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Preface

Plants are the basis of life on earth as we know it and have not only shaped, to
a large extent, diverse ecosystems but also provide food, feed, fibers, and fuel
for human subsistence. Understanding how plants evolved and develop to
fulfill their manifold functions is thus of great fundamental and applied
interest.
It will soon be 10 years since the last volume of Current Topics in
Developmental Biology focused on plant development. Since then, biology
has seen a revolution in methods for the investigation of genes, transcripts,
and proteins, allowing a wide variety of “omics” approaches. Also, the tools
to investigate gene function using forward-genetic, reverse-genetic, and
alternative methods, such as artificial microRNAs and RNA interference,
have provided deep insights into the molecular processes that regulate plant
development. The arsenal of the plant biologist has recently been extended
by the CRISPR/Cas9 system, which allows functional studies also in non-
model systems, providing new insights into developmental diversity and
plasticity. Clearly, focusing efforts on a model system, in this case Arabidopsis
thaliana, has greatly accelerated progress in understanding plant function. On
the other hand, a tunnel vision focusing on a single species is bound to miss
important aspects of development that are not found in this model system.
Certainly, plant biology has profited a lot from the powerful genetics of Zea
mays (maize), which started well over a hundred years ago, and the recent
focus on Oryza sativa (rice) as one of the most important crop plants. Over
the last years, the range of plant species used in fundamental research has
greatly expanded, providing insights into diverse developmental processes.
Unraveling the molecular control of plant development has also
provided the material to look at the evolution of gene regulatory networks,
investigating how preexisting mechanisms were co-opted for new develop-
mental programs or how novelties arose during land plant evolution. Such
evo-devo studies have provided great insights into the evolution of the
enormous diversity of plants that exists today. It is more than timely then
to provide a broad overview of plant development and evolution, as it is only
possible in a multichapter book bringing together experts in diverse field to
share their views on specific plant developmental processes and their evolu-
tion. The chapters provide a wide diversity of perspectives on both vegeta-
tive and reproductive development, summarizing the deep functional and

xvii
xviii Preface

mechanistic insights that have been gained in many fields of development.

The chapters focusing on evolutionary aspects provide an excellent view on
the directions the evo-devo field will take in the future by being able to do
functional studies in nonmodel systems that promise to provide invaluable
insights into the evolution of plant development.
I was delighted that so many of my colleagues enthusiastically accepted to
contribute to this volume of Current Topics in Developmental Biology and I am
thankful for their detailed reviews and insightful discussions on a wide range
of topics in plant development and evolution. No doubt the next decade will
see enormous progress and further deepen our understanding of plant func-
tion. Nevertheless, I am convinced that the readers will find this volume
interesting and timely and hope they will enjoy reading it.
UELI GROSSNIKLAUS
Department of Plant and Microbial Biology & Zurich-Basel Plant
Science Center, University of Zurich, Zurich, Switzerland
 CHAPTER ONE

Evolution of the plant body plan

ter Szo
Pe € ve

nyi*, Manuel Waller, Alexander Kirbis
Department of Systematic and Evolutionary Botany & Zurich-Basel Plant Science Center, University
of Zurich, Zurich, Switzerland
*Corresponding author: e-mail address: [email protected]

Contents
1. The significance of land plants 2
2. Understanding evolution of the plant body plan 2
3. Phylogenetic relationships of land plants and evolution of the land plant
body plan 3
4. Developmental patterns predating the origin of land plants 5
5. Developmental innovations of land plants 7
5.1 Alternation of haploid (gametophyte) and diploid (sporophyte)
generations 7
5.2 Evolution of three-dimensional growth in the haploid and diploid phases 10
5.3 Origin of spores, sporangia, and sporopollenin in land plants 12
5.4 Origin of unbranched sporophyte forms 12
5.5 Evolution of bifurcating axes 14
5.6 Evolution of indeterminacy 15
5.7 Evolution of meristems 16
5.8 Origin of leaves 17
5.9 Evolution of rooting systems 20
5.10 Roots 22
6. Conclusions and perspectives 24
Acknowledgments 25
References 25

Abstract
Land plants evolved about 470 million years ago or even earlier, in a biological crust-
dominated terrestrial flora. The origin of land plants was probably one of the most sig-
nificant events in Earth’s history, which ultimately contributed to the greening of the
terrestrial environment and opened up the way for the diversification of both plant
and non-plant lineages. Fossil and phylogenetic evidence suggest that land plants have
evolved from fresh-water charophycean algae, which were physiologically, genetically,
and developmentally potentiated to make the transition to land. Since all land plants
have biphasic life cycles, in contrast to the haplontic life cycle of Charophytes, the
evolution of land plants was linked to the origin of a multicellular sporophytic phase.
Land plants have evolved complex body plans in a way that overall complexity

Current Topics in Developmental Biology, Volume 131 # 2019 Elsevier Inc. 1

ISSN 0070-2153 All rights reserved.
https://doi.org/10.1016/bs.ctdb.2018.11.005
2 P
eter Sz€
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enyi et al.

increased toward the tip of the land plant tree of life. Early forms were unbranched, with
terminal sporangia and simple rhizoid rooting structures but without vasculature and
leaves. Later on, branched forms with lateral sporangia appeared and paved the route
for the evolution for indeterminacy. Finally, leaves and roots evolved to enable efficient
nutrient transport to support a large plant body. The fossil record also suggests that
almost all plant organs, such as leaves and roots, evolved multiple times independently
over the course of land plant evolution. In this review, we summarize the current knowl-
edge on the evolution of the land plant body plan by combining evidence of the fossil
record, phylogenetics, and developmental biology.

1. The significance of land plants

Land plants evolved about 470 million years ago, or potentially even
earlier, in the Cambrian (515 Ma) to Early Ordovician (473 Ma) when the ter-
restrial flora was dominated by a crust-forming assemblage of algae lichens,
bacteria, cyanobacteria, and fungi (Berbee, James, & Strullu-Derrien, 2017;
Morris et al., 2018; Salamon et al., 2018). The origin of land plants was
one of the most significant events in the history of the globe, which paved
the road for diversification of non-plant lineages in the terrestrial environment
(Bateman et al., 1998; Berner, 1997). Among others, this was enabled by
an increasing oxygen concentration produced through photosynthesis, ini-
tially by cyanobacteria and then eukaryotic algae and land plants (Kenrick,
Wellman, Schneider, & Edgecombe, 2012). The resulting ozone layer
reduced the amount of UV irradiation, which was probably necessary for
plants to leave the water (Rozema et al., 2002; Rozema, Blokker, Mayoral
Fuertes, & Broekman, 2009). Furthermore, land plants have also significantly
contributed to the weathering of rocks and the formation of soil by preventing
transportation of sediments by wind and water (Lenton, Crouch, Johnson,
Pires, & Dolan, 2012; McMahon & Davies, 2018; Perkins, 2018). Altogether,
plants have tremendously contributed to the greening of the planet and to the
rapid increase of terrestrial biodiversity.

2. Understanding evolution of the plant body plan

Three major types of information can be utilized to investigate the
main trends in the evolution of the land plant body plan, and to uncover
the underlying genetic mechanisms. Phylogenetic analyses of extant land
plants can provide information on the ancestor-descendant relationships
of major lineages, and thus a rough timeline of their evolutionary origin.
Evolution of the plant body plan 3

Nevertheless, available phylogenetic evidence is mostly based on nucleotide

data and includes only extant plants, providing little information about the
combined evolutionary history of extant and fossil forms.
Another line of evidence is provided by the fossil data. Unfortunately,
fossil record of the first land plants is sparse, and their interpretation can
be ambiguous (Boyce & Kevin Boyce, 2010; Kenrick, 2018; Tomescu,
2009). Ambiguity in the assessment of morphological characters and their
potential homology with body plans of extant land plants make the evolu-
tionary placement of fossils questionable and combined analysis of data on
fossils and extant land plants is challenging (Bateman, 1996; Puttick et al.,
2018; Salamon et al., 2018; Stewart & Rothwell, 1993; Seward, 2011).
Despite this, careful comparative morphological and evolutionary analyses
of the fossil record and extant land plants gave rise to major theories about
the evolutionary origin of main modular units of the land plant body plan,
providing testable hypotheses (Chomicki, Coiro, & Renner, 2017).
Comparative analysis of fossils and phylogenetic relationships of extant
land plants is necessary to reveal evolutionary trends and formulate testable
hypotheses, but do not allow experimental validation. Comparative devel-
opmental biology and genetics/genomics of extant representatives of land
plants can be used to experimentally test hypotheses on the developmental
and genetic mechanisms underlying the evolutionary trends in land plant
body plan evolution (Boyce & Kevin Boyce, 2010; Rothwell, Wyatt, &
Tomescu, 2014; Tomescu, Wyatt, Hasebe, & Rothwell, 2014).
In this review, we provide insights into the recent advancement surround-
ing the evolution of the land plant body plan by evaluating evidence from
phylogenetics/phylogenomics, the fossil record, comparative genomics, and
the developmental biology of plants. We do so by summarizing recent
evidence in these scientific fields and their implications for each major inno-
vation in the evolution of the land plant body plan (see Fig. 1).

3. Phylogenetic relationships of land plants

and evolution of the land plant body plan
Phylogenetic analysis of extant members of green plants unambigu-
ously suggests that the lineage of land plants (embryophytes) together with
several lineages of streptophytic algae (charophycean algae) forms the mono-
phyletic group of streptophytes (Becker & Marin, 2009; Gitzendanner,
Soltis, Wong, Ruhfel, & Soltis, 2018; Liu, Cox, Wang, & Goffinet,
2014; Wickett et al., 2014; Wodniok et al., 2011). The streptophytes are
4 P
eter Sz€
ov
enyi et al.

