishing Co., for which I am especially grateful.
Finally, thanks to all those who were involved in administering my PhD
scholarship (PRAXIS XXI/BD/19533/99, FundacZo para a Ciencia e a
Tecnologia, Portuguese Government) and other grants and/or awards received
during the last years, without whom this work would really not had been
possible.

Thanks to ALL of you!!!

 Contents

Preface
Acknowledgement vii
1. Catfishes: Introduction
1.I Phylogenetic Position within Teleostei
1.2 Catfish Families
1.3 Historical Overview of Higher Level Phylogeny of Catfishes
1.4 Catfish, an Exceptional Biological Group
2. Methodology and Material
2.1 Phylogenetic Methodology
2.2 Delimitation of Terminal Taxa
2.3 Material, Techniques and Nomenclature
3. Phylogenetic Analysis
3.1 Character Description and Comparison
3.2 Cladistic Analysis, Diagnosis for Clades, and
Comparison with Previous Hypotheses
3.3 Character State Changes for Individual Genera
3.4 Results of Phylogenetic Analysis: Major Outlines
4. Higher-level Phylogeny and Macroevolution of Catfishes:
A Discussion
4.1 Structures Associated with Movements of the Mandibular
Barbels
4.2 Pectoral Girdle Complex
4.3 Adductor Mandibulae Complex
4.4 Palatine-maxillary System
4.5 Suspensorium and Associated Structures
4.6 Elastic Spring Apparatus
4.7 A Discussion on the Origin and Biogeographic Distribution
of Catfishes
x Morphological Evolutior?, Aptations, Homoplasies, Constraints and Evolutionary Trends

5. Catfishes, Case Study for General Discussions

of Phylogenetic and Macroevolutionary Topics
5.1 Primary Homologies, Secondary Homologies, and a Priori
Versus a Posteriori Explanations in Evolutionary Biology
5.2 Homoplasies, Consistency Index, and Complexity of
Macroevolution
5.3 Functional Uncouplings and Morphological Macroevolution
5.4 Myological Versus Osteological Characters in
Phylogenetic Reconstructions
5.5 Analysis of Distinct Anatomical Regions in Phylogenetic
Reconstructions and the Coding of Multistate Characters
5.6 Aptations, Exaptations and Adaptations in
Macroevolutionary Studies
5.7 Parallelisms, Convergences and Constraints in Macroevolution
5.8 Cordelia's Dilemma, Historical Bias, and General
Evolutionary Trends
5. 9 Punctuated Equilibrium, Speciation, Living Fossils and
Macroevolution
References
List of Abbreviations -
Index
 Catfishes: Introduction

1.1 PHYLOGENETIC POSITION WITHIN TELEOSTEI

The catfishes, or Siluriformes, a distinctive order comprising 34 families, about
437 genera and more than 2700 species (de Pinna, 1996,1998; Ferraris and de
Pinna, 1999; Teugels, 2003), represent about one-third of all freshwater fishes
known worldwide. They are one of the economically important groups of
fresh and brackish fishes in the world (Teugels, 1996), as well underscored by
the 'Asian-American economical catfish war' very much in the business
newspapers these days (see, e.g., Roy, 2003). Catfish are characterised, as
their name indicates, by the presence of barbels surrounding the snout region
(see Fig. 1.1).They have a wide geographic distribution and are found in

Fig. 1.1 Maxillary, mandibular and nasal barbels of a generalised catfish, Chysichthys polli. A)
Lateral view of the body. B) Dorsal view of the head. C) Ventral view of the head
(modified from Risch, 1987).
2 Rui Diogo

North, Central and South America, Africa, Eurasia, South-East Asia, Japan
and Australia, with fossil catfishes having been recorded even in Antarctica
(Grande and Eastman, 1986).
The Siluriformes have long been included in the Ostariophysi. This
superorder contains slightly more than 25% of the teleost species and about
80% of all freshwater fishes, including, apart from the Siluriformes, the
Gonorynchiformes, Cypriniformes, Characiformes and Gymnotiformes (Fig.
1.2) (Nelson, 1994; Teugels, 1996) [Note: as a precautionary measure, the
extraordinary and remarkable tentative new ostariophysan order
Sorbininardiformes of Taverne (1999) has not been included here because
more data and a greater general consensus concerning its status are needed.]
The ostariophysans are recognised, as their name indicates (osteon= bone;

TELEOSTEI
-tsome basal teleostan fossil
-SUPERCOHORT Elopomopha
-ORDER Elopiformes
-ORDER Albuliformes
-ORDER Anguilliformes
-SUPERCOHORT Osteoglossocephala
-COHORT Osteoglossomorpha
-ORDER Osteoglossoidei
-COHORT Clupeocephala
-SUBCOHORT Ostarioclupeomorpha
-SUPERORDER Clupeomorpha
-ORDER tEllimmichthyiformes
-ORDER Clupeiformes
-SUPERORDER Ostariophysi
-SERIES Anatophysi
-ORDER Gonorynchiformes
-SERIES Otophysi
-ORDER Cypriniformes
-CLADE Characiphysi
-ORDER Characiformes
-CLADE Siluriphysi
-ORDER Siluriformes
-ORDER Gymnotiformes
-SUBCOHORT Euteleostei

Fig. 1.2 The phylogenetic position of the Siluriformes within the Teleostei, based on Arratia's
1997 classification of the basal Teleostei.
 Catfishes: Introduction 3

fysis= bladder), by a specialised set of anterior vertebrae associated with the

swim bladder. This specialisation is significantly less developed in members
of the series Anatophysi (Gonorynchiformes),however, than in members of
the other ostariophysan series, the Otophysi (Cypriniformes,Characiformes,
Gymnotiformes and Siluriformes), where a true chain of Weberian ossicles
interconnects the swim bladder and the labyrinth organ (see, Fink and Fink,
1981,1996;Gayet and Chardon, 1987; Poyato-Ariza, 1996; Grande and Poyato-
Ariza, 1999).
The phylogenetic position of the catfishes within the Ostariophysi was the
subject of extended discussion in the past century. For much of the century,
and in a large number of studies concerning the interrelationships of
ostariophysans, the Siluriformes were seen as the most plesiomorphic
otophysans, the Characiformes and the Gymnotiformes often considered sister-
groups (see Regan, 1911a, b; Greenwood et al., 1966; Rosen and Greenwood,
1970; Roberts, 1973). Such a view was sternly called into question in 1981,
however, when Fink and Fink promoted the first explicit cladistic analysis of
ostariophysan phylogeny. According to their study, later reinforced by another
study by these authors (Fink and Fink, 1996), the Siluriformes are, in fact, a
specialised group of Otophysi constituting, together with the Gyrnnotiformes,
the Siluriphysi (Fig. 1.2). The characiforms, often viewed as related to the
gymnotiforms, were placed by Fink and Fink (1981,1996) as the sister-group
of the Siluriphysi. The cypriniforms are placed at the very base of the
otophysans (Fig. 1.2).
Fink and Fink (1981: Fig. 1.1) presented several non-homoplasic
synapomorphies to support the sister-group relationship between catfishes
and gymnotiforms, namely: 1)dorsal portion of mesethmoid compressed and
appears slender in dorsal aspect; 2) absence of intercalar; 3) reduction of eye
size; 4) absence of sclerotic bones; 5) infraorbital series consisting largely or
entirely of canal-bearing portions of bones; 6) supraorbital bone absent;
7) ectopterygoid greatly reduced posteriorly or absent; 8) endochondral portion
of the metapterygoid triangular and appearing to be equivalent to the anterior
half of the metapterygoid in primitive otophysans; 9) Opercle with triangular
shape; 10)primordial ligament attaching on the anterodorsal, and not posterior
tip of anguloarticular; 11) presence of only one pharyngobranchial tooth-
plate; 12) articular process of intercalarium absent; 13) third neural arch with
anteroventral process articulating with or fused to a dorsal prominence on
the second centrum; 14) transverse process of fourth centrum with ovoid
anterolateral face; 15) os suspensorium with elongate anterior horizontal
process; 16)horizontal projection of pleural ribs; 17)well-developed Baudelot's
ligament attaching to skull; 18) posterior fin rays offset posteriorly from
anterior ray; 19) flanges for muscle attaching proximally on ventral ray halves
about equal in size to those on dorsal ray halves; 20) medial radial ossification
absent along entire length of both dorsal anal fin pterygophores; 21) when
caudal fin present, anal fin rays articulate directly with proximal radials, and
4 Rui Dingo

distal radials reduced; 22) presence of electroreception; 23) anterior lateral

line nerve with recurrent branch innervating electroreceptors of trunk (for
more details concerning these characters, as well as their phylogenetic impli-
cations, see Fink and Fink, 1981).
Some of Fink and Fink's (1981) characters have, however, been sharply
criticised, in particular by Mireille Gayet, a palaeontologist who has elabo-
rated several criticisms, some of which very pertinent and incisive (Gayet,
1986a, b; Gayet and Chardon, 1987). Other characters presented by Fink and
Fink (1981) but not mentioned in Gayet's critique, such as those related with
catfish suspensorium, for example, appear to be based on a seemingly erro-
neous interpretation of the skeletal components of this complex structure in
the Siluriformes (see Diogo et al., 2001a; Diogo and Chardon, 2003). It is also
noteworthy that neither of the two major independent studies which
purportedly supported Fink and Fink's work (Dimmick and Larson, 1996;
Arratia, 1992) in actuality has directly supported their phylogenetic hypoth-
esis on ostariophysan interrelationships. The independent analyses performed
in these two studies resulted, as a matter of fact, in a phylogenetic scenario
quite different from that proposed by Fink and Fink (1981).Only a posteriori
recombination of the results of these independent analyses with the latter's
characters allowed a consensus with the latter's scenario. The phylogenetic
analysis based on genetic data performed by Dimmick and Larson (1996)
pointed out a clade formed by the Characiformes and the Gymnotiformes,
with the catfishes a sister-group of this clade and the Cypriniformes the most
basal Otophysi. The phylogenetic analysis based on morphological data per-
formed by Arratia (1992) pointed out a clade formed by the Gymnotiformes
and the Siluriformes, as shown by Fink and Fink, but in Arratia's study the
clade was the sister-group of a clade formed by both the Characiformes and
the Cypriniformes (for more details, see Arratia, 1992; Dimmick and Larson,
1996).
However, all the contra points listed above notwithstanding, it has to be
acknowledged that concerning the sister-group relationship between
siluriforms and gymnotiforms, the numerous synapormorphies presented by
Fink and Fink (1981), the additional characters presented by these authors in
1996, and the lack of studies substantively challenging this sister-group
relationship, renders Fink and Fink's scenario the most acceptable one avail-
able at the present time (see Fig. 1.2).
With respect to the phylogenetic position of the Ostariophysi within the
Teleostei, the case is somewhat similar to that referring to the phylogenetic
position of the catfishes within the otophysans. During much of the last century
the Ostariophysi were seen as somewhat derived teleosts belonging to the
Euteleostei and, hence, not closely related to the Clupeiformes (see, e.g.,
Lecointre, 1995, for an historical overview of this subject). However, in the
last decades this view was severely disputed through independent studies
undertaken by various researchers analysing different types of data. These
 Catfishes: Introduction 5

strongly support a sister-group relationship between ostaryophysans and

Clupeiformes and thus the exclusion of Ostariophysi from Euteleostei (see,
e.g., Lecointre, 1995, and references therefrom; Arratia, 1997; Inoue et al.,
2003; Saitoh et al., 2003).
The phylogenetic position of catfishes within ostariophysans and of the
latter fishes within the teleosts can thus be resumed by the basal teleostean
classification presented by Arratia (1997), and given here in Figure 1.2 as the
hierarchical framework for the present work.

1.2 CATFISH FAMILIES

As mentioned above, 34 catfish families are commonly recognised today, of
which two, tHypsidoridae and tAndinichthyidae, comprise fossil remains
exclusively. A brief presentation of each of these 34 families, based to a great
extent in, but, of course, less detailed than, the excellent up-to-date systematic
overview recently published by Teugels (2003), is given below. The
presentation is complemented by Table 1.1 which overviews the principal
published cladistic studies providing relevant information on the
autapomorphies of each of these families, as well as on the relationships
among their genera (for a more detailed presentation, see Teugels, 2003). It
should be noted that this presentation does not focus on the interrelationships
among the different families since this issue is taken up in the next Section.

Familv Akvsidae
This family includes 4 genera and about 27 species. Its monophyly was
supported by the studies of Mo (1991) and de Pinna (1996) and its phylogeny
studied by de Pinna (1996) (see Table 1.1). Two subfamilies are presently
recognised, the Akysinae and the Parakysinae. With respect to external
morphology, the akysids, small-sized (about 10 cm) stream catfishes known
from southern China and South-East Asia, are recognised by the presence of
four pairs of barbels, strong dorsal and pectoral spines, and a long adipose
fin (absent or represented by a ridge in Parakysis). The head, body and fins
are covered with unculiferous plaques, with some of those situated on the
body greatly enlarged and arranged in longitudinal rows.

Family Amblycipitidae
The monophyly of amblycipitids, which include 3 genera and about 22 species,
is well supported (see Table 1.1). Their interrelationships were studied by
Chen and Lundberg (1994). These torrent catfishes occur in fresh water in
southern Asia, from India to southern Japan, and are recognised externally
by a small-size, robust body, depressed head, four pairs of barbels, short
dorsal and pectoral spines, smooth body covered with thick skin, and an
adipose fin more or less confluent with the caudal fin.
6 Rui Diogo

Table 1.1 List of principal cladistic studies published to date providing relevant
information on the phylogenetic relationships among the genera and/or
on the autapomorphies of the various catfish families. Note: a (p) after
the reference of a certain study indicates that the respective study only
provides information about the relationships among part of the family;
the three subfamilies of the Pimelodidae, i.e., Pimelodinae,
Pseudopimelodinae and Heptapterinae, are separately presented here (for
explanations, see text).