Fig. 1 Hypothetical phylogenetic relationship of extant and fossil land plants and Char-
ophytes. The left bar shows the timescale of evolution. Names of extant groups are in
black while extinct taxa/lineages are shown in gray. Dashed horizontal lines represent
ambiguous phylogenetic relationships. Dotted vertical lines indicate the uncertainty in
the timing of the origin of embryophytes and lineages of bryophytes. Current data using
phylogenetic dating suggests that the deep splits in the tree may be considerably older
than depicted here (Morris et al., 2018). Pictograms show the architectural features of
major plant groups. Open elliptic structures refer to sporangia while gray filled structures
depict leaves. Redrawn, extended and modified from Kenrick, P., & Crane, P. (1997). The origin
and early diversification of land plants: A Cladistic Study. Science, 389(4), 33–39.

further embedded into the large clade of the green lineage (green plants
including green algae), the Viridiplantae. Three lineages of the paraphyletic
grade of streptophyte algae, the Charophyceae, Coleochaetophyceae, and
the Zygnematophyceae, are the closest relatives of land plants and share
the presence of phragmoplast with land plants, a structure central to the for-
mation of a new cell wall after cell division. Of the streptophyte algal line-
ages, the Zygnematophyceae appear to be the closest relative of land plants
(Delwiche & Cooper, 2015; Timme, Bachvaroff, & Delwiche, 2012;
Wickett et al., 2014; Wodniok et al., 2011). Extant Zygnematophyceae
consist of unicellular and filamentous algae, occupying freshwater and ter-
restrial habitats, whose developmental features are very difficult to compare
with the complexity of land plant body plans (Delwiche & Cooper, 2015).
Evolution of the plant body plan 5

Therefore, having the Zygnematophyceae as sister to land plants makes recon-

struction of the character states of the shared common algal ancestor with land
plants particularly challenging (de Vries & Archibald, 2018). Altogether, the
sister relationship of Zygnematophyceae with land plants provides very little
information on the potential order and extent of evolutionary transformations
that led to the evolution of the complex body plans of early land plants.
Phylogenetic analysis of extant land plants (embryophytes) implies that
land plants consist of the clade of vascular plants, which include the monophy-
letic groups of lycophytes, monilophytes, gymnosperms and angiosperms,
and a group of three lineages, the mosses, liverworts, and hornworts, collec-
tively referred to as bryophytes (Cox, Li, Foster, Martin Embley, & Civáň,
2014; Gitzendanner et al., 2018; Liu et al., 2014; Wickett et al., 2014;
Wodniok et al., 2011). The phylogenetic inter-relationships of the bryophyte
lineages and their relationship to vascular plants are still highly debated. For
instance, earlier analyses, mainly based on molecular data, recovered all pos-
sible topologies between the three lineages of bryophytes and the rest of land
plants. Either mosses, hornworts, or liverworts were reconstructed as sister to
the rest of the land plants, and the three lineages of bryophytes were mainly
resolved as a paraphyletic grade with some exceptions (Bremer, Humphries,
Mishler, & Churchill, 1987; Chang & Graham, 2011; Cox et al., 2014; Finet,
Timme, Delwiche, & Marl
etaz, 2010; Fiz-Palacios, Schneider, Heinrichs, &
Savolainen, 2011; Karol, 2001; Laurin-Lemay, Brinkmann, & Philippe, 2012;
Mishler & Churchill, 1984; Qiu et al., 2006; Wodniok et al., 2011; Zhong,
Liu, Yan, & Penny, 2013). Conversely, recent phylogenomic analyses impli-
cate that mosses and liverworts are very likely monophyletic (Gitzendanner
et al., 2018; Liu et al., 2014; Morris et al., 2018; Puttick et al., 2018;
Wickett et al., 2014; Wodniok et al., 2011). Furthermore, some analyses give
strong support to the monophyly of bryophytes revealing the deepest split
between hornworts and a clade consisting of the liverworts and mosses
(Cox et al., 2014; Gitzendanner et al., 2018; Liu et al., 2014; Wickett
et al., 2014; Wodniok et al., 2011; Zhong et al., 2013).

4. Developmental patterns predating the origin

of land plants
Recent evidence suggests that the genetic basis for cellular processes that
allowed plants to colonize and successfully cope with the terrestrial environ-
ment were already present in the charophycean algae and, therefore, their
evolution predated the origin of land plants (de Vries & Archibald, 2018;
6 P
eter Sz€
ov
enyi et al.

de Vries, Curtis, Gould, & Archibald, 2018; Harholt, Moestrup, & Ulvskov,
2016; Selosse, Strullu-Derrien, Martin, Kamoun, & Kenrick, 2015). Similarly,
it was found that most transcription factor families, key regulators of
developmental processes thought to be specific to land plants, were already
present in the charophycean algae (Catarino, Hetherington, Emms,
Kelly, & Dolan, 2016; Wilhelmsson, M€ uhlich, Ullrich, & Rensing, 2017).
This suggests that some developmental innovations relevant to land plant evo-
lution may have been already present in the charophycean algae.
Charophycean algae are highly diverse in their development. For
instance, some are unicellular, but others have attained multicellularity with
thallose or filamentous forms (Delwiche & Cooper, 2015; Domozych,
Popper, & Sørensen, 2016). They may or may not show apical growth,
with some groups exhibiting complex apical growth and branching. There-
fore, developmental patterns such as apical growth, branching, and
multicellularity were present already in the charophycean algae. It is assumed
that some of these developmental patterns, and likely the corresponding
mechanisms, were retained through the evolution of early land plants. For
instance, the thallose gametophytes of liverworts resemble those of the char-
ophytes, and the underlying developmental mechanisms may have been pre-
sent and retained from their common ancestor (Ligrone, Duckett, &
Renzaglia, 2012a; Renzaglia, Duff, Nickrent, & Garbary, 2000). Similarly,
the presence of both unicellular and multicellular forms in charophytes sug-
gests that developmental processes for multicellularity might have been pre-
sent in the common ancestor of charophytes and land plants and retained.
Intriguingly, some experiments suggest that the multicellular-unicellular
transition can be achieved in moss protonema by targeting genes involved
in protein prenylation. Such mutants have unicellular cells undergoing unpo-
larized divisions and resemble those of unicellular algal cells (Antimisiaris &
Running, 2014; Thole, Perroud, Quatrano, & Running, 2014). Further-
more, recent experiments with the alga Chlamydomonas reinhardtii suggest that
multicellularity can be relatively easily attained under certain selection regimes
or by altering the regulation of cell cycle genes (Boyd, Rosenzweig, & Herron,
2018; Hanschen et al., 2016; Herron, 2016; Niklas, 2014; Olson & Nedelcu,
2016; Ratcliff et al., 2013). The observation that the basic genetic tool kit of
land plants is present in the genome of the charophyte Klebsormidium flaccidum,
and that most key transcription factor families are present in various char-
ophycean algae, further suggest that a set of developmental mechanisms and
their genetic regulators were already present in the common ancestor of
charophytes and land plants (Hori et al., 2014; Wilhelmsson et al., 2017).
Evolution of the plant body plan 7

Altogether, this implies that the common ancestor of land plants and char-
ophytes was genetically and likely developmentally prepared to achieve the
complex body plan of land plants through relatively few evolutionary steps.
Nevertheless, except some studies on MADS box genes (Tanabe et al.,
2005), information on the genetic regulation of specific developmental pro-
cesses of charophytes is lacking, and future research will provide information
about the potential homology of basic developmental mechanisms in land
plants and charophytes.

5. Developmental innovations of land plants

Transition to the terrestrial environment brought some radical changes
to the basic body plan of plants. The appearance of biphasic life cycles, spo-
ropollenin and sporangia, three-dimensional growth and simple unbranched
sporophytes appeared very early in land plant evolution.

5.1 Alternation of haploid (gametophyte) and diploid

(sporophyte) generations
All charophycean algae had unicellular ancestral forms and a life cycle in which
the haploid phase dominates, and the syngamy of gametes is immediately
followed by meiosis of the zygote (Bowman, Sakakibara, Furumizu, &
Dierschke, 2016; Niklas & Kutschera, 2009; Qiu, Yin-Long, Taylor, &
McManus, 2012). Such life cycles are haplontic because all mitotic divisions
occur in the haploid phase, and the diploid phase is only represented by a sin-
gle cell, the zygote. The body plan of land plants radically broke this pattern
by evolving multicellular alternating haploid (gametophytic) and diploid
(sporophytic) phases. Relative dominance of the two phases has also changed
during the course of land plant evolution, in such a way that the haploid phase
dominated early in evolution, followed by the elaboration of the diploid and
reduction of the haploid phase later in evolution (Bowman et al., 2016;
Niklas & Kutschera, 2009; Qiu et al., 2012). Therefore, the origin of land
plants is linked to the evolution of the multicellular sporophytic phase.
Current phylogenetic evidence, consistently resolving charophycean algae
with a purely haplontic life cycle as sister to all land plants, gives overwhelming
support to this assertion. This finding is consistent with the antithetic theory
proposing that land plants evolved from an algal ancestor with a haplontic life
cycle with zygotic meiosis. In parallel, phylogenetic evidence rejects the
homologous theory of Bower, assuming an algal ancestor with an isomorphic
alternation of haploid and diploid generations. Altogether, current evidence
8 P
eter Sz€
ov
enyi et al.

implies that the multicellular sporophyte was an evolutionary innovation of

land plants and originated by the intercalation into the ancestral haplontic life
cycle of mitotic divisions in the zygote prior to meiosis (Bower, 1890;
Haig, 2008).
Therefore, the major difference between the haplontic life cycles of
charophytes and the biphasic life cycles of land plants is whether the zygote
proceeds to meiosis without mitotic divisions or it proliferates mitotically
prior to meiosis. Intensive research on this topic over the last years suggests
that part of the genetic toolkit responsible for this developmental switch seems
to be deeply rooted in the history of the green lineage (Bowman et al., 2016).
Theory further suggests that the evolution of this toolkit may date back to the
origin of green algal mating types, which likely evolved to impose a stringent
control on the timing of developmental switches (Wang & Dohlman, 2005).
The molecular toolkit controlling these critical developmental aspects of the
biphasic life cycle of land plants seems to have evolved by co-opting
the genetic network controlling meiosis in the zygote of simple unicellular
algae, such as C. reinhardtii (Bowman et al., 2016; Lee, Lin, Joo, &
Goodenough, 2008). In C. reinhardtii, heterodimerization of the BEL1-LIKE
(BELL) homeodomain transcription factor GSP1 with the KNOTTED1-
LIKE HOMEOBOX (KNOX) family transcription factor GSM1 is necessary
to initiate zygotic gene expression and meiosis (Bowman et al., 2016; Lee
et al., 2008). Homologs of both BELL and KNOX gene families are present
in all land plants and they do heterodimerize (Floyd & Bowman, 2007;
Frangedakis, Saint-Marcoux, Moody, Rabbinowitsch, & Langdale, 2017;
Hay & Tsiantis, 2010; Horst et al., 2016). KNOX and BELL function is best
studied in the moss Physcomitrella patens, in which a single BELL gene,
PpBELL1, is sufficient and necessary to induce the sporophytic program.
It is also known that KNOX genes went through a duplication prior to
the origin of land plants, giving rise to Class 1 and Class 2 KNOX genes
(Floyd & Bowman, 2007; Frangedakis et al., 2017; Furumizu, Alvarez,
Sakakibara, & Bowman, 2015; Hay & Tsiantis, 2010). The Class 1 KNOX
gene of P. patens, MKN2, is necessary for regulating sporophyte development
in the moss, while the Class II KNOX genes appear to suppress the gameto-
phytic program in the sporophytic phase (Sakakibara et al., 2013; Sakakibara,
Nishiyama, Deguchi, & Hasebe, 2008; Sano et al., 2005). It is not yet
clear with which KNOX protein PpBELL1 heterodimerizes with (Horst
et al., 2016). Conversely, components of the Polycomb Repressive
Complex 2 (PRC2), such as the homologs of the Arabidopsis thaliana
proteins FERTILIZATION-INDEPENDENT ENDOSPERM (PpFIE)
Evolution of the plant body plan 9

and CURLY LEAF (PpCLF), suppress the sporophytic program in the game-
tophytic phase (Mosquna et al., 2009; Okano et al., 2009; Pereman et al.,
2016). Furthermore, transcriptomic data suggest early activation of meiosis-
related genes in the apical cells of the sporophyte in the moss P. patens
(Frank & Scanlon, 2015a, 2015b). Nevertheless, the link between the gene
networks regulating the haploid-diploid switch and that of sporangial
development is unknown. These findings suggest that an ancient regulatory
network, whose evolution coincided with the origin of mating types in green
algae, is deeply conserved across the green lineage. The putative ancestral
function of this network was to enable a stringent control on the developmen-
tal switch between haploid and diploid programs (Bowman et al., 2016).
This network was then used, and its regulatory role was extended, to govern
various aspects of sporophytic development in land plants.
Although the core regulatory network controlling initiation of the sporo-
phyte and gametophyte developmental programs is known, there is very little
information available about the genetic program enabling multicellularity and
three-dimensional growth in the sporophytic phase. It is possible that these
developmental mechanisms were partly recruited from the gametophytic
phase (Frank & Scanlon, 2015a, 2015b; Szovenyi, Rensing, Lang, Wray, &
Shaw, 2010). Alternatively, they could have evolved de novo in land plants.
It is known that zygotes of double mutants disrupting the FLORICAULA/
LEAFY (FLO/LFY) homologs PpLFY1 and PpLFY2 arrest and are unable
to divide mitotically (Tanahashi, Sumikawa, Kato, & Hasebe, 2005). Further-
more, P. patens zygotes lacking activity of the two WUSCHEL-RELATED
HOMEOBOX 13-LIKE (PpWOX13LA/B) genes are unable to elongate and
initiate the apical cell of the embryo (Sakakibara et al., 2014). Transcriptomic
evidence from P. patens suggests that multicellularity and three-dimensional
patterning may have been, at least partially, recruited from gametophytic pro-
grams (Frank et al., 2015; Frank & Scanlon, 2015b; Whitewoods et al., 2018).
Nevertheless, many aspects of the evolution of the sporophytic phase are still
unclear, and the interconnection of reproductive and proliferative programs is
unknown.
Although, multiple lines of evidence support the origin of land plants
from a charophycean ancestor, there is much ambiguity surrounding the
evolutionary relationship of land plant lineages. The debate about the phy-
logenetic relationship of bryophyte lineages and their relationship with
vascular plants has significant impact on how land plant body plan changes
are interpreted. This includes the evolution of the biphasic life cycle and the
evolution of other morphological traits, such as stomata, vascular tissues, etc.
10 P
eter Sz€
ov
enyi et al.

For instance, studies suggesting a paraphyletic relationship of the three

bryophyte lineages (Chang & Graham, 2011; Qiu et al., 2006, 2012) are com-
patible with a haploid-dominant ancestral land plant life cycle, which was
retained in the paraphyletic grade of bryophytes but was followed by the
dominance of the diploid phase in vascular plants (Bowman et al., 2016;
Haig, 2008; Niklas & Kutschera, 2009; Qiu et al., 2012; Tomescu et al.,
2014). In contrast, a monophyletic bryophyte clade may suggest that the
haploid-dominant life cycle could have been a unique innovation of the bryo-
phyte clade (Cox et al., 2014; Gitzendanner et al., 2018; Puttick et al., 2018;
Wickett et al., 2014). According to this hypothesis, life cycle of the ancestral
land plant could have been haploid-dominant, diploid-dominant, or equally
dominant with close to isomorphic haploid and diploid phases. Furthermore,
it is possible that the common ancestor of all land plants may have been more
trachaeophyte-like, which may explain the origin of conducting tissues in
mosses. The earliest fossil remains of land plants are currently interpreted as
being stem trachaeophytes (Kenrick, 1994, 2018; Kenrick & Crane, 1997;
Taylor, Kerp, & Hass, 2005). Nevertheless, it is possible that some of these
fossils represent the common ancestor from which the monophyletic group
of bryophytes evolved (Boyce & Kevin Boyce, 2010; Kenrick, 2018;
Puttick et al., 2018). Therefore, there is considerable uncertainty surrounding
the evolutionary origin of land plants, which makes reconstruction polarity of
key characters in land plants challenging.

5.2 Evolution of three-dimensional growth in the haploid

and diploid phases
Most charophycean algae grow in a planar form along a two-dimensional axis,
while land plants evolved three-dimensional growth, enabled by the presence
of a continuously rotating division plane in the stem cells (Delwiche &
Cooper, 2015; Domozych et al., 2016; Langdale, 2008). The ability of flexibly
changing division planes is missing from the algal relatives of land plants. The
genetic mechanisms underlying this evolutionary transition can only be stud-
ied in land plants with a life cycle including both two- and three-dimensional
growth patterns.
Importantly, some mosses exhibit a filamentous juvenile life cycle phase
(protonemata) of the gametophyte, which is maintained by two-dimensional
growth as seen in charophycean algae. After this stage, a change to three-
dimensional growth may occur, giving rise to leafy shoots. This system is
suitable to investigate the genetic mechanisms enabling the transition from
Evolution of the plant body plan 11

two- to three-dimensional growth. Nevertheless, this approach assumes that

the ontogeny of the moss P. patens recapitulates the evolutionary trajectory
that occurred between algae and land plants (Harrison, Roeder,
Meyerowitz, & Langdale, 2009). There are multiple genetic factors known
to affect initiation of the three-dimensional growth pattern in P. patens.
The NO GAMETOPHORES1 (PpNOG1) gene regulates the transition to
three-dimensional growth by inducing the degradation of proteins that likely
repress the P. patens APB transcription factors, belonging to the AP2 class tran-
scription factors homologous to A. thaliana AINTEGUMENTA, PLETH-
ORA, and BABY BOOM (APB). The PpAPB transcription factors are
necessary and sufficient to initiate three-dimensional buds (Aoyama et al.,
2012; Moody, Kelly, Rabbinowitsch, & Langdale, 2018) The PpNOG1 gene
is likely also necessary in directing division planes in the apical cell, together
with the DEFECTIVE KERNEL1 homolog PpDEK1, to achieve proper
three-dimensional growth (Demko et al., 2014; Olsen, Perroud,
Johansen, & Demko, 2015; Perroud et al., 2014). PpNOG1 and related genes
occur only in land plants; therefore, it may be one of the key factors that reg-
ulate this innovation (Moody et al., 2018). Given that PpDEK1 and AP2 class
transcription factors regulate similar processes in flowering plants and the
moss, it is possible that this ancient network was already present in the com-
mon ancestor of land plants, and that its evolution coincided with the evolu-
tion of three-dimensional growth. Finally, homologs of the CLAVATA
(CLV) pathway, crucial for meristem maintenance in the flowering plant
shoot apical meristem, were recently shown to be critical to the correct ori-
entation of division planes in the transition from two- to three-dimensional
growth in P. patens (Whitewoods et al., 2018). The role of the CLV pathway
in orienting division planes seems to be conserved between A. thaliana and
P. patens and may represent the ancient function of this pathway. It is thus
possible that the CLV pathway was recruited to regulate the proper orienta-
tion of division planes in two independent contexts: in the apical cell of the
moss gametophyte and in the shoot apical meristem of flowering plants. Phe-
notypic effects of PpDEK1, PpNOG1 and the moss homologs of the CLV
pathway are overlapping, suggesting that they are likely members of a gene
network governing the proper development of the gametophyte apical cell
in P. patens. In line with these findings, transcriptomic evidence also suggests
that the genetic mechanisms involved in the evolution of three-dimensional
growth in gametophytes may have been partially recruited to support three-
dimensional patterning in the sporophytic phase (Frank & Scanlon, 2015a).
12 P
eter Sz€
ov
enyi et al.