Family Relationships among the dlfierenf Autapomorphies fo support

genera of the family monophyly of fhe family
Akysidae De Pinna, 1996 Mo, 1991; De Pinna, 1996
Amblycipitidae Chen and Lundberg, 1994 Mo, 1991; Chen and Lundberg, 1994;
De Pinna, 1996; Diogo et al., 2003b
Amphiliidae He et al., 1999; Diogo, 2003b Diogo, 2003b
Andinichthyidae Family with only a single genus Gayet, 1988
Ariidae NA* Mo, 1991; Oliveira et al., 2002
(but see comments in text)
Aspredinidae De PiAnna,1998 De Pinna, 1996; Diogo et al., 2001b
Astroblepidae Family with only a single genus Schaefer and Lauder, 1986; Schaefer,
1990; Howes, 1983a; De Pinna, 1998
Auchenipteridae Curran, 1989; De Pinna, 1998; Curran, 1989; De Pinna, 1998; Diogo
Soares-Porto, 1998 (p) et al., in press-a
Austroglanididae Family with only a single genus Mo, 1991
Bagridae Mo, 1991; Maeda et al., 1994 (p); Mo, 1991; Diogo et al., 1999
Ng, 2003
Callichthyidae Reis, 1998a Schaefer, 1990; Reis, 1998a
Cetopsidae NA De Pinna and Vari, 1995
Chacidae Family with only a single genus Brown and Ferraris, 1988
Clariidae NA Diogo and Chardon, in press
Claroteidae Mo, 1991 Mo, 1991
Cranoglanididae Family with only a single genus Diogo et al., 2002a
Diplomystidae Arratia, 1987, 1992 Arratia, 1987, 1992
Doradidae De Pinna, 1998 De Pinna, 1998
Erethistidae De Pinna, 1996 De Pinna, 1996; Diogo et al.,
in press-b
Heptapterinae Ferraris, 1988a (p); Lundberg et al., Lundberg and McDade, 1986;
1991a; Bockmann, 1994 (p) Ferraris, 1988a; Lundberg et al., 1988,
1991a; De Pinna, 1998
Heteropneustidae Family with only a single genus Diogo and Chardon, in press
Hypsidoridae Family with only a single genus Grande, 1987; Arratia, 1992 (but see
Mo, 1991: 195; Grande and De Pinna,
1998: 471)
Ictaluridae Lundberg, 1975a, 1982, 1992 Grande and Lundberg, 1988;
Lundberg, 1992
Loricariidae Howes, 1983a (p); Schaefer, 1987, Howes, 1983a; Schaefer and Lauder,
1991 (p), 1998 (p); Armbruster, 1986, 1996; Schaefer, 1987, 1990; De
1998 (p); Montoya-Burgos et al., Pinna, 1998
1997 (p), 1998
Malapteruridae Family with only a single genus Mowes, 1985a
Mochokidae NA* Mo, 1991
(Contd.)
 Caffshes: Infroducfion 7

(Confd.)
Nematogenyidae Family with only a single genus Arratdia, 1992; De Pinna, 1998
Pangasiidae Pouyaud et al., 2000 (p) Diogo et al., in press-c
Pimelodinae Lundberg et al., 1991b; De Pinna, Lundberg et al., 1988, 199:Lb;
1998 De Pinna, 1998
Plotosidae NA" Oliveira et al., 2001
Pseudopimelodinae NA" Lundberg et al., 1991a;
De Pinna, 1998
Schilbidae NA" NA"
Scoloplacidae Family with only a single genus Schaefer et al., 1989; Schaefer, 1990
Siluridae Bombusch and Lundberg, Bombusch, 1991b; Howes
1989 (p); Bombusch, 1991a (p), and Fumihito, 1991
1995; Howes and Fumihito,
1991 (p)
Sisoridae De Pinna, 1996 (p); He, 1996 (p) De Pinna, 1996; Diogo et al., 2002b
Trichomycteridae De Pinna, 1988 (p), 1989ab (p), De Pinna, 198913, 1992, 1998
1992, 1998; De Pinna and Stames,
1990 (p); Costa, 1994 (p);
Costa and Bockmann, 1994 (p)
"NA: not available.

Family Amphiliidae
The monophyly of this family including 9 genera and about 60 species was
supported (contra He et al., 1999: see below) by a phylogenetic study of Diogo
(2003b) in which three monophyletic subfamilies were recognised:
Amphiliinae, Doumeinae and Leptoglanidinae. The amphiliids, endemic to
tropical African fresh waters and with a maximum total length of 195 mm
but usually much smaller, have three pairs of barbels, the nasal pair being
absent. Dorsal and pectoral fins are absent (except in Leptoglanis, Trachyglanis
and Zaireichthys) but adipose fin present (in Trachyglanis it is preceded by a
spine). Some specialised genera present a series of imbricate bony scutes on
the body.

Familv tAndinichthvidae
Family thdinichthyidae comprises a single genus, tAndinichthys, established
for a single fossil species, tAndinichthys bolivianensis, from the Maastrichtian
of Tiupampa, Bolivia. The most remarkable feature of this species is the
presence of a well-developed, deep 'supratemporal commissure' on the
posterior region of the cranial roof (for more details, see Gayet, 1988; Arratia
and Gayet, 1995; Gayet and Meunier, 2003).

Familv Ariidae
This family includes 21 genera (including the genus Ancharius: see below)
and about 153 species. The monophyly of Ariidae was supported by studies
of Mo (1991) and Oliveira et al. (2002),but its interrelationships remain largely
8 Rui Diogo

unknown (see Table 1.1).Moreover, it should be noted that the characters

listed by these authors for the monophyly of Ariidae refer to all members of
the group except the puzzling genus Ancharius from Madagascar, which is
traditionally considered an ariid but has been placed by some authors in its
own family (Anchariidae)or in the family Mochokidae (see Glaw and Vences,
1994; Ferraris and de Pinna, 1999; Teugels, 2003, for more details on this
subject).Most ariids are marine, but some are confined to fresh waters, as, for
example, precisely the members of genus Ancharius. They are found worldwide
in tropical and subtropical regions and are externally recognised by a robust
body compressed posteriorly, three pairs of barbels (nasal pair missing), dorsal
and pectoral fins with a strong spine and a forked caudal fin.

Family Aspsedinidae
Includes 13 genera and about 35 species. Its monophyly was supported by De
Pinna (1996) and Diogo et al. (200:lb) and a cladogram of its interrelation-
ships (based on an unpublished Ph.D. thesis, by Friel) is presented in De
Pinna (1998: Fig. 1.17). Three subfamilies are recognised: Bunocephalinae,
Aspredininae and Hoplomyzontinae. The aspredinids are distributed through-
out most of South America, with some members (the aspredinins) occurring
in the sea, in brackish water and in estuaries and tidal portions of rivers.
Externally, these 'ugly', peculiar catfishes are recognised by broadening of
the head and anterior part of the body and a slender compressed tail. Their
body usually bears knobs and sometimes a series of small plates is present
along the lateral line and base of the dorsal and anal fins. There are three
pairs of barbels (nasal pair missing). In addition to the three pairs of barbels,
some species have numerous small barbels on the anterior part of the body.
The dorsal fin is small and often spineless, adipose fin absent and the leading
pectoral ray spiny.

Family Astsoblepidae
The monophyly of this family comprising a single genus, Astroblepus, of about
54 species, is well supported (see Table 1.1). Astroblepid catfishes, known
from montane regions in Panama and western South America up to Peru,
have an elongated body presenting a dorsal fin with a strong spine and an
adipose fin that may be present or not. Their mouth is inferior, forming a
sucker disc. They possess two pairs of barbels, the maxillary and nasal ones.

Family Auchenivteridae
This large family includes 21 genera and about 107 species. Members are
small to medium in size and confined to fresh waters in South America and
Panama, although some are tolerant to brackish and salt water. The monophyly
of the family is well supported and its interrelationships likewise relatively
 Catfishes: introduction 9

well studied (see Table 1.1), with two subfamilies being recognised, the
Centromochlinae and the Auchenipterinae. The auchenipterids have an elon-
gated, laterally compressed body. They usually have three pairs of barbels
(nasal pair missing), except in Ageneiosus, Tetranematichthys and one species
of Entomocorus, which have only a maxillary pair (sometimes even rudimen-
tary). The dorsal fin is small, adipose fin may be present (small) or absent,
anal fin may be very long and pectoral fin exhibits a strong spine.

Family Austroglanididae
This small family includes a single genus, Austroglanis, whose three species
are known only from the Orange-Vaal and the Olifants river systems in
southern Africa. Its monophyly was well supported by the study of Mo (1991).
Austroglanidids are small catfishes externally recognised by the presence of
three pairs of barbels (nasal pair missing), strong dorsal and pectoral spines,
a rather small adipose fin positioned posteriorly on the body and some
rheophilic adaptations.

Family Baaridae
The Bagridae is a large family, including 18 genera and about 144 species,
which occurs in fresh waters in Central, Southern and South-East Asia except
for species of genus Bagrus, endemic to Africa. Mo (1991),Maeda et al. (1994)
and Ng (2003) have provided a cladistic account of the interrelationships of
its members, and Mo's 1991 and Diogo et al.'s 1999 studies supported its
monophyly (see Table 1.1). Two subfamilies are recognised, Ritinae and
Bagrinae. With respect to external morphology, bagrids are recognised by a
moderately elongated body, compressed posteriorly and depressed in the
head region. Four pairs of barbels are present (two in Rita), the dorsal and
pectoral fins have spines, and an adipose fin is present.

Family Callichthyidae
The callichthyids are mostly known from forest streams in a large part of
South America and from Panama and Trinidad. The monophyly of this family
including 8 genera and about 172 species, was supported by Schaefer (1990)
and Reis (1998a), and its phylogeny studied in detail by Reis (1998a). Two
subfamilies are recogrused, Callichthyinae and Corydoradinae. Callichthyidae
are characterised by a relatively short body covered with two rows of bony
plates, up to two pairs of maxillary barbels and one pair of mental barbels,
and the eventual presence of fleshy flaps. The snout is blunt, the mouth
inferior, the dorsal and adipose fins present a strong spine, and in some
genera the pectoral fin also has a strong spine. Some callichthyid species can
practice aerial respiration and are able to move on land.
10 Rui Diogo

Family Cetopsidae
The monophyly of this family including 2 subfamilies, Helogenidae and
Cetopsinae, 6 genera and about 22 species, was well supported in the work of
De Pinna and Vari (1995), but the interrelationships of its members are not
known (see Table 1.1). The cetopsids, confined to fresh waters in South
America, are externally recognised by an elongated, naked body, three pairs
of barbels (nasal pair missing), and dorsal and pectoral fins lacking pungent
spines. Anal fin base long and fin with numerous rays; adipose fin minute or
absent in adults.

Familv Chacidae
Chacidae includes the single genus Chaca with its three species. Its monophyly
was supported by Brown and Ferraris (1988). Chacids occur in fresh waters
from the Ganges in India to Borneo in South-East Asia, and are characterised
by a unique, long, broad and flattened head, posteriorly compressed body,
three pairs of barbels (minute nasal barbel may be present), numerous
cutaneous flaps or cirri on head and body, and dorsal and pectoral fins
preceded by a spine.

Family Clariidae
Clariidae includes 15 genera and about 89 species. However, its monophyly
seems only to be corroborated when the genus Heteropneustes of family
Heteropneustidae is also included in it as some of its members are seemingly
more closely related with Heteropneustes than with other clariids (for more
details see Chardon, 1968; De Pinna, 1998; Diogo and Chardon, in press).
Although clariids are the subject of extensive study, no cladograms of their
interrelationships based on explicit cladistic analyses have been published
thus far (see Table 1.I).Clariids are known as air-breathing or walking catfishes
(most, but not all, present a well-developed suprabranchial organ, formed by
extensions of the second and fourth epibranchials).They occur in fresh waters
in Africa, extending to Syria and southern Turkey, the Indian subcontinent
and in South-East Asia, with only Clarias being common to both continents.
Concerning their external morphology, clariids are characterised by an
elongated body with a long, spineless dorsal fin, long anal fin, adipose fin of
moderate to large size in some genera (e.g. Heterobranchus, Dinotopterus),
pectoral fin with a leading spine, and presence of four pairs of barbels. In
some extremely elongated genera (e.g. Channallabes, Gymnallabes) the paired
fins are reduced or absent and the dorsal and anal fins are confluent with the
caudal fin.

Family Claroteidae
Mo (1991) described this family, which includes 13 genera and about 78
species, for part of the genera previously included in Bagridae. In the same
 Catfishes: Introduction 11

work, the author provided support for its monophyly, as well as an account
of its interrelationships, with two subfamilies recognised, Claroteinae and
Auchenoglaninae. The external morphology of claroteids is similar to that in
bagrids, with the body moderately elongated and compressed posteriorly,
and the head depressed and usually presenting four pairs of barbels (three in
Auchenoglanis).Dorsal and pectoral fins with strong spines and an adipose fin
present.

Family Cranoglanididae
Diogo et al. (2002a) listed some autapomorphies to define this poorly studied
small family including the single genus Cranoglanis and three species known
from fresh waters in Yunnan Province in China and in North Vietnam (see
Table 1.1).Externally, cranoglanidids are recognised by the large, inferiorly
placed eyes, four pairs of barbels, strong dorsal and pectoral spines, a small
posteriorly placed adipose fin and a high number of anal fin rays (35-41).

Family Diplomystidae
According to Teugels' 2003 overview the family includes a single genus,
Diplornystes, with about six species endemic to the Austral subregion of South
America and occurring in fresh waters in central and southern Chile, and
from San Juan to Patagonia in Argentina, most of which are highly threat-
ened. The phylogenetic relationships among these species were studied by
Arratia (1987, 1992). Arratia also provided good support for the monophyly
of the family as a whole. Diplomystids are easily recognised by the presence
of more than one row of functional teeth along most of the ventral margin of
the maxilla (see below the different phylogenetic interpretations of this
character by different authors), exclusive presence of maxillary barbels and
whole body covered with large papillae and tubercles. Dorsal and pectoral
fins with strong leading spine and a relatively long adipose fin present.

Family Doradidae
This large family of South American catfishes includes 30 genera and about
71 species. Its monophyly was supported by De Pinna (1998) (based on an
unpublished Ph.D. thesis by Higuchi). A cladogram of its interrelationships
(also based on Higuchi's unpublished thesis) is presented in De Pinna (1998:
Fig. 1.14), with three subfamilies being recognised, Platydoradinae,
Astrodoradinae and Doradinae. Doradids are externally recognised by a thick-
set body, generally covered laterally with a row or series of bony plates
which may bear strong, spiny scutes, three pairs of barbels (nasal pair miss-
ing), dorsal and pectoral fins presenting a strong spine and usually an adi-
pose fin.
12 Rui Diogo

Familv Erethistidae
The monophyly of this small Asian family, which includes 6 genera and
about 13 species that were previously assigned to the Sisoridae, was supported
by De Pinna (1996) and Diogo et al. (in press-b). Its interrelationships were
studied by De Pinna (1996), with two subfamilies being recognised,
Erethistinae and Continae. Externally, erethistids are somewhat similar with
sisorids, presenting four pairs of barbels, a small dorsal fin that sometimes
bears a spine, and an adipose fin.

Family Heteropneustidae
The Heteropneustidae, as presently defined (see commentaries above, in the
presentation of the family Clariidae), includes a single genus, Heteropneustes,
and two species occurring in fresh waters in southern Asia from Pakistan to
Thailand. Some autapomorphic features exclusive of Heteropneustes were
described by Diogo and Chardon (in press). Externally, members of this genus
are characterised by an elongated body, four pairs of barbels, short spineless
dorsal fin, very long anal fin that may be confluent with the caudal fin, and a
pectoral spine connected with a venom gland.

Family tHyvsidoridae
Family tHypsidoridae includes a single genus, tHypsidoris, which includes
two fossil species, tHypsidoris farsonensis from the Eocene Green River
Formation of Wyoming and tHypsidoris oregonensis from the Eocene Clarno
Formation of central Oregon. The most remarkable features of these two
fossil species are the presence of maxillary teeth and the highly developed
coronoid process of the lower jaw (for more details and commentaries
concerning these characters, see Grande and De Pinna, 1998).

Family Ictaluridae
The monophyly of this North American family including 7 genera and about
46 species was supported by studies of Grande and Lundberg (1988) and
Lundberg (1992). Its phylogeny was also the subject of cladistic analyses by
the latter author (Lundberg, 1975a; 1982; 1992).Ictalurids have a moderately
elongated body, four pairs of barbels and an adipose fin. The dorsal (except
in Prietella) and pectoral fins have a strong spine and, in some species of
Noturus, the pectoral spine has a poison gland at its base. Some ictalurid
species are troglobitic and share morphological features that seemingly
correlate with their subterranean habitats, such as a depigmented body,
absence of eyes and reduced lateral line.

Family Loricariidae
This is the largest and one of the most studied catfish families; it includes six
subfamilies-Lithogeninae, Neoplecostominae, Hypoptopomatinae,
 Catfishes: Introduction 13

Loricariinae, Ancistrinae and Hypostominae, about 95 genera and more than

650 species. Its monophyly is well supported and the phylogenetic
relationships among its members have been the subject of numerous cladistic
studies (see Table 1.I). Loricariids are freshwater catfishes from Central and
South America, with some genera reported from brackish water, however.
Externally, loricariid catfishes are recogrused by an elongated body compressed
ventrally and generally covered with bony plates, a ventrally placed mouth
often modified into a sucking disc and a pair of maxillary barbels (sometimes
extremely reduced) connecting the upper and lower lip. Nasal and mandibular
barbels absent; dorsal and pectoral fins have a flexible spine with numerous
odontodes on these spines as well as in the snout region.