5.3 Origin of spores, sporangia, and sporopollenin

in land plants
All land plant spores are covered by sporopollenin, whereas algae only have
heavy-walled zygotes that germinate via meiosis. Therefore, sporopollenin
coated spores are an innovation specific to land plants. This assertion is in line
with the finding that the earliest fossils with land plant affinities are the
so-called cryptospores (470 million years old), whereas sporangial fragments
are younger (450 million years old) (Edwards, Morris, Richardson, &
Kenrick, 2014; Steemans et al., 2009). Cryptospores occur either in monads,
diads, or in tetrads (see chapter “Evolution and co-option of developmental
regulatory networks in early land plants” by Bowman et al., this issue). Their
wall structure resembles that of some liverwort spores, such as the diads and
tetrads of the genera Haplomitrium and Sphaerocarpos, respectively, but their
affinity to extant bryophyte species is highly debated (Edwards, Richardson,
Axe, & Davies, 2012; Gensel, 2008; Renzaglia et al., 2015). It is thought that
sporopollenin-covered spores evolved earlier than sporophytes, likely via
modification of the timing of meiosis and the deposition of sporopollenin,
which significantly affected spore viability (Brown & Lemmon, 2011).
Genetic pathways and their components are partially conserved across land
plants, but conservation of the genetic components underlying the develop-
mental process and its evolutionary trajectory is not known (Harrison,
Alvey, & Henderson, 2010).

5.4 Origin of unbranched sporophyte forms

The earliest non-spore fossils assigned to land plants are believed to resemble
present day vascular plants with bifurcating axes (Bowman, 2013; Boyce &
Kevin Boyce, 2010; Kenrick, 2018; Kevin Boyce, 2005; Gensel, 2008;
Langdale, 2008). Nevertheless, phylogeny and the earliest fossils of stem
group polysporangiophytes suggest that early sporophytes may have been
unbranched, with terminal sporangia such as in extant mosses. Furthermore,
the earliest sporophytes are believed to be obligate matrotrophic (nutrition-
ally supported by the gametophyte), with a well-developed photosynthetic
gametophyte phase (Gensel, 2008; Kenrick, 2018; Qiu et al., 2012; Remy,
Gensel, & Hass, 1993). Therefore, it is hypothesized that fossils of the earliest
multicellular unbranched sporophyte forms are missing from the fossil
record, as all known fossils show some bifurcation. Nevertheless, their small
size and the lack of vascularization correspond to the organizational level
of bryophyte sporophytes (Boyce & Kevin Boyce, 2010; Gensel, 2008;
Evolution of the plant body plan 13

Gerrienne et al., 2006; Remy et al., 1993; Taylor et al., 2005). It is assumed
that these unbranched forms are retained in extant bryophytes, especially in
the lineages of mosses and hornworts, which can be used to gain information
about their developmental biology and evolution. Unfortunately, develop-
ment of the sporophyte in the three lineages of bryophytes is strikingly dif-
ferent and, therefore, it is difficult to establish which features are ancestral to
the group (Ligrone et al., 2012a; Ligrone, Duckett, & Renzaglia, 2012b)
(Fig. 2).
Importantly, elongation of the liverwort sporophyte is almost exclusively
due to cell expansion, and there is no apical or localized meristem activity with
divisions occurring all over the sporophyte (Ligrone et al., 2012a; Renzaglia
et al., 2000). In contrast, sporophyte growth is dominated by cell divisions in
both hornworts and mosses, involving the activity of well-localized intercalary
meristems and/or apical cells (Ligrone et al., 2012a; Villarreal & Renzaglia,
2015). Hornwort and moss sporophytes share the presence of a so-called mul-
ticellular proliferative zone within the sporophyte (Ligrone et al., 2012b).
This proliferative zone produces cells upward, giving rise to the full body
of the sporophyte in hornworts, whereas it produces cells downward in

Fig. 2 Schematic representation of sporophyte development in the three lineages of

bryophytes. The bottom row shows the three cell layers (basal, middle and upper) of
the early embryo. The upper row depicts the developing sporophytes. Actively dividing
cells are shown in gray and arrows refer to the direction of cell production. Inactive but
preformed regions are shown in black. Randomly arranged arrows in the liverwort
embryo symbolize cell divisions occurring across the whole embryo. Gray arrow heads
in the developing sporophyte of liverworts depict cell elongation and a lack of cell divi-
sions leading to the elongation of the seta.
14 P
eter Sz€
ov
enyi et al.

mosses, giving rise to the stalk (seta) of the sporophyte (French & Paolillo,
1975b; Villarreal & Renzaglia, 2015). Developmental homology of these
structures and their relation to the probably ancient uniaxial structures is
unexplored. Nevertheless, historical experiments of moss sporophytes suggest
that cytokinin may be involved in regulating activity of the intercalary meri-
stem. Retarded growth of the sporophyte after decapitation can be alleviated by
exogenous cytokinin application, which may represent a conserved mechanism
with the angiosperms shoot apical meristem (SAM) (French & Paolillo, 1975a,
1975b; Ligrone et al., 2012b; Snipes et al., 2018; Truskina & Vernoux, 2018).
It is not yet clear, how developmental mechanisms of unbranched forms
are linked to the origin of branched shoots of early vascular plants. There are
two competing hypotheses explaining the evolutionary origin of early vascular
shoot systems, the interpolation/sterilization and apical growth hypotheses
(Bowman, 2013; Ligrone et al., 2012b; Tomescu et al., 2014). The apical
growth hypothesis assumes that early shoots evolved from the extended mer-
istematic activity of the transient apical cell of a moss-type embryo. In contrast,
the interpolation/sterilization hypothesis suggests that the common ancestor
of bryophyte and vascular plant sporophytes had apical reproductive activity,
which went through a sterilization process to produce the axis of the shoot.
Further comparative study on the unbranched sporophytes of bryophytes and
that of vascular plants may clarify whether any of these hypotheses can be
supported by experimental data.

5.5 Evolution of bifurcating axes

The earliest vascular plant remains had independent gametophyte and spo-
rophyte generations, with bifurcating sporophyte axes that terminated in
sporangia. This is in contrast to the unbranched and matrotroph sporophytes
of extant mosses (Gensel, 2008; Kenrick, 2018; Remy et al., 1993; Taylor
et al., 2005; Tomescu et al., 2014). Indeterminate sporophytes appear only
much later in the fossil record, implying that evolution of bifurcation pre-
dates that of indeterminacy. Information on how branching evolved is
provided by observations on fern and bryophyte sporophytes. In P. patens
plants, deletion either of the TEOSINTE BRANCHED1/CYCLOIDEA/
PROLIFERATING CELL FACTOR1 (TCP) transcription factor class
gene PpTCP5, the TERMINAL EAR1-LIKE (TEL) gene PpTEL, or
PINFORMED1 (PIN1) homolog PpPINB gene will increase the proportion
of bifurcating sporophytes compared to the wild type (Bennett et al., 2014;
Ortiz-Ramı́rez et al., 2016; Vivancos et al., 2012). PpFLO/LFY mutants have
Evolution of the plant body plan 15

a similar phenotype (Tanahashi et al., 2005). At present, it is unclear when

exactly branching takes place and whether it occurs at the zygote or at the
sporophyte-seta stage. It is also unclear whether the effect of these genes is
achieved via similar developmental mechanisms. Observations on fern sporo-
phytes show that branching is achieved by segregation and amplification of
stem cells at the shoot apex (Harrison, Rezvani, & Langdale, 2007). Trans-
criptomic evidence suggests that genetic components necessary for this process
are present in fern genomes and shoot apices, but the genetic mechanisms are
still unclear (Evkaikina et al., 2017; Frank et al., 2015; Harrison, 2015). In
contrast, gametophytic branching in mosses is regulated by the co-option
of ancient hormonal effects involved in sporophyte branching in angiosperms,
such as auxin, cytokinins, and strigolactons (Coudert, Bell, Edelin, &
Harrison, 2017; Coudert et al., 2015). Polar auxin transport is essential for
branching in flowering plants but in the moss bi-directional transport is
required for normal branching (Coudert, 2017; Harrison, 2017).

5.6 Evolution of indeterminacy

The fossil record suggests that all indeterminate sporophyte axes have later-
ally arranged sporangia (Boyce & Kevin Boyce, 2010; Tomescu et al., 2014).
Therefore, the evolution of indeterminacy of sporophytic axes and the
lateral displacement of sporangia are linked, indicating spatially and tempo-
rally separated activity of reproductive and vegetative functions (Kenrick,
2018). Information on the molecular mechanisms underlying indeterminacy
is exclusively coming from investigations on the determinant sporophyte of
P. patens. It was shown that two components of the moss PRC2, encoded by
PpFIE and PpCLF, are necessary to repress the meristematic activity of the
sporophyte apical cell (Kenrick, 2018; Mosquna et al., 2009; Pereman et al.,
2016). In mutants disrupting PRC2, the sporophytes produced branched
structures by continuous proliferation. Class I KNOX genes are known
to be responsible for the maintenance of the meristematic activity of the spo-
rophyte apical cell (Sakakibara et al., 2008). That is, the interaction of PRC2
and Class I KNOX gene activity is key in the regulation of determinant and
indeterminant growth. Class II KNOX genes are also expressed in and nec-
essary for the development of the sporophyte by repressing the gameto-
phytic program (Sakakibara et al., 2013). These observations imply that
function of the PRC2 in repressing pluripotent sporophytic cells is con-
served across land plants. Whether the antagonistic effect of Class I and II
KNOX genes seen in flowering plants is also conserved in the development
16 P
eter Sz€
ov
enyi et al.

of the sporophyte of mosses is unknown (Furumizu et al., 2015). Neverthe-

less, it is possible that evolution of indeterminacy is partially achieved by the
proper coregulation of Class I and Class II KNOX activities, which may
connect reproductive and vegetative proliferative activities.