Familv Malapteruridae
The monophyly of this African family comprising a single genus, Malapterurus,
with 3 species was supported by Howes (1985a). Morphologically, the
malapterurids or electric catfishes (they possess an electric organ of muscular
origin that produces violent electric discharges up to 450 volts) are recog-
nised by a more or less elongated cylindrical body, three pairs of barbels
(nasal pair missing), absence of dorsal fin and presence of adipose fin, and
spineless pectoral fin.

Family Mochokidae
The monophyly of Mochokidae, a large African family including 10 genera
and about 177 species, was supported by Mo (1991). However, the
interrelationships within the family are not known (see Table 1.1). The
mochokids present a robust body, slightly compressed posteriorly and three
pairs of barbels (nasal pair missing), with mandibulary barbels and sometimes
maxillary ones also, being branched in some genera. Dorsal fin with strong
spine, followed by an adipose fin; caudal fin generally forked and pectoral
fins presenting a well-developed pectoral spine. In Chiloglanis the upper and
lower lip expand and unite to form a sucker.

Family Nematonenyidae
Some unique autapomorphies have been described by Arratia (1992) and De
Pinna (1998)to diagnose the single species included in this family, Nematogenys
inermis from central and southern Chile. The members of this species, highly
endangered, are externally recognised by an elongated body, three pairs of
barbels (only one mandibular pair), dorsal fin situated midbody, pectoral fin
with leading spine, and absence of a dorsal spine and an adipose fin.

Familv Panrrasiidae
Pangasiidae includes 4 genera and about 28 species occurring in southern
and South-East Asia. Some autapomorphies of this family have been described
14 Rui Diogo

by Diogo et al. (in press-c), and a cladistic analysis of part of this family was
published by Pouyaud et al. (2000) (see Table 1.1). The external morphology
of pangasiids consists of a laterally compressed body, short dorsal fin with
one or two spines, small adipose fin, long anal fin with numerous fin rays,
strong pectoral spine and two pairs of barbels (maxillary and mandibular).

Familv Pimelodidae
Pimelodidae, a family of freshwater catfishes from South America, Central
America, southern Mexico and the Caribbean Islands comprising 54 genera
and about 312 species, is one of the largest and most diverse Neotropical
groups. The external morphology of members of this family shows a high
degree of diversity. The body is naked and somewhat elongated; there are
three pairs of barbels (nasal pair absent) and in some genera, maxillary bar-
bels are longer than the body. The dorsal spine is sometimes absent, adipose
fin always present, and pectoral spine may/may not be present. Achially,
nowadays most authors attribute the notable diversity of Pimelodidae to the
fact that the family is a heterogeneous assemblage comprising 'three major
well-defined monophyletic groups, currently ranked as subfamilies, namely,
Pimelodinae, Heptapterinae and Pseudopimelodinaef that do not form a
monophyletic 'Pimelodidae' clade (De Pinna, 1993 : 313). That is why I decided
in a recent detailed overview of the phylogeny and systematics of this clade
to treat these three subfamilies as separate families but retain their subfamilial
names to avoid unnecessary nomenclatural complication (Table 1.I). It can be
seen from this Table that all three pimelodid subfamilies have been the subject
of several studies with reference to their monophyly but only Pimelodinae
and Heptapterinae are relatively well studied with respect to their phylogenetic
intrarelationships.

Family Plotosidae
Plotosidae includes 10 genera and about 32 species occurring in the western
Pacific and the Indian Ocean from the east coast of Africa to Australia. About
half of the species are confined to fresh water and occur in Australia and
New Guinea. Some autapomorphies of this family have been described by
Oliveira et al. (2001).The phylogenetic relationships among the plotosid genera
have not been studied so far (see Table 1.1)The external morphology of
plotosids consists essentially of an elongated body with a pointed tail, four
pairs of barbels, two dorsal fins, and a pectoral fin with a strong spine.

Family Schilbidae
The monophyly of this large and diverse family, containing 15 genera and
about 55 species of pelagic fishes from the fresh waters of Africa and southern
Asia was questioned by Mo (1991). Regan (1911b) recognised three schilbid
subfamilies, namely Schilbinae, Siluranodontinae and Ailiinae. Schilbids are
 Catfishes: Introduction 15

externally recognised by a laterally compressed body, two to four pairs of

barbels, dorsal and adipose fins that may/may not be present, a very long
anal fin with numerous fin rays and a pectoral fin presenting a strong leading
spine.

Family Scoloplacidae
Schaefer et al. (1989) and Schaefer (1990) provided strong evidence to support
the monophyly of this small family including the single genus Scoloplax and 4
species endemic from fresh waters of Brazil, Peru and Bolivia. Scoloplacids
are small catfish with a somewhat compact body, covered with two bilateral
series of odontode-bearing plates. Maxillary barbels are well developed and
mandibular barbels may/may not be present. Dorsal fin, as well as pectoral
one, with spine; adipose fin absent.

Family Siluridae
The phylogeny of this Eurasian family including 10 genera and about 79
species is relatively well studied, and its monophyly well supported (see
Table 1.I). As mentioned by Teugels (2003),the external morphology of silurids
differs somewhat from that found in other catfishes. The head and body are
compressed, nasal barbels absent, one and in some genera two pairs of
mandibular barbels present, dorsal spine flexible, pectoral spines usually weak,
anal fin very long, and adipose fin absent.

Familv Sisoridae
Sisoridae is a large family including 16 genera and about 97 species. Its
members are bottom-dwelling catfishes ranging in size from 20 mm to 2 m
and occur in mountain rapids but also in large rivers in southern and eastern
Asia, with one genus (Glyptothorax) also known from the Tigris-Euphrates
basin in Turkey, Syria, Iraq and Iran and from the Black Sea drainage of
Turkey. Monophyly of the family was supported by De Pinna (1996) and
Diogo et al. (2002b) and its phylogeny was the subject of cladistic analyses by
He (1996) and de Pinna (1996), with the latter author recognising two
subfamilies, Sisorinae and Glyptosterninae. Sisorids present a more or less
thickened leathery skin with unculiferous tubercles or plaques, four pairs of
barbels, small dorsal fin, adipose fin and, in genera inhabiting fast-flowing
mountain streams, a mouth developed into a sucker and a belly with special
adhesive modifications.

Family Trichomycteridae
The last family to be presented, Trichomycteridae, is a very large and diverse
family including 41 genera and about 183 species occurring in Costa Rica,
Panama and South America, including southern Patagonia. Eight subfamilies
16 Rui Diogo

are presently recognised--Copionodontinae, Trichogeninae, Trichomycterinae,

Vandelliinae, Stegophilinae, Tridentinae, Glanapteryginae and Sarcoglanidinae.
Although trichomycterids are usually cited as an example of parasitic catfishes,
only a few are truly parasitic, with the vast majority being free-living predators
(see de Pinna, 1998, for more details on this subject). The phylogeny of the
Trichomycteridae was the subject of some cladistic analyses and the
monophyly of the family is well supported (see Table 1.1). With respect to
their external aspect, trichomycterids usually have an elongated naked body,
two pairs of maxillary barbels and a pair of nasal barbels (mandibular barbels
may/may not be present), a spineless dorsal fin, and usually also spineless
pectoral fins.

1.3 HISTORICAL OVERVIEW OF HIGHER LEVEL PHYLOGENY OF

CATFISHES
Several studies on the relationships between the various siluriform families
have been undertaken since the catfishes were designated a formal group per
se for the first time by Rafinesque in 1815. Important classic studies are, for
example, Cuvier (1817)' Bleeker (1852, 1862), Giinther (1864), Gill (1872),
Eigenmann and Eigenmann (1890)' Bridge and Haddon (1893)' Boulenger
(1904), Goodrich (1909), Regan (1911b) and Chardon (1968).
However, the number of studies devoted to this subject has considerably
increased in the last three decades due to the renewed impetus provided by
the advent of cladistics in the second half of the last century (De Pinna, 1998;
Diogo, 2003a). This Section provides an overview of those cladistic studies
published in these last decades dealing with the higher level phylogeny of
the Siluriformes, based mainly on an extensive Chapter published by the
author (Diogo, 2003a) in the two-volume book "Catfishes". As mentioned in
that Chapter, when the author refers to phylogenetic studies dealing with the
relationships between various catfish families, he refers only to published
cladistic studies. Among the several reasons for following this procedure,
detailed in that chapter, one of the most important is to minimise the confusion
associated with such a puzzling issue as the phylogeny of Siluriformes. In
fact, despite their evident importance, many of the 'precladistic' (see De Pinna,
1998) studies dealing with catfish relationships are highly confusing, group-
ing certain taxa due to the presence of both plesiomorphic and highly
homoplasic characters, or simply (several times) without giving a clear expla-
nation for doing so. Therefore, to compare clearly, objectively and coherently
the conclusions of all these complex, highly confusing studies using such
'precladistic' methodologies with the phylogenetic results of those studies
following a cladistic methodology is extremely demanding. Furthermore, an
excellent and detailed extensive historical overview of the most important
'precladistic' works on the taxonomy and classification of siluriforms (see
above) was just recently published by De Pinna (1998).
Of the explicit cladistic works published to date on catfish phylogeny, the
vast majority concerned the intrarelationships within a particular catfish
 Ca+shes: Introduction 17

family (see Table 1.I). The only published cladistic studies presenting original,
explicit cladograms on the interfamilial relationships of either a part or the
whole of the order Siluriformes are those of Howes (1983a), Grande (1987),
Schaefer (1990), Mo (1991), Arratia (1992), De Pinna (1992, 1996, 1998),
Lundberg (1993) and He et al. (1999).A pr6cis of each is given below:

Howes, 1983a (Fig. 1.3)

In his Figure 1.22, Howes presents a hypothesis on loricarioid relationships
(shown in Fig. 1.3 here), based on both his own observations and an
unpublished thesis by Baskin, 1972 (Howes, 1983a: 341-342). According to
this hypothesis, Astroblepidae and Loricariidae form a clade that is the sister-
group of Scoloplacidae, with these three families constituting in turn the
sister-group of Callichthyidae. Also, according to this hypothesis, the clade
formed by these four families is the sister-group of Trichomycteridae, with
Nematogenyidae the next sister-group and hence, the most basal of the six
loricarioid families. In order to support the phylogenetic hypothesis illustrated
in his Figure 1.1, Howes (1983a: 342) advanced several derived morphologi-
cal characters (involving mainly the osteology, myology and external mor-
phology of the cephalic region, the Weberian apparatus and the caudal fin) to
support its different nodes. The clade including all six loricarioid families is
diagnosed as 'swim bladder encapsulated, divided into separate vesicles;
some part of the cranium contributing to encapsulation'. The clade including
all loricaroid families excluding the Nematogenyidae is defined by 'claustrum
and intercalarium lacking from Weberian vertebral series'. The characters
uniting Callichthyidae, Scoloplacidae, Loricariidae and Astroblepidae are:
'posttemporal contributing to distal portion of swim-bladder capsule; de-
rived hypural fusion pattern; low number of principal caudal fin rays'. Fi-
nally, the Scoloplacidae, Loricariidae and Astroblepidae are united due to the
presence of a 'connecting bone between the 1st rib and the 2nd pterygophore'.
The Loricariidae and Astroblepidae are grouped into a monophyletic clade
by the presence of a retractor premaxillae muscle, the medial division of the
protractor hyoideus, as well as the 'reverted lip', 'lateropterygium' and the '6
fused anterior centra'.

Fig. 1.3 Hypothetical relationships among the loricarioid families (the family Diplomystidae is
used as the outgroup) according to Howes' 1983a paper.
18 Rui Diogo

However, Howes called attention to some derived characters that conflict

with his phylogenetic hypothesis. The single derived character defining the
clade including all non-nematogenyid loricaroids (see above), for example,
conflicts with the 'lateral insertion of the dilatator operculi muscle', a derived
character shared by both Nematogenyidae and Trichomycteridae (Howes,
1983a: 341-342).

Grande, 1987 (Fig. 1.4)

In 1987, Grande, based on his own osteological observations of a fossil catfish
from the Eocene Green River Formation of Wyoming originally described by
Lundberg and Case (1970), tHypsidorisfarsonensis, and comparisons with other
catfishes, proposed an original hypothesis on the higher level phylogeny of
Siluriformes (Fig. 1.4). According to this hypothesis the fossil catfish family
tHypsidoridae is the sister-group of all other non-diplomystid catfish families
(= Siluroidea), with the clade formed by the latter plus tHypsidoridae
(= Siluroidei)being the sister-group of Diplomystidae (Fig. 1.4).Grande (1987:
48) listed three characters to diagnose the suborder Siluroidei, namely: 1) '17
or fewer principal caudal rays'; 2) 'an extension of lamellar bone over the
ventral surface of the fifth centrum'; 3) 'fifth centrum joined closely to the
complex centrum'. He (1987: 48) listed five characters to diagnose the
superfamily Siluroidea: 1) loss 'of maxillary teeth'; 2) loss or reduction of
'distal expansion of the maxilla'; 3) loss of 'elongate mesial process of the
maxillaf; 4) reduction of 'palatine either to an extremely small bone, or to a
rod-shaped bone'; 5) 'long, interdigitating sutural contacts between the
ceratohyal and epihyal'. Grande's 1987 hypothesis contradicts the phylogenetic
hypothesis formulated in the original description of tHypsidorisfarsonensis, in
which this fossil species was described as a member of the family Ictaluridae
(Lundberg and Case, 1970: 451-456).

. . (
m -
1
Wyps#lfonbae {Siluroidsi,HypsWmii)
All other f a m i t i (Situ~,SUWOidsa)
Fig. 1.4 Hypothetical relationships among the Diplomystidae, tHypsidoridae and the other
catfish families according to Grande's 1987 paper.

Schaefer, 1990 (Fig. 1.5)

Schaefer (1990) undertook a phylogenetic analysis to infer the relationships
among loricarioid families (Fig. 1.5) as well as between the scoloplacid species.
This study was based on osteological, myological and arthrological structures
of the cephalic region and osteological structures of the axial and caudal fin.
The numerical analysis (using PAUP) of a data matrix of 72 characters
x 9 terminal taxa, which did not include autapomorphic characters, resulted
in a single, most parsimonious cladogram with 77 steps and a Consistency
Index (CI) = 0.842. Concerning the interrelationships between Loricariidae,
 Catfshes: introduction 19

Fig. 1.5 Hypothetical relationships among the loricarioid families according to Schaefer's 1990
paper.

Astroblepidae, Scoloplacidae and Callichthyidae, this cladogram (Fig. 1.5) is

similar to that of Howes (1983a). The synapomorphies listed by Schaefer
(1990: 204) to diagnose the clade including these four families were: 1) 'loss
of the mesethmoid lateral cornua'; 2) 'fusion of pterotic and supracleithrum';
3) 'loss of canal in the lachrimal-antorbital'; 4) 'loss of tight attachment of
premaxillae with the neurocranium'; 5) 'four or fewer branchiostegal rays'; 6)
'dorsal hypurals fused with the compound caudal centrum'; 7) 'presence of
mesethmoid-maxillary ligament'; 8) 'presence of mesethmoid-premaxillary
ligament'; 9) 'presence of retractor tentaculi muscle'. Synapomorphies listed
by Schaefer (1990: 204) to define the clade formed by astroblepids, loricariids
and scoloplacids included: 1)'loss of open cranial fontanels'; 2) 'loss of cranial
aperture which receives the cleithral dorsal process'; 3) 'presence of bifid jaw
teeth'; 4) 'loss of interoperculum'; 5) 'ventrolateral shift in articulation of rib
on sixth centrum'; 6) 'presence of lateral bone'; 7) 'loss of pterygoethmoid
ligament'; 8) 'loss of cranial attachment'; 9) 'loss of interoperculomandibular
ligament'; 10) 'shift in origin of retractor tentaculi muscle'; 11)'bifurcation of
hyohyoideus muscles'. Finally, the synapomorphies listed by Schaefer (1990:
204) to support the sister-group relationship between loricariids and
astroblepids included: 1) 'loss of contact of mesethmoid posterior process
with the frontals'; 2) 'presence of a hyomandibula-metapterygoid suture';
3) 'ventromedial rotation of the mandibles'; 4) 'ankylosis or suture between
sixth centrum and Weberian complex central; 5) 'loss of vertebral
parapophyses'; 6) 'presence of expanded transverse shelf on first anal fin';
7) 'geniohyoideus bilaterally subdivided'; 8) 'presence of an expanded oral
disk'; 9) 'right and left sides of lower jaw not tightly associated at midline';
10) 'presence of intermandibular cartilage'; 11)'presence of juxtaposed nostrils'.
The most significant difference between Schaefer's (1990) and Howes'
(1983a) studies is that Schaefer (1990) did not place the Trichomycteridae as
the sister-group of the clade formed by these four families, but instead in an
unresolved trichotomy including that clade, the Trichomycteridae and the
Nematogenyidae (Fig. 1.5).However, it should be noted that, as stressed by
Schaefer (1990: 174)' such an unresolved trichotomy was not the consequence
of his own phylogenetic results, but of considering the relationships between
the nematogenyids, trichomycterids and the remaining loricarioids as
'unresolved a priori'.
20 Rui Diogo

Mo, 1991 (Fig. 1.6)

One year after ihe publication of Schaefer's 1990 paper, Mo (1991) published
a study dealing mainly with the phylogenetic relationships of Bagridae,
including a generic-level revision and phylogeny of the family. In addition,
Mo (1991) included a somewhat brief (46 pages in a total of 216 pages plus 63
unnumbered figures) analysis of the higher level phylogeny of siluriforms.
This analysis, based on osteological characters of the cephalic region, the
Weberian apparatus, pectoral girdle and the various fins, but also on a few
soft and/or myological characters, resulted in two considerably different
cladograms. One (Fig. 1.6A), based on an 'unweighted' numerical analysis

- Mabptentridae
- Iaaluridae

A B
Fig. 1.6 Hypothetical relationships among the major groups of the Siluriformes according to
Mo's 1991 paper. A) Cladogram produced from the numerical analysis of 126
unweighted characters. B) Cladogram produced from the numerical analysis of 126
characters with a weighting (4) on one of them, namely the "number of vertebrae
united to the complex vertebra" (Mo, 1991: 193).
 Catfishes: Introduction 21

(using Hennig 1986) of a data matrix of 126 characters x 40 terminal taxa

(which did not include autapomorphic characters), had 602 steps, CI = 0.31,
RI (Retention Index) = 0.64 and was only poorly resolved. The other (Fig.
1.6B) was based on a 'weighted' numerical analysis (also using Hennig 1986)
of the same data matrix in which to one of the 126 characters ('number of
vertebrae united to complex vertebra') was given a phylogenetic weight 4
times that of the other characters. It had 549 steps, CI = 0.36, RI = 0.72 and,
apart from two trichotomies, was completely resolved. According to Mo (1991:
193), this 'weighting' was due to the "morphological stability and consistent
distribution of this character in comparisons with those conflicting features".
One of the main conclusions of Mo's (1991) work was the separation of
'Bagridae' into three monophyletic units, namely Bagridae (sensu Mo, 1991),
Claroteidae, and Astroglanididae; according to Mo, these are more closely
related to other catfish families than to each other (Fig. 1.6).
Another important conclusion of this work is the suggestion that Cetopsidae
occupies a markedly basal position in the order Siluriformes (although the
'Helogeneidae', 'Cetopsidae' and 'Hemicetopsis' of Mo were not, as commonly
accepted today, grouped in a single monophyletic taxon, they were placed in
a rather basal position among Siluriformes in both Mo's cladograms). In Mo's
cladogram I (which he clearly seems to prefer), the cetopsids and diplomystids
are distinguished from all other siluriforms by the presence of two characters:
1) 'interdigital union of the two coracoids'; and 2) 'ramus mandibularis nerve
runs inside hyomandibular for a distance' (Mo, 1991: 204). Of these two
characters, only the latter is listed in Mo's cladogram I1 to diagnose the clade
composed by all non-diplomystid, non-cetopsid catfishes.
Another important, but confusing, conclusion drawn by Mo (1991) is the
phylogenetic position of the fossil catfish tHypsidoris. In his cladogram I1
(Fig.16B),tHypsidoris is placed in a far more derived position than in Grande's
1987 cladogram (Fig. 1.4).However, in his cladogram I (Fig. 1.6A),tHypsidoris
is placed in an unresolved polychotomy, with the phylogenetic position of
this genus thus uncertain.
With respect to the other catfish groups, their phylogenetic position is also,
with just a few exceptions, quite uncertain not only as a consequence of the
poor resolution of Mo's cladogram I (Fig. 1.6A), but more importantly the
significant differences between it and his cladogram I1 (Fig. 1.6B).These few
exceptions are discussed below.
One exception concerns the relationships among the loricarioid families,
which are essentially similar to those proposed by Howes (1983a) and Schaefer
(1990); the only difference is that in Mo's cladograms Trichomycteridae and
Nematogenyidae are considered sister-groups (Fig. 1.6). This sister-group
relationship is supported, according to Mo (1991: 204, 208), by the fact that
trichomycterids and nematogenyids, contrary to other loricaroids, have 'nasal
barbels situated at anterior nostrils'.
In both of Mo's cladograms (1991) the loricarioids and amphiliids are
grouped in a clade closely related to the Sisoridae, Akysidae, Amblycipitidae,
22 Rui Diogo

Clariidae, Heteropneustidae, Aspredinidae and Chacidae (Fig. 1.6). In

cladogram I the clade including Amphiliidae and Loricaroidei is diagnosed
by the 'posterior portion of the palatine reduced into a bony lamina or short
spinelike process without distal cartilage' (Mo, 1991: 204), while in cladogram
I1 no character defines this clade. In Mo's cladogram I the clade including the
loricaroids, amphiliids, sisorids, akysids, amblycipitids, clariids,
heteropneustids, aspredinids and chacids is justified by a 'computer generated
node' (Mo, 1991: 204), while in cladogram I1 the clade including all these
groups is diagnosed by the 'absence of extrascapular' (Mo, 1991: 207).
Both Mo's cladograms suggest a monophyletic clade consisting of the
Auchenipteridae, Doradidae, Mochokidae, Ariidae, Hypophthalrnidae and
Pimelodidae (although the 'Auchenipteridae', 'Ageneiosidae' and
'Centromochlidae' of Mo were not, as is now commonly accepted, grouped
in the family Auchenipteridae, all these three groups were placed in this
clade in both of Mo's cladograms). In Mo's cladogram I this clade is defined
by a single character, namely the 'anteriorly thickened and rounded or convex
mesethmoid' (Mo, 1991: 204). In his cladogram I1 this clade is defined not
only by this character, but also by two other features, namely the presence of
'four infraorbitals' and the 'enclosed aortic canal in the complex vertebra'
(Mo, 1991: 208).

De Pinna, 1992 (Fig. 1.7)

One year after the publication of Mo's work, De Pinna (1992) described a
new subfamily of the Neotropical catfish family Trichomycteridae, the
Copionodontinae. In the same work, he provided a phylogenetic analysis of
the interrelationships among trichomycterids, as well as among those catfishes
and other loricarioids. The 27 characters included in that analysis consisted
mainly of osteological characters of the cephalic region, Weberian apparatus,
dorsal fin, pelvic fin and pectoral girdle, but also included a few soft and/or
myological characters. The handmade comparison of these characters resulted
on a fully resolved cladogram with a CI (autapomorphic characters not
included) = 0.78 that, like both cladograms of Mo (1991), suggested a sister-
group relationship between Trichomycteridae and Nematogenyidae.However,
it should be noted that De Pinna's 1992 phylogenetic analysis was completely
independent of Mo's (1991) (when writing his paper, De Pinna was unaware
of Mo's results). In fact, the main reason that led De Pinna to propose the

Remining laricahids
Nematogenyidae

Trichogenes
Remaining trichomycterids
Fig. 1.7 Hypothetical relationships among the trichomycterids, as well as among these fishes
and other loricarioids according to de Pinna ' s 1992 paper.
 Calfishes: lnlroduclion 23

sister-group relationship between Trichomycteridae and Nematogenyidae was

'to a major extent induced by the inclusion of copionodontines and Trichogenes
in the analysis of lower loricarioid relationships' (De Pinna, 1992: 175). Ac-
cording to De Pinna (1992: Fig. 1.23), this inclusion indicated that, of the four
derived characters traditionally used to support a sister-group relationship
between trichomycterids and the other non-nematogenyid loricarioids, only
one ('transformator process of tripus absent') indeed represented the
plesiomorphic situation for trichomycterids. The other three ('intercalarium
absent', 'ductus pneumaticus absent' and 'superficial ossification covering
ventral surface of articulation between complex vertebrae and basioccipital')
represented instead, an apomorphic configuration exclusively present in a
restricted group of derived trichomycterids (Fig. 1.7: 'Remaining
trichomycterids'). Consequently, the grouping of all non-nematogenyid
loricaroid families, that could no longer be supported by more than a single
derived character, was parsimoniously discarded by De Pinna (1992) in
favour of the sister-group relationship between Trichomycteridae and
Nematogenyidae, supported by three derived characters. These three charac-
ters are: 1) 'mesial juncture between scapulo-coracoids without
interdigitations'; 2) 'first dorsal fin pterygophore inserted posterior to neural
spine of ninth free vertebra'; 3) 'absence of dorsal fin spine and locking
mechanism' (De Pinna, 1992: Fig. 1.23).

Arratia. 1992 (Fig. 1.8)

Arratia's 1992 work is a detailed, extensive study dedicated to the develop-
ment, morphological variation and homologies of the suspensorium of cer-
tain siluriform and non-siluriform ostariophysans.It also provides an analysis,

Fig. 1.8 Hypothetical relationships among certain catfish taxa according to Arratia's 1992 paper.
24 Rui Diogo

based on the suspensorial features examined, as well as some other charac-

ters described previously in other studies (Fink and Fink, 1981; Arratia, 1987;
Grande, 1987), of the phylogenetic relationships among the different
ostariophysan orders, and also among certain catfish groups. With respect to
the relationships among certain siluriforms, Arratia's (1992) analysis resulted
in four practically identical cladograms, the only difference concerning the
relationships among diplomystids. As this Section essentially concerns the
interfamilial relationships of the Siluriformes, I shall only refer to the
cladogram illustrated in Arratia's (1992) Figure 46A (for a detailed explanation
of the methodology followed to produce the other three cladograms, as well
as a discussion of the differences between them, see Arratia, 1992: 126-129).
This cladogram (Fig. l.8), based on a numerical analysis (using PAUP) of a
data matrix of 75 characters x 15 terminal taxa, in which 69 characters were
ordered and 6 unordered, corresponds to the consensus of two equally
parsimonious trees (CI = 0.672) with 137 steps. It supports Grande's (1987)
hypothesis, according to which the fossil catfish tHypsidoris occupies a rather
basal position within the Siluriformes. Arratia (1992: Fig. 46A) listed eight
uniquely derived characters and two homoplasic ones to diagnose the clade
constituted by all non-diplomystid, non-hypsidorid catfishes examined in her
study. These characters are: 1)maxilla without long anterior process; 2) maxilla
rudimentary; 3) articulation between autopalatine and maxilla double,
lateroventrally oriented; 4) absence of autopalatine extension dorsal to dermal
entopterygoid; 5) absence of dermal ectopterygoid; 6) absence of dermal
entopterygoid; 7) presence of link between 'entopterygoid' and prevomer
(homoplasic); 8) loss of notch separating processus basalis and posterodorsal
part of metapterygoid (homoplasic); 9) presence of three or four pairs of
barbels; 10) absence of a supraneural bone above the Weberian apparatus in
adults.
Arratia's cladogram (Fig. 1.8)also supports Mo's (1991) phylogenetic results,
according to which the ariids are somewhat closely related to the pimelodids.
The six characters uniting Parapimelodus and the two ariid genera, Bagre and
Galeichthys, in Arratia's cladogram (Fig. 1.8) are: 1) presence of a sesamoid
ectopterygoid joining the autopalatine and 'entopterygoid'; 2) presence of
ectopterygoid process of metapterygoid (homoplasic); 3) blood vessels run-
ning in tubelike lamellar formation ventral to the Weberian apparatus
(homoplasic); 5) fusion of hypurals 1 and 2 (homoplasic); 6) branched sen-
sory canals (homoplasic).
However, Arratia's cladogram (Fig. 1.8) attributes a rather basal position
to Nematogenyidae and the Trichomycteridae, two families that occupy a
rather derived position in Mo's (1991) cladograms (Fig. 1.6).Indeed, Arratia
(1992: Fig. 46A) listed nine derived characters to separate the diplomystids,
thypsidorids, nematogenyids aitd trichomycterids from all the other catfishes
represented in her cladogram. These are: 1)presence of a rod like autopalatine;
2) no articulation between autopalatine and prevomer (homoplasic);
3) presence of a ligament and/or connective tissue between 'entopterygoid'
 Caffishes: Infroducfion 25

and lateral ethmoid (homoplasic); 4) presence of a metapterygoid-

'entopterygoid' ligament (homoplasic); 5) hyomandibula articulating with
autosphenotic; 6) absence of prootic in the hyomandibular fossa; 7) presence
of bony extension over the ventral surface of the fifth centrum (homoplasic);
8) presence of suture between pterosphenoid and parasphenoid (homoplasic);
9) blood vessels in a groove partially surrounded by lamellar walls in the
ventral part of the Weberian apparatus (homoplasic).
Another significant aspect of Arratia's cladogram is the fact that the
heptapterin genus Rhamdia appears as more closely related to the clade formed
by Parapimelodus (Pimelodinae),Bagre (Ariidae) and Galeichthys (Ariidae) than
to the heptapterin genus Heptapterus. The features listed by Arratia (1992: Fig.
46) to unite the genera Galeichthys, Bagre, Parapimelodus and Rhamdia, and
thus to separate these genera from Hepapterus are: 1) fusion of abdominal
centra 2-6 or more; 2) presence of a small, elongate pharyngobranchial attached
to the epibranchial and medial aspect of the hyomandibula (homoplasic).

Lundberg, 1993 (Fig. 1.9)

In an overview of certain clades formed by African and South American
freshwater fishes and their respective implications for the continental drift
theory, Lundberg (1993) provided a phylogenetic hypothesis (Fig. 1.9)
concerning the relationships among some catfish taxa. This hypothesis was
based on a handmade analysis of 12 osteological and myological characters
of the cephalic region, dorsal fin and Weberian apparatus previously described
by other authors and/or personally observed by Lundberg (Lundberg, 1993:
180).Lundberg's hypothesis (Fig. 1.9) is practically identical to Mo's (Fig. 1.6)'
with the addition of the Eocene fossil catfish "tTitanogIanis" as the sister-
group of the clade constituted by Mocholudae, Auchenipteridae and Doradidae
(the 'Auchenipteridae' and the 'Ageneiosidae' of Lundberg correspond to the
Auchenipteridae of this work). It should be noted, however, that Lundberg
(1993) was seemingly unaware of Mo's study since he makes no mention of it.
The two characters listed by Lundberg (1993: 180) to support the sister-group
relationship between "tTitanoglanis" and the clade including Mochokidae,
Auchenipteridae and Doradidae were: 1) 'posterior edge of supraoccipital
truncated, not draw out to form a process'; 2) 'middle nuchal plate with
anterolateral processes contacting posttemporal-epioccipitalregion of skull'.