5.7 Evolution of meristems

Plant form and architecture are tightly linked to the activity of meristems.
Indeterminate meristems have independently evolved in the gametophyte
phase of bryophytes and ferns, and in the sporophyte of vascular plants.
Gametophyte meristems considerably differ from those of sporophyte mer-
istems, because the former are composed of a single stem cell, while the latter
consist of a single stem cell, or a group of cells, overlaying several layers of
proliferative cells (Ambrose & Vasco, 2016; Harrison et al., 2007, 2009;
Langdale, 2008; Sanders, Darrah, & Langdale, 2011). Meristematic activity
is not recognizable in the fossil record of early vascular plants. Furthermore,
fossils at the divergence of vascular plants and bryophytes are missing, which
makes it impossible to decide whether sporophytic indeterminant meristems
have evolved once or multiple times independently. Therefore, all informa-
tion is coming from observations made on extant taxa.
Among the extant representatives of the earliest diverged lineages, mosses,
hornworts and some liverworts have multicellular proliferative sporophytic
regions, which may be—to some extent—homologous to proliferative regions
of the indeterminate meristematic regions of extant vascular plants (Bowman,
2013; Ligrone et al., 2012b; Tomescu et al., 2014). Importantly, hornwort
basal meristems seem to be indeterminate, while moss intercalary meristems
show only transient activity (Bowman, 2013; French & Paolillo, 1975b;
Langdale, 2008; Ligrone et al., 2012a, 2012b; Villarreal & Renzaglia, 2015).
It remains to be seen whether these structures are homologous to one another,
and whether their proliferative activity shares common regulatory activity with
that of the proliferative regions of vascular plant meristems.
The structure of indeterminate meristem activity of sporophyte shoots in
lycophytes and ferns is highly variable, but it usually consists of a single apical
cell, or groups of cells, overlaying a deeper layer of proliferative cells
(Ambrose & Vasco, 2016; Frank et al., 2015; Vasco et al., 2016). Expression
of key genes of the SAM in ferns and lycophytes suggests that their sporo-
phytic meristems are multicellular structures, and their core regulatory
mechanisms may be homologous to that of the flowering plant SAM
(Evkaikina et al., 2017; Friedman, 2011). Nevertheless, transcriptomic
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after them. Yet, even in this rude way, so large a force contrived to
establish itself among the sand-hills by the evening of the 24th, that
all apprehension of a renewed attack from the enemy was laid aside.
Nevertheless the artificers were anxiously pressed to render the
bridge trustworthy with as little delay as possible; and they
strenuously exerted themselves to meet the wishes of the general.
Meanwhile, about fourteen or twenty thousand men of the Spanish
army took post along those faces of the town and citadel which
looked towards Helletre and the Joyeuse. The left of this semicircular
line, resting upon the heights where, during the late affair, I stood in
safety to watch the progress of the skirmishers on both sides of me,
swept round, through the abandoned intrenchments, to the brink of
the river. Here, the stream being narrow, a pontoon-bridge was
formed, and the line recommencing on the opposite bank, wound on
till it touched a corps of Portuguese at the back of the citadel. But as
yet the chain was continued from that point only by occasional
patrolling parties; and through this opening the enemy daily sent out
his foragers and brought in supplies. Such a state of things could not
be permitted long to last. It was essential to the prosecution of Lord
Wellington's future operations that the gap should be filled up
previous to the renewal of hostilities between his army and that of
Soult; nor was much time wasted in making preparations for driving
the garrison within the walls. Working parties laboured hard, not
only during the day, but during the whole night of the 24th; and at
dawn on the 25th it was reported that infantry might cross the
floating bridge with safety. This was the signal for action; and hence
the 25th proved to be, at least to part of the army, a day of hostile
employment.
 CHAPTER XXII.
A direct communication between the opposite banks of the river
being thus established, the remaining battalions of the Guards, the
chief part of the King's German Legion, together with a
proportionate force of cavalry and artillery, marched at daybreak on
the 25th to join their comrades among the sand-hills. The whole of
the besieging army being at the same time put in motion, the gap
which prior to this date had existed in the line of investment was
filled up. Little or no fighting took place on that occasion. The
enemy, perceiving our design, offered no serious resistance, but
evacuating the village of Boucaut, after having exchanged a few
shots with our skirmishers, established their pickets about half a mile
in its rear. As yet, therefore, a good deal more of open space was
granted to them than they could long hope to enjoy; but all
opportunity of corresponding with Marshal Soult, as well as of
adding to the stock of grain and provisions already in their
magazines, was cut off.
The running and irregular fire which had been maintained
throughout the morning gradually died away, and ceased altogether
about noon. From that hour till after nightfall everything continued
quiet. A feverish excitement, necessarily consequent even upon a
trifling skirmish, prevailed indeed amongst us; nor did we venture to
take off our accoutrements, or return to our usual employments,
during the remainder of the day. But we might have done so, had
we felt disposed, with perfect safety; because the enemy were too
well satisfied with being permitted to retain what they still held of
territory beyond the glacis to endanger its loss by a useless attempt
to regain what had been wrested from them. Still we were anxious;
and the anxiety which pervaded us all the day ceased not to operate
at night.
The garrison of Bayonne, we were well aware, was at once
numerically powerful and composed of the best troops in the French
army. From all that we could learn, Soult had by no means
calculated upon the plan of operations adopted by Lord Wellington.
Concluding that the Marquess would halt after the passage of the
Adour, and invest that important place with the whole of his forces,
he had thrown into it fifteen thousand picked men, assigning the
command to General Thouvenot—an officer who, by his successful
defence of Burgos on a former occasion, appeared worthy of so
delicate a trust on the present. Lord Wellington, however, knew too
well how much depended in war on celerity of movement to waste
his time under the walls of Bayonne. He therefore left Sir John Hope
to mask the place with two British divisions, and taking the
remainder with himself, hung upon the rear of the retreating enemy.
These two divisions, which composed his left wing, were indeed
supported by a considerable force of Spaniards. But not even now
could much reliance be placed on Spanish troops; though it is just to
add, that they were much upon the alert at the outposts, and
patient under privations and hardships. The task assigned to Sir
John Hope was not therefore an easy one. With some thirty
thousand men of all arms, of whom only one-half might be fully
trusted, he drew his lines of circumvallation round a well-fortified
town, the enceinte of which could not measure less than four
English miles, and towards which he did not as yet venture to push
his pickets nearer than one mile, or perhaps more, from the glacis.
It was felt by him, and indeed by all, that vigilance could not under
the circumstances be too keen or too constant. The besieged,
moving upon an interior line, had it in his power to throw at any
moment a superior force upon the besiegers; hence, especially for
the first few days and nights, officers and men alike kept
themselves, so to speak, constantly on the stretch of expectation.
On the 25th, for example, as all seemed to be quiet in front, we lay
down at the usual hour and slept. Our camp was pitched under the
lee of a sand-hill; and just over its brow, and at the base beyond,
our pickets were posted. It was still early, an hour or two from
midnight, when a musket-shot in the direction of the pickets roused
us. There was no time to consider, because the enemy, if a sortie
was intended, would be upon us in a moment; so we sprang from
our pallets, and each dressing hastily, and seizing his weapons, we
ran to the place of muster. And now another and another shot broke
the stillness of the night. The bugles began to sound, the baggage
was hastily packed, the horses were saddled, and all the bustle and
hurry attendant upon the preparations for battle took place. For
myself, having seen that my men were in their ranks, I ran to the
top of the hill, whence I saw the flashes of several muskets half-way
between our sentinels and those of the enemy; but no sound of
advancing columns met my ear, neither was the fire returned by our
own soldiers. The perplexity occasioned by this state of affairs was
not, however, of long continuance. The officer in command of the
outposts sent in a messenger to say that no symptom of an attack
was discernible, but that several deserters had come into his lines, at
whom the French sentries were firing. This account was confirmed
soon afterwards by the arrival of the deserters in the camp; and the
troops accordingly laid aside their weapons and returned to their
tents.
The alarm in that direction had hardly subsided, when another and
not less serious one arose in a different quarter. A sentry who was
posted by the bank of the river reported to his officer when visiting
him that boats were moving and oars splashing in the water.
Apprehensions were immediately excited for the safety of the bridge,
against which we naturally concluded that some attempt was about
to be made. To oppose it as far as possible, of whatever nature it
might be, three field-pieces which were attached to our brigade
limbered up, and galloped to the water's edge. These I
accompanied; and certainly the splash of oars was very audible,
though the darkness would not permit us to distinguish whence the
sound proceeded. A few shots were, however, fired in the direction
of the sound, just by way of hinting to the enemy that we were
awake; and whether it was that the hint was not lost upon them, or
that they never seriously entertained the idea of assailing the bridge,
an immediate cessation of rowing was the consequence. Having
watched, therefore, for half an hour, and neither hearing nor seeing
anything indicative of danger, I left the gunners to themselves, and
returning to my cloak and blanket, wrapt myself closely up, and slept
soundly and securely till the morning.
The whole of the 26th passed over without the occurrence of any
event worthy of mention. By myself it was spent, not very profitably,
in sauntering about among the pine-woods, where little or no game
was to be found. For the troops in general, as well within as without
the walls of the beleaguered city, it might be accounted a sort of
armed truce. Hardly a cannon-shot was fired, from sunrise till
sunset, on either side; but matters were drawing to a crisis. Stores
and ammunition were conveyed day and night across the river in
large quantities; and it was manifest that even the few miles of open
country which the garrison still held would before long be taken
away from them. It was therefore no unexpected communication
which reached me on the morning of the 27th, that the corps was to
stand to its arms forthwith, and that the enemy were to be driven in
all directions within their works.
Having in a former chapter described the nature of the ground in our
immediate front, the reader will understand why no serious advance
on our part was intended. We were already within range of the guns
on the ramparts, and between the ramparts and the camp no broken
ground nor village, nor any other species of cover, existed. We could
not therefore hope to establish ourselves had we been pushed on;