"Titanoglanis"

rDoradiiae

Fig. 1.9 Hypothetical relationships among certain catfish taxa according to Lundberg's 1993
paper.
26 Rui Diogo

De Pinna, 1996 (Fig. 1.10)

Based on a phylogenetic comparison of a considerable number of characters
of the cephalic region, Weberian apparatus, pectoral fins and girdle, vertebrae,
dorsal fin, pelvic fins and girdle and caudal fin, De Pinna (1996) developed a
hypothesis on the relationships among the Asiatic Amblycipitidae, Akysidae,
Sisoridae and Erethistidae and the South American Aspredinidae. He also
examined the relationships among some genera of these families. His numerical
analysis (using Hennig 1986) of a data matrix that had 112 characters x 21
terminal taxa resulted in a single, completely resolved, most parsimonious
cladogram with 167 steps, CI = 0.70 (autapomorphic characters included) and
RI = 0.79 (Fig. 1.10).

Amblyceps (Arnbtyapitrdae)
Liobagrus (AmbtycipitWae)

Breitensteinia (Akysibae)
Acmdwrdonichthys (Akyhiidae)
Glyptothorax (Sisoridae)
Pgauctscheneis (Sitidas)
g l y p t ~ ~ o i (Siscrridae)
ds

.
I

Nangra (Siisoridae)

I Conta (Erethistidae)
laguvia (ErethisMae)
PswdaIlaguvia (Erethistidag)
Erethistukb (ErethisMae)
Ham (Erethistidae)
EMistes (Erethistidae)
Fig. 1.10 Hypothetical relationships among the Sisoroidea according to de Pinna's 1996 paper.

One of the most significant conclusions of De Pinna's work was that the
Sisoridae of previous authors was a paraphyletic assemblage, with a subunit
of it (subsequently named Erethistidae by De Pinna) being more closely re-
lated to the Neotropical Aspredinidae than to the remaining taxa previously
assigned to the Sisoridae (Fig. 1.lo). Five synapomorphies were listed by De
Pinna (1996: 64) to diagnose the clade constituted by Erethistidae and
Aspredinidae, of which only the last is non-homoplasic: 1) 'mandibular
laterosensory canal absent'; 2) 'second hypobranchial unossified'; 3) 'anterior
margin of pectoral spine with serrations'; 4) 'internal support for pectoral fin
rays small in size'; 5) 'anterior portion of lateral line running closely in paral-
lel to lateral margin of Weberian lamina'. In turn, ten synapomorphies were
listed by de Pinna (1996: 61) to diagnose the clade formed by these two
 Catfishes: Introduction 27

families plus the Sisoridae sensu stricto, eight of which are homoplasic:
1) 'posterior portion of supracleithrum ankylosed to margin of Weberian
lamina' (homoplasic); 2) 'parapophysis of fifth vertebra strongly flattened
and expanded' (homoplasic); 3) 'parapophysis of fifth vertebra long, almost
or quite reaching lateral surface of body wall'; 4) 'humeral process or region
around it connected to anterior portion of vertebral column by well-defined
ligament-state 3' (homoplasic); 5) 'posterior part of Weberian lamina exten-
sively contacting parapophysis of fifth vertebra'; 6) '(reversal of) anterior half
of segments of pectoral-fin spine elongate, almost parallel to axis of spine'
(homoplasic); 7) 'coracoid with ventral anterior process' (homoplasic); 8) '(re-
versal of) second dorsal fin spine with medial ridge along its anterior surface,
forming bilateral longitudinal pouches' (homoplasic); 9) 'ventral arms of first
dorsal-fin spine with posterior subprocesses attached dorsal to their tip'
(homoplasic); 10) 'basipterygium with ventral longitudinal keel, anteriorly
extending alongside internal arm' (homoplasic).
In addition, De Pinna's (1996) work suggested the existence of a
monophyletic clade formed by the Sisoridae, Erethistidae, Aspredinidae and
Akysidae which, in turn, together with family Amblycipitidae formed
superfamily Sisoroidea (Fig. 1.10).Three synapomorphies were listed to define
the clade including Sisoridae, Erethistidae, Aspredinidae and Akysidae,
namely: 1) 'supratemporal fossae present' (homoplasic); 2) 'supracleithrum
strongly attached to skull'; 3) 'posterior nuchal plate with anterior process
forming facet for articulation with anterior nuchal plate' (De Pinna, 1996: 60).
With respect to superfamily Sisoroidea, De Pinna (1996: 59-60) listed seven
synapomorphies, namely: 1) 'posterior center of ossification of palatine
compressed and expanded vertically' (homoplasic); 2) 'articular region of
lateral ethmoid elongated as a process, with articular facet for palatine at tip';
3) 'parapophysis of fifth vertebra strong and attached to ventral side of
centrum, directed directly transversely to centrum'; 4) 'humeral process or
soft tissue around it connected to anterior portion of vertebral column by
well-defined ligament'; 5) 'segments of pectoral fin spine very oblique, almost
parallel to axis of spine, not evident' (homoplasic); 6) 'dorsal spine with
medial ridges along anterior surface, forming bilateral longitudinal pouches'
(homoplasic);7) 'ventral tip of first dorsal fin pterygophore and correspond-
ing neural spines with contacting facets'.

De Pinna, 1998 (Fig. 1.11)

Two years after the publication of his 1996 work, De Pinna published an
overview of the phylogenetic relationships of the Neotropical Siluriformes,
which included a not completely resolved cladogram expressing the
relationships among the major groups of the whole order (Fig. 1.11). As
explained by him (1998: 289-290), this cladogram was mainly derived from
his 1993 unpublished thesis, with: 1) 'some resolution added on the sisoroid-
aspredinidid part of the tree based on the results of De Pinna (1996)';
28 Rui Dioao

Fig. 1.11 Hypothetical relationships among the major groups of the Silurifomes according to
de Pinna's 1998 paper.

2) 'position of the Ariidae from Lundberg (1993)'; and 3) 'position of

tHypsidoridae left unresolved'.
One of the most remarkable aspects of de Pinna's 1998 cladogram is the
fact that although mainly derived from his unpublished thesis (de Pinna,
1993)' of which the greater part was realised parallelly and thus independently,
of Mo's (1991)' several points of this cladogram coincide with Mo's
phylogenetic results. In fact, small differences excepted, both De Pinna (1998)
and Mo (1991) agree: 1) in a rather basal position of Cetopsidae within the
siluriforms; 2) in the relationships among the different loricarioid families
 Catfishes: Introduction 29

and a close relationship between these families and Sisoridae (sensu lato),
Akysidae, Aspredinidae and Amblycipitidae; and 3) in the relationships among
Mochokidae, Auchenipteridae, Doradidae and Ariidae (however the position
of Ariidae in De Pinna's 1998 cladogram is based on Lundberg's 1993paper).
But there are also some significant differences between the cladogram of
De Pima (1998) and the phylogenetic results of Mo (1991), of which one of
the most notable is De Pinna's suggestion that both the Bagridae and
Claroteidae sensu Mo (1991) are polyphyletic groups.
Another important aspect of De Pinna's 1998 cladogram (Fig. 1.11) is that
it constitutes the first published cladogram providing an explicit hypothesis
concerning the phylogenetic position of the three Pimelodidae groups, i.e.,
Pseudopimelodinae, Pimelodinae and Heptapterinae. In this cladogram the
pseudopimelodins form, together with the loricarioids and sisoroids, a
monophyletic unit that is the sister-group of a clade with the heptapterins
and some bagrids as its more basal taxa (Fig. 1.11). With respect to the
pimelodins, the cladogram suggests a sister-group relationship between these
catfishes and some bagrids, with the clade formed by these two groups being
included with the claroteins, schilbids, pangasiids, Horabagrus and
austroglanidids in a clade included in a large, unresolved pentatomy (Fig.1.11).
Unfortunately, except for the interrelationships among the loricaroid
families, as well as some other specific cases, De Pima's 1998 paper does not
directly provide the phylogenetic characters that support the interfamilial
relationships illustrated in that cladogram (these characters are given only in
de Pinna's 1993 unpublished thesis). Consequently, neither the characters
concerning the polyphyly of the Bagridae and Claroteidae sensu Mo (1991)
nor the characters concerning the phylogenetic position of Pimelodinae,
Pseudopimelodinae and Hepapterinae within the Siluriformes are included
in De Pima's 1998 paper.

He et al., 1999 (Fig. 1.12)

The He et al. (1999) study is dedicated mainly to the phylogeny of the African
family Amphiliidae but also includes an analysis of the relationships between
this family and some other taxa, namely Diplomystidae, Amblycipitidae,
tHypsidoridae, Bagridae, Sisoridae and Lqfoglanis (the phylogenetic position
of this genus, transferred from the Bagridae to Amphiliidae in Bailey and
Stewart's 1984 paper, was considered uncertain a primi by He et al.). The
characters used in this study were osteological features of the cephalic region,
Weberian apparatus, vertebrae, pectoral girdle, dorsal fin, caudal skeleton
and the pelvic girdle. The phylogenetic study of He et al. (1999), based on a
numerical analysis (using PAUP) of a data matrix with 73 characters including
x 14 terminal taxa, resulted in a singlemost parsimonious cladogram with 190
steps and CI = 0.616 (0.603 excluding autapomorphic characters).
According with this cladogram (Fig. 1.12), neither the Amphiliidae or the
Sisoridae are monophyletic groups. The doumein amphiliids are more closely
30 Rui Diogo

Amphilius (Amphiliida~,Amphifiinae)
Pararnpttilius (Arnphitiidae,Amphiliinae)
Euchitogfanis(Sisoridae)
Glyptothorax (Sisoridae)

Andmmia (Amphiliidae, Doumeinae)

Dournea (Amphitiidae, Doumeinae)
ractura (Amphiliidae, Doumeinae)
&twqbnis (Arnphitiidae, Dourneinae)
I- Trachyglanis (Amphiliidae, Doumeinae)
Fig. 1.12 Hypothetical relationships among the amphiliid genera, as well as among these
genera and some non-amphiliid taxa, according to He et al.'s 1999 paper.

related to Leptoglanis and the sisorid Glyptothorax than to either the sisorid
Euchiloglanis or the amphiliin amphiliids (Fig. 1.12).The characters listed by
He et al. (indirectly given in their table 1) to support the clade composed by
the doumein amphiliids, Leptoglanis and the sisorid Glyptothorax were: 1) no
posterior fontanel (homoplasic); 2) posterodorsal process of supraoccipital
short, slightly forked at its posterior end (homoplasic); 3) short maxillary
without enlarged fanlike or forked posterior part; 4) fourth and fifth
parapophyses of Weberian apparatus partly fused, thin and long (homoplasic);
5) proximal 1 and 2 of dorsal fin with independent nuchal plates; 6) all units
of second dorsal spine fused.
Another interesting aspect of He et al. (1999) is placement of the
Amblycipitidae in an unresolved trichotomy leading to this family, the
Diplomystidae, and a clade constituted by the remaining catfishes examined,
including the fossil catfish family tHypsidoridae (Fig. 1.12).

1.4 CATFISH, AN EXCEPTIONAL BIOLOGICAL GROUP

As can be seen and as emphasised in the above Sections, catfishes are a
remarkably diverse and widely distributed group, being 'the most diverse
and widely distributed of ostariophysan groups' (De Pinna, 1998: 280). But
catfishes are clearly not only remarkable in what concerns their amazing
taxonomic diversity and biogeographic distribution. The diversity of these
fishes is, in fact, remarkable in several other aspects, such as their anatomy,
ethology, ecology or functional morphology.
For example, catfishes can attain lengths of up to 5 metres and weights of
330 kg, as does the European Wells Silurus glanis (see Fig. 1.13), rivalling
even the famous, non-catfish osteoglossid Arapaima gigas in dimensions
 Catfishes: Introduction 31

.. .,- I

Fig. 1.13 An example of giant catfish, the European Well Silurus glanis (for more similar
photos on giant, as well as miniature, catfish, see the excellent survey given by
Burgess, 1989).

(Burgess, 1989). The giant South American catfish Paulicea lutkeni has been
said to approach and even surpass these sizes (Burgess, 1989). In contrast,
there are some fully mature catfishes at lengths of no more than 35 mm, such
as Coydoras pygmaeus (Burgess, 1989).
Several siluriforms present no structures related to sound production, while
several others, e.g. the 'croaking' catfishes of family Doradidae, are famous
for their sound production (for more details on this subject, see the up-to-
date overview on catfish sound production given by Fine and Ladich, 2003).
Also, although most catfishes are not poisonous, some, such as the plotosids,
are particularly famous for the strong and painful poison released from the
sharpened extremity of their pectoral spines (for more details on poisonous
catfishes, see the recent overview on this subject by PerriPre and PerriPre,
2003). Some siluriforms are also renowned for their trophic performance.
This is the case of species of the trichomycterid subfamily Vandeliinae which,
by being exclusively hematophagous suckers of the blood from the gills of
other fishes, are the only exclusive hematophagous jawed vertebrates other
than some bats (De Pinna, 1998). Some of the vandellin catfishes are even
popularly known in the Brazilian Amazon for their 'accidental' penetration
of the urethras of humans and other mammals (see Burgess, 1989, for more
details and some photos of these remarkable catfishes). Within the Siluriformes,
some other groups are also famous for their 'aberrant' ecological preferences,
e.g. members of the loricariid genus Cochlidon renowned for eating wood (see
32 Rui Diogo

Baras and Laleye, 2003, for a recent overview on the ecology and behaviour
of catfishes). Some catfishes are also ecologically famous for their subterra-
nean habits, as well documented by Trajano (2003).
Biogeographically, catfishes are also an amazingly interesting, and highly
puzzling, biological group. As already mentioned, they are primarily
freshwater fishes, but families Plotosidae and Ariidae have considerable
representation in marine environments. Catfishes are found in all continents,
with catfish fossil remains having even been reported in Antarctica (see Grande
and Eastman, 1986). One of the most striking and exemplary cases of puz-
zling biogeographic issues within the Siluriformes is the seemingly
phylogenetic position of the South American aspredinids in the very middle
of the Asian sisoroids hypothesised by De Pinna (1996, 1998) (see above).
This, as stated by De Pinna himself, clearly would constitute an 'intriguing
biogeographical phenomenon' (De Pinna, 1996: 77). Some palaeontological
discoveries also provide such intriguing issues, as the finding of a seemingly
ictalurid fossil form in Mongolia (see Stucky, 1982), when actually extant
ictalurid catfishes are exclusively restricted to North American fresh waters.
Or the finding of seemingly clariid and bagrid fossils forms in Europe when
these two latter groups are actually exclusively present in Africa and Asia
(for more details, and other similar, puzzling palaeontological discoveries,
see the detailed up-to-date overview on catfish palaeontology and
palaeobiogeography of Gayet and Meunier, 2003).
Representative examples of the high diversity and complexity of the
Siluriformes could fill a whole book. In fact, the examples given in this chapter
are only a very brief attestation of the indisputably amazing variation and
complexity of this remarkable group of fishes, either from a taxonomic,
anatomical, morphofunctional, ethological, biogeographic, or ecological point
of view. That is why the Siluriformes constitute a very appropriate case study
for a broader discussion on general macroevolution and phylogeny.
 Methodology and Material