whereas the French general, by opening the sluices from the river,
might at any moment lay the whole level under water. On the
opposite side of the Adour the case was different. There the most
forward British pickets were very little in advance of the village of
Boucaut; and the village of Boucaut is full four miles from the
citadel. The face of the country also between the two points being
rugged and broken, numerous positions could be taken up by the
besiegers, in which, whilst they were themselves secure from the fire
of the place, they could easily prevent the garrison from venturing
beyond the ditch. Moreover, the relative situations of the town and
fortress rendered the former secure against active annoyance till
after the latter should have fallen into our hands. Though, therefore,
it was understood that the whole of our line was to be drawn
somewhat more tightly round the city, we were all aware that the
trenches would be opened, and breaching batteries thrown up
against the citadel alone.
The men being accoutred and the baggage packed, we stood quietly
in our ranks behind the sand-hill, till a gun from the opposite side of
the stream sounded the signal of attack. Upon this we extended our
files so as to give to a single weak battalion the appearance of an
entire brigade, and, ascending the heights, we stopped short where
the tops of our bayonets and caps just showed themselves over the
ridge. Similar demonstrations were likewise made by the corps which
filled Anglete and crowned the rise in connection with it; whilst
occasionally a shout was raised, as if at length the order of attack
had been given, and we were preparing to rush on. All this was done
for the purpose of drawing the attention of the enemy to many
different points at the same time, and thus hindering them from
opposing with the entire strength of the garrison the forward
movements of those who were appointed to invest the castle.
Whilst we and the divisions near us were thus amusing ourselves
and the enemy with the pomp and circumstance, rather than with
the reality, of war, the Guards and light Germans, with a corps of
Portuguese infantry, were very differently occupied on the other
bank of the river. As our situation was a commanding one, it enabled
us to obtain a tolerably distinct view of their proceedings. We saw
one column of British troops form on the sands beside Boucaut. In
front of it was a body of German riflemen, who pressed leisurely
forward in skirmishing order till they reached a picket of the French
troops. Of the enemy, on the other hand, a heavy column showed
itself upon the high ground, where it halted, and continued to send
out numerous parties to support the outposts, between whom and
the Germans a hot skirmish began. But it could not be said that any
decided advantage was gained by either party during several hours.
The column which we descried upon the sands beside Boucaut was
not of great strength; indeed the numbers of our own people
discernible by us were very inconsiderable. The fact, as I afterwards
learned, was, that the side of the hill visible to us was by far the
most rugged and least assailable of any; consequently the main
attack was to be made in another direction, the attack in this waiting
till the other should have in part succeeded. Hence the trifling
progress made by our skirmishers, who seemed to be kept back
rather than animated forward by their officers; and hence the
apparently obstinate resistance of the French pickets. But it was,
nevertheless, an exceedingly interesting spectacle, to the beauty of
which the uneven and picturesque scenery around added not a little.
I wish I could convey to the mind of the reader some notion of the
scene as it then appeared, and is still remembered by myself. Let
him imagine himself, then, lying with me upon the brow of a sand-
hill, and looking down first upon the broad and deep waters of the
Adour, and over them upon a sandy bank, which speedily ends, and
is succeeded by a green hill, having on its side—the side upon which
we are gazing—frequent cuts or gullies, or glens, some of them
bare, others wooded, with here and there a white cottage showing
itself from among the trees. Let him imagine that he sees on the
summit of the heights, and immediately in a line with himself, a
portion of an armed mass, with a single field-piece pointed towards
the river's mouth. About a mile to the rearward let him figure to
himself a green field, more level than any other part of the hillside—
a sort of table-land, as it were, having a hedge along that face of it
which is turned towards Boucaut, and a precipitous red bank under
the hedge. In this field he will observe about three hundred infantry
soldiers dressed in grey greatcoats and broad caps or shakos, who
carry hairy knapsacks on their backs, and are armed with long clear
muskets, which have bayonets screwed to their muzzles. These are
Frenchmen. Under the red bank let him farther suppose that there is
a picturesque valley stocked with tall and shadowy cork-trees, about
the middle of which is a neat mansion something larger than a
farmhouse, yet hardly deserving the name of a chateau. That house
is full of light Germans; and almost every tree about it affords cover
to a rifleman, who fires as a good aim is presented to him at the
persons behind the hedge. From the windows of the house, likewise,
many shots are from time to time discharged; and the sudden flash
and uprising of smoke from various parts of the hedge show that the
French tirailleurs are not less active than their assailants, or disposed
to receive their salute without returning it. In this skirmish little
change of ground takes place. Occasionally, indeed, a single rifleman
will steal on, running from tree to tree, till he has reached a
convenient spot; whilst a Frenchman will as often rise, and having
watched him through a brake or over a bush, will fire whenever he
exposes himself to observation. But no grand rush is made on either
side, nor is any decided loss sustained either of ground or in men.
All this while the exertions of our people were, as far as might be,
aided by a well-served cannonade from the three pieces of artillery
which had kept their station near the bank of the river since the
evening of the 25th. The fire of these guns was directed chiefly
against a large house—apparently some public work or manufactory
—which stood by the brink of the water, and was filled with French
troops. Neither were the enemy's batteries opposite to us idle.
Having wasted about twenty or thirty round-shot without effect, they
brought a couple of mortars, with a howitzer or two, to bear upon
us, from which they threw shell after shell among our ranks. But
from the effects of the cannonade the nature of the soil secured us,
the shells either burying themselves in the sand to the extinction of
the fuze, or exploding when we were all snugly laid flat, and
therefore safe from their fragments.
Matters had continued thus for an hour or two, and we were
beginning to fear that some part of our General's plan might have
gone wrong, or that the enemy were in too great force to be driven
in by the divisions opposed to them, when a sudden stir in the
French column which had hitherto stood quietly upon the heights
attracted our attention. The field-piece was all at once wheeled
round, and turned in the direction of the opposite country; the
infantry collected into compact order, and were gradually hidden
from us by the brow of the hill. By-and-by a few musket-shots were
fired; then about a dozen more; then came the report of one, two,
or three field-pieces; and, lastly, a roar of cannon and small-arms.
This was kept up, hot and rapid, for half an hour. Every moment the
sound came nearer and nearer. Now the smoke, which had at first
followed each report after the interval of a few seconds, rose at the
same instant with the noise; then the glancing of arms over the high
ground was distinguishable; next came the French troops, some
retiring slowly, and firing as they fell back—others fleeing in extreme
confusion. Mounted officers were galloping over the ridge, and
apparently exerting themselves to restore order; but all would not
do. The enemy were in full flight. Down they rushed towards the
river, and away along the sands in the direction of the citadel, whilst
our three guns poured in round-shot among them, some of which
we could distinctly perceive take effect. And now the green field on
which my reader and I have so long looked was abandoned. The
tirailleurs fled—the riflemen pursued—the little column in scarlet
pushed on in good order and with a quick pace—and on the brow of
the height above a British ensign was held up as a signal for our
battery to cease firing. The signal was obeyed, and we had nothing
farther to do during the remainder of the day than to watch, which
we eagerly did, the progress of our victorious comrades.
The enemy having fled as far as the manufactory, were there joined
by reinforcements from the garrison. Here, then, the battle was
renewed with great obstinacy; but stern as was the resistance
offered, it became every hour less and less effectual. At length the
building took fire; it was abandoned, and its defenders fled; after
which the entire scene of action was hidden from us, and we were
enabled to guess at the state of affairs only by the sound of firing
and the direction which it took. That inclined every moment more
and more towards the ramparts. But it continued without
intermission till darkness had set in, when both parties were
compelled to desist because they could not distinguish friends from
foes.
In this affair the loss on both sides was considerable; but we were
completely successful. The enemy were driven within their works,
and our advanced-posts were established in the village of St Esprit,
about half-pistol-shot from the nearest redoubt. In other directions
little change of ground occurred. Some Spanish divisions took up a
position, I believe, somewhat less distant than formerly from the
walls of Bayonne; but neither we nor the divisions in communication
with us were in any degree affected by it. We returned, on the
contrary, to our tents, having lost by the cannonade only one man
killed and three wounded.
I stated in another part of my narrative that, except on one
occasion, I could not tax my memory with any symptom of violent or
permanent grief on the part of a soldier's wife at the death of her
husband. The case to which I then alluded occurred to-day. A fine
young Irishman, the pay-sergeant of my own company, had brought
his wife with him to the seat of war. He married her, it appeared,
against the wish of her relatives, they considering themselves in a
walk of life superior to his. To what class of society they belonged I
cannot tell; but she, I know, was a lady's-maid to some person of
rank, when the handsome face and manly form of M'Dermot stole
her heart away. They had been married about a year and a half,
during the whole of which time she had borne the most unblemished
character, and they were accounted the happiest couple in the
regiment. Poor things! they were this day separated for ever.
M'Dermot was as brave and good a soldier as any in the army; he
was at times even foolhardy. Having observed a recruit or two cower
down in no very dignified manner as a cannon-ball passed over
them, M'Dermot, by way of teaching them to despise danger, threw
himself at his ease on the summit of the sand-hill, with his head
towards the enemy's guns. He was in the act of laughing at these
lads, assuring them that "every bullet has its billet," when a round-
shot struck him on the crown of the head and smashed him to
atoms. I shall never forget the shriek that was raised. He was a
prodigious favourite with all ranks; and then all of us thought of his
poor young wife, so spotless, and so completely wrapped up in him.
"Oh, who will tell Nance of this?" said another non-commissioned
officer, his principal companion. "Poor Nance!" cried the soldiers, one
and all; so true is it that virtue is respected, and a virtuous woman
nowhere more beloved than among British soldiers. But there was
no hiding it from Nance. The news reached her, heaven knows how,
long before we returned to our tents, and she was in the midst of us
in a state which beggars all description in five minutes after the
event took place.
I cannot so much as attempt to delineate the scene that followed.
The poor creature was evidently deranged, for she would not believe
that the mangled carcass before her was her husband; and she
never shed a tear. "That! oh, that is not he!" cried she; "that
M'Dermot!—my own handsome, beautiful M'Dermot! Oh no, no—
take it away, or take me away, and bring me to him!" She was
removed with gentle violence to the camp, and the body was buried,
a young fir-tree being planted over it.
Several days passed before Mrs M'Dermot was sufficiently calm to
look her situation in the face. But at last the feeling of utter
desolation came over her; and instead of listening, as women in her
position generally do, to the proposals of some new suitor, all her
wishes pointed homewards. To her home she was accordingly sent.
We raised for her a handsome subscription, every officer and man
contributing something; and I have reason to believe that she is now
respectably settled in Cork, though still a widow.
 CHAPTER XXIII.
From the date of the occurrences just described—namely the 27th of
February—the siege of Bayonne may be said to have fairly
commenced. To follow in regular detail the events of each day as it
proceeded would not, I am sure, greatly interest my readers; and to
lay such detail before them would be to myself an occupation little
less irksome than it sometimes was to kill the tedious hours of a ten
weeks' blockade. I may be permitted, therefore, to state generally,
and in few words, that the strictest investment was continued all the
while, and that an extremely harassing kind of duty was imposed
upon us till the siege and the war were brought to an end together
by the hoisting of the white flag on the 28th of the following April.
Premising this, I shall merely take the liberty of narrating, without
regard to dates or natural order, such incidents and adventures as
appeared to myself best deserving of record at the time.
In the first place, then, it may be observed that, while on our side of
the river no other works were erected than such as appeared
indispensable for strengthening our own position, and rendering the
bridge and the highroad, and the stores brought up by them, secure,
the Guards and Germans on the other side were busily employed in
digging trenches, and pushing forward active operations against the
citadel. These, as may be imagined, they were not permitted to
carry on without being annoyed in every practicable manner by the
besieged. A continual, or rather a dropping and irregular fire of
cannon, was kept up upon their parties from the ramparts, to which
even the darkness of the night brought no cessation; for blue-lights
were thrown out wherever the people were at work, the flame of
which guided the artillerymen in taking aim. Nor were we wholly
exempt from that species of entertainment. On the contrary, as the
erection of a three-gun battery on the top of our hill was deemed
necessary, we worked at it by turns till it was completed; and, as a
matter of course, we worked under the fire of all the cannon and
mortars which could be brought to bear upon us. These working
parties are by far the most unpleasant of all the employments to
which a soldier is liable. There is in them nothing of excitement, with
a great deal of danger; and danger, where there is no excitement,
no man would voluntarily choose to incur for its own sake. Let me
describe one of these mornings' amusements.
It fell to my lot frequently to superintend the people when at work.
The spot on which we laboured was high, and therefore completely
exposed to the view of the enemy. It was the top of the hill opposite
to them. Immediately on our arrival, a four-gun battery, with one
howitzer and two nine-inch mortars, began to play upon us. They
were well served, for the balls hit apparently in every quarter except
the particular spots on which each of us stood. On such occasions, if
there be no very pressing demand for the completion of the work,
you generally station one of your party to watch. As soon as he
perceives a flash he calls out—"Shot," or "Shell," as the case may
be. If it be simply a cannon-shot, you either toil on without heeding
it, or having covered yourself as well as you can till the ball strike,
you start up again and seize your tools. If it be a shell, you lie quite
still till it burst. The unmilitary reader may perhaps question whether
it be possible to tell the nature of the missile which is coming against
you, when as yet it has barely escaped from the muzzle of the gun,
and is still a mile or more distant; but he who has been in the habit
of attending to these matters will entertain no such doubt. Not to
mention the fact that an experienced eye can trace, by means of the
burning fuze, the whole journey of a shell through the air, from its
expulsion till its fall, the more perpendicular flight of the smoke may
of itself inform him who takes the trouble to observe when it issues
from a mortar; and there is a sharpness in the report of a gun which
will effectually distinguish the one from the other, even if the sense
of sight should fail. I have heard men assert that they can trace, not
only a shell, but a cannon-ball through the air. This may be possible;
but if it be, it is possible only to those whose sense of sight is far
more acute than mine.
Though abundantly annoying, the cannonade of which I am
speaking proved but little destructive of human life. I do not believe,
though continued for several weeks, it cost us five men. Neither was
an attempt which the enemy by-and-by made to shell us out of our
position more successful. Hidden from their view by the sand-hills,
we lay exposed only to such aim as a calculation of distances might
enable their gunners to take; and this, besides that it was not always
very accurate, had, in the nature of the soil on which our tents
stood, much to defeat it. Shells falling among loose sand bury
themselves so far into the ground as to take away a good deal from
the force of their destructiveness. Probably the French artillerists
guessed, from the perfect indifference with which we treated them,
that they were wasting their ammunition. At all events, they
slackened their fire by degrees, and by-and-by restrained it
altogether.
Unless my memory greatly deceive me, the chief subject of
complaint amongst us at this time was that we were fettered to one
spot, and that there was not in our situation peril or excitement
enough to hinder us from feeling the confinement as a grievous
restraint. Though tolerably secure, from the nature of the ground,
our post was one of vast importance—that is to say, had the enemy
succeeded in forcing it, they might have easily made their way to the
bridge ere any fresh troops could be brought to oppose them. Under
these circumstances, it was considered imprudent to wander far or
frequently from the tents; and hence even the resource of fishing
and shooting was in a great measure taken away from us. My friend
and I did indeed occasionally venture into the woods; but these
excursions were too rare to be very profitable, and our limits too
confined to furnish an abundance of game.
The reader is not, however, to suppose that all our days and night
partook of the same tame character. Independently of the usual
round of outpost duty (a duty which, to me at least, was never
irksome, because it always served to keep my interest awake), a
deserter would, from time to time, come over, and bring with him
rumours of sorties intended. One of these brought with it a more
than ordinary degree of excitement. We were sitting one Sunday
evening, Grey and myself, in the upper loft of an old mill, where, by
way of an indulgence, we had established ourselves. The
commanding officer had read prayers to the battalion about half an
hour previously, and the parade had just been dismissed, when a
sergeant clambered up the ladder to inform us that the servants and
batmen were commanded to sleep accoutred, that the horses were
to be saddled, and the baggage in readiness to move at a moment's
notice. On inquiring into the cause of this order, we learned that a
French officer had arrived in camp, that he had brought with him
intelligence that a sortie would take place a little before midnight,
and that the garrison were already making preparations for the
attack. As may be supposed, we put everything in proper trim
forthwith; and having seen that our men lay down, with knapsacks
buckled up, and pouches and bayonets slung on, we too threw
ourselves on the floor in our clothes.
It might be about eleven o'clock when we were startled from our
repose by the firing of cannon. The sound was, however, distant: it
evidently came from the opposite side of the river, and it was
followed by no musketry. We watched it, therefore, for a while,
anxiously enough, and sat up prepared to issue forth as soon as our
presence might be required. But no bugle sounded, nor was any
other summons given; so we lay down again, and the night passed
by in peace. I have reason to believe, however, that the French
officer had not deceived us. An attack upon our position had been
seriously intended, and the plan was abandoned only because this
officer being missed, it was conjectured that we should be fully
prepared to repel it.
Another little affair took place soon afterwards. Whether our posts
on the left of Anglete had been of late pushed somewhat more in
advance than formerly I cannot tell; but the enemy sent a message
one morning by a flag of truce to the officer in command, desiring
that he would fall back, otherwise they would compel him. The
demand was of course met by a peremptory refusal, and they,
having allowed him an hour to change his mind, proceeded to carry
their threat into execution. A considerable body of light troops
attacked the post, and a sharp skirmish ensued. The sound of firing
soon drew assistance to our picket; and the result was, that the
French once more retired within their works, leaving us in possession
of the disputed ground. This event, with many others which I have
not recorded, because they have in them even less of interest,
occurred during the remaining days of February and the whole of
March. On the 1st of April our position was changed, and we took
from that period a more active part in the conduct of the siege.
The change of ground to which I now allude proved, at least for a
day or two, extremely agreeable to the corps in general. My friend
and myself had, indeed, as I have already stated, fixed our abode in
an old mill—close to the camp, yet sufficiently apart from it to be
freed from the bustle. It was a ruinous and dilapidated mansion, I
admit; our living and sleeping chamber consisting simply of one half
of a loft, and only of one half, because the flooring of the other half
had given way; to which we ascended by means of a ladder or trap-
stair, and from which we looked down upon our horses and mules
that occupied the basement story. But in that old mill, the tiling of
which was unsealed, and can hardly be said to have been proof
against the weather, I spent some weary and many more pleasant
evenings; for, ruinous as it was, it appeared comfortable to men who
repaired to it from the sandy ground on which they had previously
spent several days and nights under cover of the canvas. Though,
therefore, I cannot accuse myself of murmuring at the removal of
the camp, it is quite certain that I partook not in the general
rejoicing which the occurrence produced among my comrades; or
that the beauty of the spot to which my tent was transferred at all
compensated for the loss of a boarded floor and a detached
habitation.
It was, however, a delightful change to the majority. During the last
week or ten days the heat of the sun had become exceedingly
oppressive, beating, as it did, through the white canvas, and having
its rays reflected back on all hands from a grey sandy soil. Not a tree
grew near to shelter us; nor was there a blade of grass within sight
on which the weary eye could repose. On the 1st of April we retired