2.1 PHYLOGENETIC METHODOLOGY

As already mentioned, much of this book deals with an analysis of the higher
level phylogeny of catfishes. The systematic procedure employed in this analy-
sis is cladistic methodology, as extensively presented and discussed in the
excellent book of Kitching et al. (1998). Parsimony was employed to find the
hypothesis best supported by the analysed data, using both the Hennig86
(Farris, 1988) and the Nona Winclada (Nixon, 2002) computer programs.
The Implicit Enumeration algorithm (ie*) of Hennig86 was employed in the
search for the most parsimonious cladograms, with Nona Winclada used to
check the most parsimonious results found with this algorithm. Hennig86
was chosen mainly because of its efficiency in calculation time (more specific
Information on the options and capacities of the program are provided in
Fitzhugh, 1989). Tree manipulations and diagnostics were done using the
ingenious and updated program Nona Winclada. Autapomorphies for the
different terminal taxa examined were actively searched for and included in
the analysis.
Multistate characters were ordered whenever possible, following the basic
principles presented in Kitching et al. (1998). The same phylogenetic weight
and hence the same phylogenetic importance was given to all characters used
in the analysis, following Wiens (2001). In fact, as pointed out by him, most
phylogenetic studies neglected the fact that by using ordered and unordered
characters without corrective weighting, a different weight and thus different
phylogenetic importance is, in reality, given to the different characters. For
example, if species A and B share state 2 of a certain ordered character X and
if species A and C share state 2 of a certain unordered character Y, this will
favour a grouping of A and B, since they share two evolutionary steps on the
cladogram (from state 0 to state 1 of character X, and then from state 1 to
state 2 of this character, since this is an ordered character), while A and C
share only one evolutionary step on the cladogram (directly from state 0 to
34 Rui Diogo

state 2 of character Y, since this is an unordered character). Thus, a different

weight and hence a different phylogenetic importance is, in reality, attributed
to the characters X and Y since, after all, A and C share, like A and B, state 2
of a phylogenetic character. Therefore, in this simple example, in order to
render the phylogenetic importance of the character Y equal to that of the
character X, the phylogenetic weight of the character X should be multiplied
by x, that is, by 1/(n-1), where n is the total number of states of the character.
In this way, the sharing of state 2 of the unordered character Y by A and C is
ranked as equally important as the sharing of state 2 of the ordered character
X: both characters have a total weight of 1. Consequently, in order to attribute
the same phylogenetic importance to all characters used in the present analysis,
i.e., in order to give an equal total weight of 1 to all these characters, the
following procedure was undertaken: a) the weight of unordered characters,
as well as of ordered characters with two states, was not changed, i.e., it
remained 1; b) the weight of ordered characters with three, four and five
states (there are no ordered characters with more than five states in the
analysis) was multiplied, as described above, by a corrective factor of
respectively, /;, and pi. However, it is important to note that the
phylogenetic outcomes resulting from the two other alternative methodologies
concerning the ordering/unordering of characters, i.e., the exclusive use of
unordered characters and the use of ordered characters without normalisation
of their total weights, will be carefully examined and discussed in Chapter 5.
Special attention was also given to the process of determining the polarity
of the different character states used in the phylogenetic analysis. In order to
facilitate and optimise this process, it was decided to use family Diplomystidae
as the outgroup of all other siluriforms. In fact, the sister-group relationship
between Diplomystidae and all the other siluriforms has long been recog-
nised (see Eigenmann and Eigenmann, 1890; Bridge and Haddon, 1893; Regan,
1911b; Alexander, 1965; Chardon, 1968; Gosline, 1975) and subsequently fully
supported by all cladistic analyses on catfish higher level phylogeny (see
Ch.1, Sec. 1.3). Therefore, defining all non-diplomystids as the ingroup and
the Diplomystidae as the basal outgroup clearly seems to constitute the wis-
est, appropriate, unambiguous and efficient option in the present work. The
configuration found in the non-siluriform ostariophysans was, of course, also
seriously taken into account for decisions on the polarity of the characters. In
fact, as an important framework for decisions on character polarity, a detailed
and extensive analysis on the homologies, anatomical variation and
plesiomorphic configuration of the different morphological features exam-
ined, taking into account the situation found in both the Diplomystidae and
the other ostariophysans, as well as data from different fields such as func-
tional morphology, ontogeny and palaeontology, was carefully undertaken.
Much of this analysis was presented in a series of papers recently published
by the author and colleagues, such as: Diogo and Chardon (2000a) (adductor
mandibulae complex), Diogo and Chardon (200Clb) (structures associated with
the mandibular barbels), Diogo and Chardon (2000~)(structures associated
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 So they fared sumptuously, and both fell fast asleep in big arm
chairs near the stove in the salon afterwards, and when Ray yawned
and woke it was close on three o’clock, and the sun had won and
the mountains all round were shining white against the clear deep
blue.
There was no one else in the salon. There seemed, in fact, no
one else in the hotel except a few officers who kept to the smoking-
room. So he kissed Lois awake, and in five minutes they were
footing it gaily up the Furka road, with the Bernese giants towering
in front and dwarfing all the lesser wonders closer at hand.
“That must be Finsteraarhorn,” said Ray, pointing to the highest
and sharpest peak. “And that one further on is probably Jungfrau,
but I know her better from the other side.”
Then they passed the fortifications and turned a corner, and the
great Rhone glacier lay below them, dappled here and there, where
the sun got into the hollows, with the most wonderful flecks of fairy
colour—tenderly vivid and lucently diaphanous blues and greens so
magically blended that Lois caught her breath at the sight.
“How beautiful! How beautiful!” she murmured. “It is a dream-
colour, but I never dreamed anything half so lovely.”
He could hardly get her along. She wanted to stop at every
second step to gloat on some fresh wonder. But they came at last,
by slow degrees, to the point, just below the Belvédère, where
sturdy pedestrians can drop from the main road into the valley and
so avoid the tedious winding-ways.
“We’ll get down here, if you think you can manage it,” said Ray.
“Then we can get right up to the glacier-foot where the Rhone
comes out. It’s worth seeing, but it’s a bit of a scramble down unless
they’ve improved the path.”
“I’ll manage it all right if you’ll go first and show me the way.”
So they started on that somewhat precarious descent, and had
gone but a little way when Ray began to be sorry he had not stuck
to the solider footing of the road.
For the apology of a path had in places disappeared entirely
under the attrition of the wet season and many heavy boots. Whole
lumps of it had slipped away and left gaps and slides down which a
rough-clad Switzer might flounder with possible impunity, but which
suggested serious possibilities to the ordinary traveller.
He had gone on hoping it would improve, but it did not. Instead
it grew worse. But if falling down such awkward slides was no easy
matter, re-climbing them to gain the high road was next to
impossible.
They bumped and slipped and floundered downwards as best
they could.
“I’m truly sorry,” he said, as he helped her down one specially
awkward place. “It was nothing like this last time I came.”
“It’s all right,” she laughed. “It’s fun—all in the day’s work. Don’t
tumble right out of sight if you can help it.”
And then he did. A lump of rock to which he had trusted his foot
came squawking out of the wet bank, and he and it went down
together a good half-dozen yards.
He brought up with his rucksac over his head and turned at once
to see to her safety.
“All right,” he shouted. “No bones broken. But I don’t advise you
to try it. Strike to the right and try and find a better place. Throw me
down your rucksac and cloak, then you’ll be free-er.”
She dropped them down to him, with a startled look on her face,
and he scrambled round, as well as he could so laden, to meet her
round the corner. But she had to make quite a long détour before
she came at last on another and less precarious path and was at last
able to join him.
“Sure you weren’t hurt?” she asked anxiously.
 “Quite sure. Bit scraped, that’s all. I suppose it’s the rains that
have boggled the path so. Now, if we keep on round here we’ll be
able to get right up to the ice-cave where the stream comes out.
Here’s the rain on again. Better put that cloak on,” and they
scrambled on over the rough detritus from the glacier and the
hillsides till they reached the ice-foot, and stood looking into the
weird blue-green hollow out of which the gray glacier water came
rushing as though in haste to find a more congenial atmosphere.
“It’s the most wonderful colour I’ve ever seen,” she said,
drinking it in with wide appreciative eyes. “It hardly looks real and
earthly. It looks as though a breath would make it vanish. I suppose
if we got inside there it would simply be all white.”
But just then, in sullen warning, a solid lump of overhanging ice
came down with a crash, and a volley of stones came shooting at
them mixed with its splinters, and they turned and went on their
way down the stony valley.
The rain ceased again just as they arrived at the big hotel, and
as Ray swung off his cloak and shook it, Lois laughed and said,
“When we get to Oberwald you must hand me over your
trousers and I’ll stitch them up.”
“Why?—what?—” and he clapped his hands to his hips to feel
the damage, while Lois still stood laughing at the rents and tears
which his cloak had so far hidden.
“I should keep my cloak on if I were you,” she suggested, and
then asked quickly, “Why—Ray? What is it? Are you more hurt than
you thought?”—for the look on his face was one of concern if not of
actual consternation.
“I am,” he jerked, with a pinch on his face, and then he felt
hastily in his other pockets and the tension slackened somewhat.
“But it’s not in my person,—only in my pocket. Would you mind
kicking me, dear? Here,—we’ll go round the corner,” and he stepped
back the way they had come. “And—would you also mind telling me
what money you have in your pocket or your rucksac.”
 “Not very much, I’m afraid. Two or three pounds, I think. Why?”
“Because,” he said, displaying the catastrophe. “That stupid slip
of mine has busted my hip-pocket and all our money’s gone. All
except the change out of this morning’s five-pounder. With that and
yours we can get to Montreux all right, and I can wire from there to
Uncle Tony, but it’s confoundedly stupid,——”
“Couldn’t we find it if we went back?”
“I’m going to try, but you’ll stop here and have some tea to pass
the time.”
“Oh no, I won’t. It’s share and share alike. Aren’t we almost man
and wife? Come along! We’ll have a hunt for our money anyway,”
and she led the way back towards the glacier.
They searched for an hour, but looking for a flat leather purse in
that stony land was like searching for the proverbial needle in the
haystack. They found the exact spot where Ray took his sudden
slide, but search below it discovered nothing. They followed step by
step the way he had taken till he met Lois and then, as well as they
could, the path they had taken to the ice-foot. But there was no sign
of the purse and he came to the conclusion that his pocket was
probably torn by the slide and the purse fell out of it later on,—
anywhere down the two-mile stretch of stony valley between them
and the hotel.
They paced it with meticulous care, searching cautiously, but
found nothing, and at last gave it up and went on,—soberly as
regards Ray, amusedly as regards Lois, who persisted in looking only
at the humorous side of the matter.
“We’ll walk all the way,” she laughed, “and pick out the
cheapest-looking hotels, and you’ll have to haggle like a German
about terms.”
“I’m awfully sick of myself for being such an ass,” he said
gloomily. “It’s hateful to be short of cash in a strange land. I often
used to run it pretty close. I remember once reaching home from
this very place with only a halfpenny in my pocket. I remember I
wanted a cup of tea on the train, more than I’d ever wanted one
before, and I had to go without.”
“Had you lost your purse then also?” asked Lois mischievously.
“No,—just stopped longer than I’d planned and ran it a bit too
fine.”
They plodded into Oberwald just before dark, and stumped
heavily up the steep wooden steps that led from the stony road to
the door of the little Furka Hotel, fairly tired out with the day’s walk,
which their diversion in search of Ray’s purse had extended, he
reckoned, to close on five-and-twenty miles, and he proceeded to
haggle with the depressed-looking landlady like any German of them
all.
She was glad enough to have them, however, even on their own
terms, and gave them a quite sufficient supper, in which three
different kinds of sausage, and veal in several guises, figured
principally; and her bed-rooms, if somewhat meagrely furnished,
were at all events clean. And they went up early to bed, tired with
their long tramp and still tireder,—as Ray expressed it, concerning
himself—of playing the fool with his money and throwing it about for
some wiser man to pick up.
The landlady knew nothing about the war, except that the
diligences had stopped running because the horses were wanted,
and most of the men had gone—to Thun, or Berne, she was not
quite sure where, but it was all because of the talk of war, and she
did not hold with any of it,—stopping business and upsetting
everybody and everything.
Oberwald, they decided, could not at the best of times be a very
inspiring place. Under the shadow of the war-cloud it was dismal.
They had early breakfast on the wooden platform outside the front
door, while the deserted village below and about them roused itself,
lazily and obviously against the grain, to its day’s work.
 But Ray was obviously not up to his usual standard, even though
Lois had borrowed needle and thread from the landlady and had
patched up his rents with deft fingers and visible enjoyment at being
of service to him.
“You’re not letting that old purse worry you, are you?” she had
asked, as they sat over their coffee and cheese and honey on the
wooden platform.
“Not at the loss of it, though the stupidity of losing anything
always annoys me. It’s the possible consequences I’m thinking of. It
came on me all in a heap in the night that it’s just possible we may
have difficulty in communicating with them at home if things are
really bad. I wish to goodness we could get some definite news. I
wanted very much to take you up the Eggishorn—it’s just close here,
and it seems a shame to pass right under it without going up. You
don’t really know what a glacier’s like till you’ve seen the Aletsch.
But....”
“I think we’d better go right on. We can come back some other
time and see all these things. Suppose they shouldn’t have got your
telegram from Leipsic! They’ll be getting frightfully anxious about us.
Let us get on as quickly as possible.”
“I’m afraid there’s nothing else for it,” he said regretfully. “Let’s
see now—it would take us at least four days to walk down the valley
to Montreux.... How much money did you say you have with you?”
“I’ve got three pounds, five shillings. I’ll get it for you.”
“No. Better keep it safe. I might lose it, you know. Well, four
days’ tramping at the lowest possible rate means at least forty
francs. It will pay us to take the train from Brigue. There’s a quick
train about mid-day, I remember ... that is, if it’s still running. They
may have taken the trains off also. It comes from Milan, you see,
through the Simplon.”
“Third class?”
 “Rather. I’ve come home by it more than once, and it’s generally
packed with Italians, who are not the pleasantest of travelling
companions. But needs must when you’re such a fool as to lose your
purse,—and they’re probably all being kept at home just now
anyway. We had a tough day yesterday, so to-day we’ll just jog
along to Fiesch. That’s another place I wanted you to stop at. Most
fascinating country, all the hillsides covered with little irrigation
channels about a foot wide, and the natives spend most of their time
turning them on and off. That’s where you strike up for the
Eggishorn ... and the Märjelen See ... and then there’s Binn.... It’s a
mighty pity to pass them all ...” and he rattled the few coins in his
pocket thoughtfully.
But—“Needs must!” said Lois firmly, anxious to get into touch
with the outer world again and especially with the folks at home.
“Wait a bit!” said Ray thoughtfully, and got down the map from
its peg in the hall, and began figuring with his pencil on the back of
the bill the landlady had just brought him, which came to 9.50 francs
for the two of them. “Just ... you ... wait ... a bit ... my child!” and
he measured and figured away with immense energy and growing
enjoyment.
“We can do it all right,” he burst out at last. “See here!—We’ve
got 160 francs left after settling up here. We’ll get Madame here to
put us up the usual trampers’ lunch,—that’s one franc each. We’ll
walk on to Fiesch and then up to the little Firnegarten Inn—small but
clean—on the Fiescher Alp, and stop the night there. That’ll be, say,
10 francs. It would cost us more down below. To-morrow we’ll make
an early start and climb up to the Märjelen See and the Eggishorn,
taking our lunch with us again. Then we’ll come down by the big
hotel,—we can only afford to look at the outside of it this time,—and
walk along the ridge to Rieder Alp. It’s wonderful,—worth coming all
the way from England for,—that and the Aletsch. Stop the night at
Rieder Alp. That will be say 12 francs, if I haggle well. And next day
we’ll walk down to Brigue and Oberried and Bitsch and the Massa,
and get the mid-day train there for Montreux,——”
 “If it’s running.”
“If not we’ll just toddle on.”
“But can we afford it?”
“Including fares and all it will come to just about as much as
four days’ tramping along the road. And two days up aloft here are
worth forty days on that road. The road’s fine but it’s not to be
compared with the bridle path along Rieder-Alp.”
He was so obviously set on it that, in spite of her anxiety to get
on, she had not the heart to raise any objection, and five minutes
later they were on the road, with the dew-drenched green slopes
above and below them shimmering like diamond-dust in the early
sunshine, and Ray’s spirits at their highest again at this getting the
better of the misfortune that would have done them out of the best
bit of the journey.
As to the fact that they would arrive in Montreux with only 120
francs between them, that did not trouble him in the slightest now
that they were going up aloft.
“I’ll wire Uncle Tony the very first thing when we get there. It’ll
be quite all right, you’ll see, my child. ‘The year’s at the Spring——’”
“Ninth of August!”
“That’s nothing. It’s our year I’m talking of, and it’s only a week
or so after New Year’s Day.... ‘The day’s at the morn. Morning’s at
seven;’——”
“Nearer eight,”—with a glance at her wrist-watch.
“‘The hillside’s dew-pearled,’——”
“Undoubtedly,”—with a comprehensive wave of the hand uphill
and down.
“‘The lark’s on the wing;’——”
“Maybe—somewhere.”
 “‘The snail’s on the thorn; God’s in His Heaven; All’s right with
the world!’”
“With your and my little world. But, oh, I wonder what’s going
on outside there, Ray! It’s terrible to think of war at any time, even
though we none of us really know what it means. But for all the
Great Powers to be flying at one another’s throats,—and England
too! I can’t realise it.”
“Don’t try, child. Rhenius may have caught some flying
nightmare by the tail. I haven’t much faith in Italian newspapers.
Anyway we’ll make the most of these few days of grace and be
thankful for them.... You see, if things really are as bad as he said,
we may be stuck for some time in Switzerland, and an extra day up
here in heaven will make no difference in the end and is all to the
good now. Learn to gather your roses while you may, my child,” and
his determined enjoyment carried the day.
They made Fiesch about noon, and Ray marched her right
through the little town to the house he had stopped at more than
once—the cosy-looking little Hotel des Alpes, near where the rushing
Fieschbach flung its gray waters into those of the Rhone.
They knew him there and were much hurt that he had not come
to stop with them again, and were greatly interested in Lois. He had
to explain matters very fully before they were pacified sufficiently to
permit him to have a bottle of Asti, with a small table and two chairs
outside in the sunshine, and the mistress and the two comely maids
hung about them all the time they ate their Oberwald lunch of bread
and sausage and cheese and biscuits, and insisted on supplementing
it with apples and pears and grapes, grumbling good-humouredly at
him and chattering and giving such news as they had.
“You’d do much better to stop with us. Firnegarten cannot keep
very much of a table up there, you know. Most people go right on to
the Jungfrau Hotel for the night——”
“I know. But we’re pauper-tramps, you see, till we get to
Montreux, and we have to look twice at every sou. You see, I was
fool enough to lose my purse up at Gletsch there——”
“Ach! To lose your purse! That was foolishness. But if you had
come to us we would have helped you.”
“It’s awfully good of you, and we’re going to come back here as
soon as ever we can. There’s heaps of things I want to show
mademoiselle,—Binn, and the Fiescher Glacier, and Ernen—oh,
heaps. But now we’ve got to get on. We’re going to get married as
soon as we reach Montreux, but I couldn’t bear to stump along the
road down here when Aletsch and the Rieder-Alp called me.
Mademoiselle is not at all sure we’re doing the right thing in not
going straight on.”
“You will never regret it, mademoiselle,” they assured her.
“Though, of course, when one is hurrying along to get married,
—” interjected one of the girls thoughtfully.
“The Great Aletsch is a thing to see before one dies,—”
continued Madame.
“Or even before one gets married, when you have to pass right
under it,” said Ray. “And the Märjelen——”
“Ach—the poor Märjelen! It is gone. It got a hole in it
somewhere and all the water has run out, and so now there is
nothing to see.”
“So! But the Aletsch is still there?”
“Och, yes! The Aletsch can never run away through a hole.
There it is and there it will remain till the world comes to an end.”
“And the war? What news have you?”
“They are fighting terribly over there, it seems,—at some place
called Liége. But we do not hear very much since the diligence
stopped. And all our visitors went away at once. We were quite full
and not one has come since. War is bad for everybody. For me, I
cannot understand what people want to fight for. It will not come
into Switzerland, do you think, monsieur?”
 “I shouldn’t think so, but when war once starts you never know
where it will stop. And I’ve no doubt Germany would be only too
glad to get hold of Switzerland if she got half a chance.”
“Ach—those Germans! No, I do not like them. Whenever I see
one come in here I say to myself, ‘Another trouble-maker!’ They are
never satisfied, and they want everything—except to pay proper
prices. No, I do not like them. If they all get killed in the fighting I
shall not care one bit.”
Their leave-taking could hardly have been warmer if Madame
had been jingling in her hand a whole month’s pension fees instead
of the price of a modest bottle of Asti, and presently they were
slowly and steadily climbing the steep and stony path to Firnegarten.
The maid in charge there was sister to one of those down below,
and she also remembered Ray. She was much astonished at their
intention of stopping the night there, and laughed merrily when Ray
proceeded to hammer her price down to his level and then explained
why he was, for once, acting like a German.
She made them very comfortable, however, in a simple way, and
obviously enjoyed their company. They went early to bed, and were
well on their way up the Fiescher Alp soon after seven next morning.
It was close on noon before they struggled up the tumbled
débris of the top, and sank down on a flat rock, with that great glory
of the Aletsch glacier sweeping down in front of them, from the
great snow-basins of Jungfrau and Finsteraarhorn, till it curled out of
sight behind the green ridges of Rieder-Alp away down below them
on the left.
“The Chariots of the Lord!” came involuntarily to Lois’s lips as
she sat gazing on it, and her eyes followed the strange dark parallel
lines which ran throughout its length and looked exactly like gigantic
wheel-tracks. “What makes them?”
“The continuous slow downward movement of the ice, I believe.
It picks off earth and stones from the sidewalls and gradually throws
them into exact lines like that. Curious, isn’t it? I remember it struck
me in just the same way the first time I saw it.”
It was long before she could be got to look at anything else.
“I can’t help expecting it all the time to do something,” she
explained.
“I know. But it never does. See!—that’s Jungfrau over there, and
that one is Finsteraarhorn. And round this other side you can see the
Matterhorn and Mont Blanc. Those big white lumps are the
Mischabels.”
In time he got her to start on her lunch, though she asserted
that it felt like eating in church,—desecration.
“I’m glad you insisted on coming,” she said softly. “It is a sight
one could never forget,” and he was radiant.
“And to think,” she said again, presently, “that over yonder the
guns are booming and men are doing everything they know to kill
one another! Isn’t it dreadful to think of—in face of this great silent
wonder which takes one’s thoughts right up to God?”
“It’s simply brutal.... I just hope whoever’s to blame for bringing
it about will get whipped out of existence.”
He could hardly drag her away. She vowed she could never
weary of that most wonderful sight, and was certain it would begin
to move if they only waited long enough. And so it was a very tired
but very well-satisfied pair that dropped into the first chairs they
came to in the homely little Riederalp Hotel, with barely enough
energy left to arrange terms on the German plan.
Next morning they came down the steep wooded ways by
Oberried and Bitsch and the Massa gorge, and reached Brigue
exactly fifteen minutes before a train started for Montreux.
The run down the Rhone Valley and up to Montreux was full of
enjoyment, tempered only by their doubts as to being able to get
any further than that.
 Ray pointed out to her all the things he knew,—the new
Lötschberg line away up on the opposite mountain-side,—the
openings of Nicolai Thal, leading to Zermatt and Saas Fée,—the Val
d’Anniviers leading to Zinal, and the Val d’Herens to Arolla, and
promised to take her to them all when the times got re-jointed. Then
they were at Martigny, and presently the flat delta and the upper
end of the lake came into sight, and Chillon, and they were at
Montreux.
Ray enquired at once from the station-master as to trains for
Paris.
“Paris, mon Dieu?” jerked that much harassed official. “Ask again
in a fortnight’s time, monsieur, and perhaps we shall know
something then!” and Ray made at once for the Post Office and
wrote out a telegram to Uncle Tony,—“Just arrived here. Both well.
Lost purse. Send cash Poste Restante.”
The young man behind the official window looked at the address
and said in excellent English, “We can send it from here, but we
cannot make sure it will ever get there. You see it must go through
France or Germany, and they are fighting and everything is
disarranged.... It is very awkward,” as they looked at one another in
dismay.
“Very awkward!” said Ray. “Please do your best. Are letters
coming through?”
“Not from England for some days. Doubtless in time matters will
arrange themselves.”
In time, doubtless! But the one thing about which there was no
doubt whatever was the fact that they were in a strange land, cut off
from communication with their own, and that the sum total of their
united funds amounted to something under five pounds,—and there
was no saying when they could procure more.
 XIV
A LMA at St Barnabas’s, and Mrs Dare at The Red House each
received a brief note from Con, from Southampton, saying he
was leaving immediately but was not permitted to say more.

He seemed in the best of spirits and said he had plenty to do.

After that the vail of war fell between him and them, and to them
was left the harder task of possessing their souls in hope, with such
patient endurance as they could draw from higher hidden sources.
Both, however,—Alma in her crowded ward, and Mrs Dare in the less
strenuous and so the more meditative sphere of home,—went about
their daily tasks with tranquil faces which permitted no sign to show
of the fears that might be in them. It was their quiet part in the
crisis to give of their best and suffer in silence, as it was the part of
the millions of other women similarly circumstanced.
Mr Dare had perhaps the heaviest burden to bear at this time,
and in spite of his attempts at cheerfulness the weight of it was
apparent in him. His business at a deadlock, valued customers
urgently claiming the fulfilment of contracts, the goods they wanted
hermetically sealed within the flaming borders of Germany and
Austria, accounts for goods sent to those countries falling due, and
no money forthcoming from abroad to meet them. No wonder he
looked harassed and aged, and at times grew somewhat irritable
under the strain.
What his wife was to him in those days none but he knew,—not
even Mrs Dare herself in full. In her own quiet fashion she would at
times draw him gently on to unburden himself to her in a way that
would have been impossible to anyone else, and her great good
sense would seek out the hopeful possibilities and tone down the
asperities of life. And when things were past speaking about she
would show, by her silent sympathy and brave face, that she
understood but still had faith in the future.
But for an unusually alert and active business man to find
himself, without warning, plunged suddenly into a perfect morass of
difficulties, for which no blame attached to anyone save to the blind
precipitancy of untoward circumstance;—to find himself helplessly
idle where his days had always been briskly over-full,—it was
enough to drive any man off his balance, and in some cases it did.
He went down to St Mary Axe each morning and stopped there
all day in gloomy exasperation. He explained his situation to irritated
clamourers for goods till he grew sick of explaining. He was grateful
when release came at night; and in the night he lay awake at times
and hugged to himself the few precious hours which still intervened
before he must shoulder his burden again. Sunday he looked
forward to, all the week long, as a dies non when business matters
ceased perforce from troubling and his weary soul could take its
rest. He longed for weeks of Sundays. At times, in his utter
weariness, the thought of the final unbroken rest made infinite
appeal to him.
The complete lack of any word from Lois and Ray added not a
little to their anxieties. The Colonel, indeed, never would admit any
possibility of mischance in the matter.
“Don’t you worry, Mrs Mother,” he would adjure her. “They’re
having the time of their lives somewhere or other, I’ll wager you a
sovereign.”
“If they’re shut up in Germany it may be a very unpleasant
time,” argued Mrs Dare.
“But they’re not. Ray’s no fool and he got out of that trap
instanter. Of that I’m certain. Where to I can’t, of course, say. Tirol
seems nearest, from the map——”
“That’s Austria,” said Mrs Dare quietly.
 “Well then, Switzerland—Russia—Italy—anywhere,—I don’t
know. But if he’s still in Germany he’s a much bigger fool than I ever
thought him. They’re all right. Don’t you worry!”—which was all most
excellent in intention but did not bring to the anxious mother-heart
the comfort that one word from the missing ones would have done.
But the Colonel was too busy to waste time and energy in
worrying, and, besides, he was not given that way. Immediately on
the declaration of war, he had donned his uniform and gone down to
Whitehall and tendered his services in any capacity whatever. His
bluff, antique enthusiasm overcame even the natural repugnance of
War-Office messengers to further the wishes of any but their own
immediate chiefs, and he succeeded in seeing Lord Kitchener, whom
he had not met since they toiled up Nile together in quest of
Gordon.
The quiet, level-eyed man, who had gone so far and high since
those days, gave him cordial greeting and expressed the hope that
the younger generation would exhibit equal public spirit, in which
case this belated creation of a sufficient fighting force would prosper
to the extent of his wishes, which he acknowledged were great,
though not more than the dire necessities of the case called for. He
tactfully switched the Colonel’s enthusiasm on to the recruiting
branch line, and the fiery little warrior had since then been devoting
himself, heart and soul, to the business of presenting Kitchener’s
Army to the youth of Willstead and neighbourhood as the one and
only legitimate outlet for its duty to its King and Country.
With his V.C. and his Crimean and Mutiny and African medals, he
made a brave show on a platform, and his fervid exhortations
persuaded many from the outer back rows to the plain deal tables
where the recruiting forms awaited them.
He toured the neighbouring villages in a motor car, and until the
muddle-headed mismanagement by the authorities of the earlier
comers cast somewhat of a chill on their waiting fellows, the Colonel
was a great success.
 Noel and Gregor MacLean were still impatiently hanging on for
the War Office to decide whether or not the London Scottish were to
be permitted to form a Second Battalion. And Noel, with the
impetuosity of youth, grew so restive under the strain at times that
he stoutly urged Gregor to enrol with him in one of the regiments of
Kitchener’s army.
“Man!” he would growl, after the usual ineffectual visit to
Headquarters. “We’re going to get left. It’ll all be over and done with
before we get a look in. Let’s join the Hussars!”
“I’m for the London Scottish, my boy, if it’s at all possible. They
say they’ll know in a week or two for certain, and we can wait all
right. I know such a lot of the fellows there and I’d sooner be among
friends. It makes a mighty difference and they’re all good chaps in
the Scottish. Besides I’ve a natural yearning for the kilt. If they shut
down on us, then we’ll sign on wherever you like.”
“Hang it, man! The fun’ll all be over.”
“Don’t you believe it, my son. K of K isn’t raking in all these men
just to amuse himself. He’s the squarest-headed chap we’ve got, and
those eyes of his see a long long way past Tipperary, you bet. We’re
up against a jolly tough job and he knows it.... Anyway we’ll be fitter
than most when they do take us on. I bet you there aren’t many
recruits can down ten out of twelve clays at two hundred yards.”
This was Noel’s top score so far. He was rather proud of it and
judicious reference to it always had a soothing effect on his feelings.
So they strenuously kept up their training, walking all the way in and
back whenever they went up to Buckingham Gate for news, and
spending much time and money at the shooting-grounds.
The girls missed them, of course, but consoled themselves as
best they could with one another. They did a round of the links each
day for health’s sake, but felt the lack of Noel’s outspoken jibes and
Gregor’s curt criticisms and all the subtle excitation and enjoyment
of the former times, and learned that golf for duty and golf for
pleasure are greens of very different qualities.
 Still they would not have had it otherwise. The boys were doing
their duty as it appeared to them, and it was their portion to miss
them and get along as best they could without them. For their sakes
they heartily wished Headquarters would make up its mind what it
was going to do, and get them settled down to actual work and
disciplined courses.
For this waiting on and on, with no definite certainty as to the
outcome, was wearing on Noel’s temper, and bits of it got out on the
loose at home at times and disturbed the atmosphere somewhat.
Like most boys of his age, when things went his way he was as
pleasant as could be. And they so generally had gone his way that
when they did not he resented it and let people know it. Like nine
boys out of every ten, whose chief concern in life had so far been
themselves and their own troubles and enjoyments, there was a
streak of natural selfishness in him, any implication of which he
would have hotly resented. He could be generous enough of his
superfluities, but so far had had to make no call on himself for the
higher virtues of self-denial or self-restraint. In short he was just an
ordinary boy merging into man, very full of himself and his own
concerns and enjoyments, and at times a little careless of others.
This odd new friendliness which had sprung up of late between
himself and Victoria Luard was all very much to the good. It came in
between him and himself and made him feel ready, and even
anxious, to do great things for her, and to consider her feelings even
before his own. But, at the same time, his feeling of personal
discrepancy with regard to her, drove him in the rebound to
occasional little displays of bearishness and boyish arrogance, the
springs of which Victoria understood perfectly and was vastly
amused at.
Gregor MacLean, with the advantages of his extra five years and
much shoulder-rubbing with his fellows, had grown out of these
youthful discordances, and he sometimes took Noel humorously to
task for his little lapses, and Noel would take more from him in that
way than from anyone else.
 Honor of course, in sisterly fashion, saw his faults and did not
pass them over in silence. Still, she also generally did it in humorous
fashion which left no more than a momentary sting even if it did not
produce much result.
Miss Mitten knitted untiringly. Victoria gravely asserted to Mrs
Dare and Honor, when they had dropped in for tea one afternoon,
that, so assured was Auntie Mitt that the outcome of the war
depended entirely on the number of body-belts and mufflers she
could complete in a given time, that she went on knitting all night
long in her sleep. And Auntie Mitt, in no way offended, though
somewhat scandalised at such public mention of her in the privacy of
her bed, only smiled and knitted harder than ever.
“The cold weather will be coming soon,” she said gently, “and it’s
cold work fighting in the trenches.”
“But, my dear Auntie Mitt, they don’t fight in trenches
nowadays,” said Vic.
“No?... They used to. I remember ... I remember hearing much
of the discomforts of the trenches in the Crimean War from those
who had taken part in it.”
“Nowadays they fire shell at you from four or five miles away
and you’re dead before you know what’s hit you,” said Honor. “It’s
low kind of fighting to my mind.”
“Or drop bombs on you from aeroplanes without a chance of
hitting back,” added Vic, “which is lower still.”
“Well ... I don’t myself agree with anything of that kind,” said
Auntie Mitt gently. “It certainly does not seem to me a very manly
way of fighting.”
“It isn’t. But unfortunately it’s the way that’s in fashion,” said Vic.
“It is very horrible,” said Mrs Dare, busy with her knitting also
and thinking of her two, one of whom would probably sooner or
later be exposed to these barbarous novelties of civilised warfare.
“But of course they respect the Red Cross men,”—in which case Con
at all events might possibly return alive.
“Oh, they’ll respect the Red Cross all right, Mrs Mother,” said the
Colonel, catching her last words as he strode in, with an early
evening paper in his hand. “They’re big fighters but they’re civilised
and they’ll fight like Christians.”
“What a horrible expression!” said Mrs Dare. “Fight like
Christians!”
“Yes,—I apologise and withdraw. You are quite right, Mrs
Mother,” with an old-fashioned little bow towards her. “It was not
happily expressed.... And yet Christians have to fight at times, and if
ever fighting was justified it is now—on our side. We’re fighting for
Right and for the rights of everybody outside Germany. Never in the
history of the world was there a more righteous war as far as we are
concerned. And so we are fighting like—or if you prefer it—as
Christians.”
“Yes, I prefer it that way. It is my only consolation when I think
of the boys. They are fighting for the Right.”
“When they get to it,” said Honor. “What’s the latest, Colonel?
Does Liége still stand where it did?”
“It stands marvellously—the forts that is. The Germans seem to
have the town, but the forts are still alive and kicking. It’s simply
marvellous how those Belgians have suddenly transformed
themselves into the pluckiest fighters the world has ever seen.
Marvellous! No one ever believed they could hold Germany’s millions
for a day, and here they’ve kept them at bay for a whole fortnight
and given France time to get herself in order. If the rest of the war
goes the same way there can be no doubt as to how it will end.”
“Doubt?” echoed Vic scornfully. “You don’t mean to dare to say
you’ve ever had any doubts as to how it would end, Uncle Tony?”
“There speaks Young England,—always cocksure of winning and
inclined to despise the enemy. If you had seen as much of war as I
have, my dear, you would be cocksure of nothing, except that you’d
do your duty to the last gasp and would have to leave the rest to
Providence. Germany is a tremendous fighting-machine. We have a
tough job before us, but we’re fighting for the Right and please God
we’ll win. It’s good to see the new spirit the war is evoking
everywhere. Great Britain and Ireland shoulder to shoulder, and
India and all the colonies rushing to help. It’s magnificent,—simply
magnificent.”
“Yes,” said Mrs Dare quietly. “It is doing good in that way, and in
matters at home also,—the matters which come home to the hearts
of us women. We’ve just formed a committee for looking after the
wives and children of the men who have to go to the front, and
every single person I’ve seen about it is keen to help,—people in
some cases who have hitherto shown no inclination for anything
beyond their own concerns.”
“There will be a good deal of distress one way and another, I
fear,” said the Colonel, nodding thoughtfully. “That is if things go on
as they usually do.”
“I’m inclined to hope they’ll go better,” said Mrs Dare. “Our men
at the head of affairs are in closer touch with the needs of the
people than yours ever have been,”—with a pacificatory little nod
towards him. “I know you don’t like Lloyd George, but you must
acknowledge that he has handled the financial situation in a
masterly way.”
“I do acknowledge it. And I’ll even go so far as to say that I
don’t believe our side would have handled the whole matter as well
as it has been done. We might. Men rise to the occasion,—as yours
have done. We might,—but I confess I don’t at the moment see
which of our men could have done what has had to be done as well
as Sir Edward Grey, and Churchill and Lloyd George and Asquith.”
“Hooray!” cried Honor. “You’ll be on the right side yet, Colonel.”
“I’m always on the side of right, anyway. What are you girls
doing to help?”
 “I’m going to knit body-belts and mufflers,” said Honor
lugubriously. “But I’m only a beginner and I’m shy of performing in
public yet.”
“And you, Victoria-who-ought-to-have-been-Balaclava?”
“Our Central Committee in town is considering how we can best
help, and as soon as they decide I’m on to it. In the meantime,
Honor is teaching me to knit body-belts and mufflers,—that is, she’s
passing on to me, the beginnings of her own little knowledge,—
though I don’t quite see the need of them. It’ll all be over in a
month, I expect.”
“If it’s all over in six months I shall be more than glad,” said the
Colonel weightily. “And there’ll be plenty of cold days and nights
before then. However, I’m glad you’re all doing what you can. It’ll do
you all good.”
 XV
“Y US!” SAID Mrs Skirrow, with an emphasis that carried
conviction. “It may seem a vi’lent utt’rance to you, mum,
but, for me, I’m bound to say I’m right down glad o’ this war. It’s tuk
my three off o’ me hands, an’ it’s givin’ me the time o’ me life.”

“Where have they got to?” asked Mrs Dare sympathetically.

“Jim and George, they’re in Kitchener’s lot at Colchester—the
Hoozars, and me old man’s back in th’ Army Transport, an’ if that
don’t mek him move his lazy bones I d’n know annything this side
the other place that will. It tired him so last time he was in it, that
he’s bin resting ever since. But it’s the thing he knows best, and
when the call come he forgot his tiredness an’ up an’ went like a
man. ‘Damn that Keyzer!’ he says,—you’ll pardon me, mum, but
them was his identical words,—‘Damn that Keyzer!’ he says. ‘He is
the limit,—walking over little Belgium with ’is ’obnails like that
without so much as a by-your-leave or beg-pardon. He’s got to be
knocked out, he has, and I’m on to help jab him one in the eye. And
you two boys,’ he says, ‘you’re onto this job too, or I’ll have the skin
off of you both before you know where y’are. Yer King and yer
country needs yer.’ An’ if you’ll b’lieve me, mum, they went like
lambs.”
“And why did they go into the Hussars? Can they ride?”
“Divv—I mean, not a bit of it, mum. But they talked it over
atween themselves, and Jim, he said, if it come to riding or walking,
he’d sooner ride any day, an’ the spurs made a man look a man. So
they went up together and they was took on like a shot. An’ I’m to
get twelve-and-six a week now and mebbe more later on, they do
say. I ain’t got it yet, but it’s a-comin’ all right, an’ then——”
“Well, I hope you’ll save all you can, Mrs Skirrow. You never
know what the future may bring, you know.”
“That’s true, mum. But I’ve worked harder than most for these
three this many a year, and I’m inclined to think I’ll mebbe tek a bit
of a holiday and have a decent rest. How long d’you think it’ll go on,
mum?”
“I’m afraid no one can tell that, Mrs Skirrow. Colonel Luard says
he will be glad if it’s over in six months.”
“Ah,—well,”—with a satisfied look on her face,—“that’s a tidy
spell. For me, if it was a year I d’n know as I’d mind. It’ll keep a lot
o’ men out o’ mischief.”
“And put many out of life altogether, I’m afraid.”
“Ay—well—mebbe! But there’s always the pension to look
forward to, an’ they do say it’s goin’ to be bigger than ever it was
before.”
“Yes, I’m sure everybody feels that everything possible should be
done for the men at the front and those they leave behind them.”
“That’s right, mum. ’Tain’t such a bad old world after all. D’you
hear about the Chilfers down the road, mum?”
“No. What about them?”
“A rare joke. Everybody’s laughing at ’em. When yon first pinch
come and it lukt ’s if we might all be starvin’ inside a week, Mr
Chilfer he went up in his big motor to th’ Stores, and he come back
with it full,—’ams and sides o’ bacon, all nicely done up, an’ flour, an’
cheeses, an’ I d’n know what all. Lukt like a Carter Paterson at
Christmas time, he did. An’ now prices is down again he wants to
get rid o’ the stuff, an’ nob’dy’ll luk at it ’cos it’s all goin’ bad on ’is
’ands. And serve him jolly well right!—that’s what I say.”
 “And I say the same. It was inconsiderate and selfish and
decidedly unpatriotic. If everybody had done like that where would
the rest of us have been?”
“That’s it, mum. But it’s them Chilfers all over. I’m glad to say
they’ve tuk his car f’r the war, and they’ve tuk all the horses they
could lay their hands on. That’s rough on some. There’s Gilling down
our way. He runs a laundry. They stopped him in the street t’other
day an’ tuk his horse and left th’ van and th’ laundry he was
delivering right there. It’ll put a stop on him I’m thinking, and folks’ll
have to go dirty, unless th’ big laundries pick up all the business.”
“There will be discomforts in all directions, I’m afraid, Mrs
Skirrow. But we’re much better off than the poor people in Belgium
who are being turned out in thousands and their homes burnt over
their heads. It’s dreadful work.”
“’Tis that, mum. An’ begging your pardon, I says like my old
man, ‘Damn that Keyzer, and put the stopper on ’im as quick as may
be!’”
“One cannot help hoping he will suffer as he deserves.”
“That’s right, mum! Bet you I’d trounce ’im if I got half a chance.
I’d twist his old neck like that, I would,”—and she wrung her wet
floor-cloth into her pail with a vehemence that imperilled its further
usefulness. “He’s an old divvle, he is, an’ th’ young one’s worse, they
say. All the same, if they c’d do it so’s none of ’em got killed, for me
I wouldn’t mind th’ war going on for quite a goodish bit.”
“And I would be thankful if it all ended to-morrow.”
“Ah! ’twon’t do that, mum,” was Mrs Skirrow’s safe prophecy.
Since Con’s post-card saying they were expecting to leave within
an hour or two, they had had no word from him, nor was any
information as to the movements of the troops permitted in the
papers. The rigid censorship dropped an impenetrable vail between
the anxious hearts at home and the active operations abroad.
 It was a time and an occasion for the exercise of unparalleled
and implicit faith and hope and trust in the powers that held the
ways, and still more in the Highest Power of all. And on all sides was
manifested an extraordinary strengthening and quickening of those
higher and deeper feelings which had become somewhat atrophied
during the long fat years of peace. The nation and the Empire drew
itself together, forgot the little family disputes which had enlivened
its existence for so long, and stood shoulder to shoulder as never
before. The waters were troubled and the sick were healed.
The Colonel, in the pursuit of his duties, was frequently at the
War Office. He heard, there and at his club, many things of which he
never spoke even to Mr and Mrs Dare in their intimate evening
confabulations.
The full bleak blackness of the days of Mons and Maubeuge
were known to him, and the peril of Le Cateau and Landrecies, and
it was as much as he could do to keep the weight of these grave
matters out of his face at times.
He saw the casualty lists as they were compiled at the office,
long before they were issued, and groaned over them in general and
in particular. Killed, wounded, missing,—many whom he had known,
and more whose people he knew, were already gone. Who would be
left when the full tale was told?—he asked himself gloomily,—when
this was barely the beginning.
Then, one day, his anxious old finger, following the list down,
name by name, stopped with a sudden stiffening on the name of
“Dare, Lieut. C., R.A.M.C.” under the head of “Missing,” and he had
to inflate his chest with a very deep breath and hold himself very
tightly, before he could mechanically get through the rest of the list.
“Missing!”—Under all the ordinary circumstances of civilised
warfare that would leave abundant ground for hope. But the
appalling stories he had been hearing of late as to the newest
German methods left only room for fear.
 They were, on the most indisputable evidence, behaving worse
than the worst of savages. Their barbarous cruelties were the result
of a deliberate system of frightfulness and terrorism inspired by
headquarters. They had shocked and wounded his soul till at times it
had felt sick of humanity at large. But they filled him also with a
most righteous anger which helped to brace him up again.
That a hitherto reputedly civilised nation could, of cold
deliberation, do such things!—and exult in them!—Faugh! It was
savages they were,—and worse than any savages he had ever come
across!
And so he feared the worst for Con, and his heart was heavy for
Con’s wife and mother and father.
He went over to his club to think it over, but found too many
friends there for his present humour. So he turned into St James’s
Park, and walked on and on, with his mind full of Con and Alma,
past the Palace and the Duke’s statue, and found himself in Hyde
Park, where the London Scottish were drilling and manœuvering
with a huge crowd looking on.
That made him think of Ray, and he wondered briefly where
those two had got to. If Ray had been at home, as he ought to have
been, he would have been among these stalwart kilties who looked
fine and fit for anything. As soon as he got home he would take his
place of course. And young Noel and Gregor MacLean,—he had
heard that very day that reserve battalions were to be raised pretty
generally. So they would be in it too. And that was all right. Duty
called, and it was the part of the young to bear the burden and heat
of this desperate life-struggle to the death.
But his heart gave a twinge, all the same, at the possibilities.
Con was possibly gone. Suppose these others went too! It would
leave a dreadful gap in their homes, and wounds in their hearts that
would never heal. This was what war meant. God help them all!
He watched the brave swing of the boys in hodden gray for a
time with approving eye, till they fell out to munch exiguous lunches
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