about a couple of miles into the heart of a pine-wood, and left the
sand-hill to be guarded by the pickets alone. Our tents were pitched
in a sweet little green vale, overshadowed with the dark foliage of
the fir-trees, and near the margin of a small lake or pond of clear
water. Here we remained in a state of comparative idleness and
enjoyment for three days—running and leaping, and causing the
men to run and leap for prizes—till an order arrived in the evening of
the 3d, that we should be under arms at daybreak on the morrow,
and cross the bridge to take part in the fatigues and dangers of the
trenches.
At an early hour on the 4th we formed into marching order and took
the direction of the bridge. This we crossed, the planks waving and
bending as the cables swung to and fro with our tread; and then
filing to the right, we halted in an open field above the village of
Boucaut, where the ground of encampment was marked out. It was
a day of heavy rain, so we were thoroughly saturated by the way;
and as several hours elapsed ere the baggage came up, we were
compelled to continue in that uncomfortable plight all the while. It
came at length, however, and our tents were pitched; after which,
having substituted dry for wet apparel, I spent the rest of the
evening in lounging among the numerous stalls and booths which
surrounded the market-place.
The village of Boucaut presented at this period a curious spectacle.
It was not deserted by its inhabitants—all, or the greater number of
whom, remained quietly in their houses. Their little shops were not
closed; the inns—for there were two in the place—so far from being
abandoned, were thronged with customers; cooks, waiters, landlady,
and mine host were all in motion from morning till night; and crowds
of peasantry came and went all day long laden with eggs, butter,
cheese, poultry, and other luxuries. These articles of merchandise
were exposed for sale in the centre of the market-place—a large
square surrounded by high walls, along the sides of which sutlers'
tents, porter booths, confectioners' stalls, and even tables loaded
with hardware, shoes, stockings, &c., were laid out in regular order.
The place was, moreover, full of people—soldiers, camp-followers,
villagers, peasants, male and female; and much laughing and great
merriment prevailed in every direction. To a mere spectator there
was constant food for amusement; in the fruitless endeavours of an
English soldier, for example, to make love to a pretty French girl; or
in the vain efforts of a staid German to overreach some volatile but
mercenary villager; whilst the ceaseless gabbling in all European
tongues—the attempts made on all hands to carry on by signs that
conversation to which the faculty of speech lent no assistance;—to
watch these, and a thousand other extravagances, furnished ample
and very agreeable employment to one who was willing to find
amusement where he could. Yet with all this apparent confusion the
greatest regularity prevailed. Not a single instance of violence to a
native, either in person or property, occurred; indeed, both men and
women scrupled not to assure us that they felt themselves far more
secure under our protection than they had been while their own
countrymen were among them.
It was our business, so long as the camp stood here, to move up
every morning to the front, and to work in turns at the erection of
batteries and redoubts, within half musket-shot of the walls of the
citadel. The spot where I usually found myself stationed when my
turn of duty came round, was a chateau situated upon the brow of
an eminence, from the windows and garden of which I obtained a
distinct view of one flank of the castle. Upon this building an
incessant fire of round-shot, shells, grape, and occasionally of
musketry, was kept up. The enemy had upon their walls a number of
long swivel-guns, which they could elevate or depress, or turn in any
direction at will, and with which as perfect an aim could be taken as
with an ordinary carbine. These threw with great force iron balls of
about a quarter of a pound weight. Beside them men were always
stationed, who watched our movements so closely that it was
impossible to show so much as your head at a window, or over the
wall, without being saluted by a shot; whilst from time to time a
nine-inch shell would tumble through the roof, and burst sometimes
before we had time to escape into another apartment. Then the
crashing of the cannon-balls as they rushed through the partitions—
the occasional rattle of grape and canister which came pouring in by
the windows,—all these things combined, produced a species of
feeling of which no words can convey an adequate notion to him
who has not experienced it. It was not terror—it can hardly be called
alarm, for we followed our occupations unceasingly, and even our
mirth was uninterrupted; but it kept the mind wound up to a pitch of
excitation from which it was by no means an unpleasant matter to
relieve it.
Ours was a mortar-battery. It was formed by heaping up earth
against the interior of the garden wall, and proceeded with great
rapidity. We likewise cut down trees, and constructed out of their
branches fascines and gabions; but we had nothing to do in the
trenches. Of these, indeed, not more than a couple were dug, the
uneven nature of the ground producing numerous valleys and
hollows, which saved us a great deal of toil, and very sufficiently
supplied their place.
Besides taking charge of working parties, it came occasionally to my
turn to command a picket. The post of which I was put in charge
was the village of St Esprit, and the church formed the headquarters
of the guard. It was a small building, but, fortunately for us,
constructed with great solidity, inasmuch as it stood under the very
muzzles of half-a-dozen field-pieces, which the enemy had placed in
a redoubt about a short stone's-throw distant. To add to its strength,
and to render it more tenable in case of attack, an embankment of
earth—of earth carried from the churchyard, and so mixed with the
mouldering bones of "the rude forefathers of the village"—was
raised inside, to the height of perhaps four feet, above which ran a
line of loopholes, cut out for the purpose of giving to its garrison an
opportunity of firing with effect. When I say that the church formed
the headquarters of the guard, I mean that the guard took up its
station there during the night. So long as daylight lasted the men
kept as much as possible concealed behind a few houses in the rear
of the building, and left only a single sentinel there to watch the
movements of the enemy.
A little to the right of my post were a couple of barricades, the one
cutting off the main road, the other blocking up the entrance to a
cross-street in the village. Beside these respectively stood a six-
pounder gun. They were, I should conceive, about long pistol-shot
from the walls of the castle, and formed our most advanced stations.
Our line of sentinels ran through the churchyard and streets, winding
away by the right and left as the shape of the place required; and
they were planted as close to one another as the occurrence of
trees, or other species of cover, would permit. For the French were
no longer the magnanimous enemy we had found them in the open
field. Every man, no matter whether a sentry or a lounger, who
could be seen, was fired at; nor could the reliefs be carried on as
under ordinary circumstances. No corporal's party marched round
here, but the men themselves stole one by one to the particular
spots allotted to them, those whom they came to relieve stealing
away after a similar fashion. Yet even thus we seldom returned to
the camp without bringing a wounded man or two back with us, or
leaving a dead comrade behind.
At night the utmost vigilance was necessary. The enemy were so
close to us that the slightest carelessness on our part would have
given them free and secure access through our chain—a
circumstance which rendered it impracticable for the vedettes to give
sufficient warning to men who should not be at every moment in a
state of preparation. No man slept, or so much as lay down. The
privates stood round the embankment within the church, as if they
had been all on watch; the officer crept about from place to place in
front of it, or listened with deep anxiety to every sound. In these
wanderings the conversation of the French soldiers could be
distinctly overheard, so near were the troops of the two nations to
each other, and so perilous, or rather so momentous, was the duty
which we were called upon to perform.
 CHAPTER XXIV.
The blockade of Bayonne being now decidedly converted into a
siege, Sir John Hope very justly determined that every brigade of
British and Portuguese troops—in other words, every brigade upon
which he could at all depend—should take by turns a share in the
fatigue and danger attendant upon the progress of operations. The
tour of duty allotted to each was accordingly fixed at three days. In
consequence of this arrangement, we, who had assumed the care of
the works and outposts on the 4th, were relieved on the evening of
the 7th; and, at an early hour on the morning of the 8th, once more
turned our faces in the direction of the pine-wood. The tents which
we had pitched in the vicinity of Boucaut were not, however, struck.
These we left standing for the benefit of a brigade of Portuguese,
which crossed the river to succeed us; and hence, instead of halting
where we originally sojourned beside the pond, and under the
shadow of the fir-trees, we pushed on as far as the outskirts of
Anglete. The morning of the 8th chanced to be uncommonly dark
and foggy. It so happened, moreover, that a man who had got drunk
upon duty the night before was doomed to suffer punishment as
early as circumstances would allow; and the battalion, having
reached what was supposed to be its ground, formed square in a
green field for the purpose. Partly in consequence of the density of
the fog, which rendered all objects at the distance of fifty yards
invisible, and partly because the country was altogether new to us,
we lost our way. Our astonishment may therefore be conceived,
when, on the clearing away of the mist, we found ourselves drawn
up within less than point-blank range of the enemy's guns, and close
to the most advanced of our own sentinels in this part of the line.
For a moment or two we were permitted to continue thus
unmolested, but not longer. The breastworks in front of us were
speedily lined with infantry; mounted officers arrived and departed
at speed; a few field-pieces, being hurried through a sally-port, were
posted upon the exterior of the glacis, and then a sharp cannonade
began. It was quite evident that the enemy expected an assault; and
the accidental appearance of two other British brigades, which
chanced at the moment to pass each other in our rear, added
strength, without doubt, to that expectation. The scene was highly
animating; but the enemy's guns were too well served to permit our
continuing long spectators of it. A ball or two striking in the centre of
the square warned us to withdraw; and as we were clearly in a
situation where we were never meant to be, as well as because no
act of hostility was on our part intended, we scrupled not to take the
hint, and to march somewhat more to the rear. There a certain
number of houses was allotted to us, and we again found ourselves,
for the space of four days, under cover of a roof.
We were thus situated when a messenger extraordinary arrived at
the quarters of the commanding officer about midnight on the 11th
of April, with intelligence that the Allies were in possession of Paris,
and that Buonaparte had abdicated. It would be difficult to say what
was the effect produced upon us by the news. Amazement—utter
amazement—was the first and most powerful sensation excited. We
could hardly credit the story; some of us even went so far for a while
as to assert that the thing was impossible. Then came the thought of
peace, of an immediate cessation of hostilities, and a speedy return
to our friends and relatives in England; and last, though not with the
least permanent influence, sprang up the dread of reduction to half-
pay. For the present, however, we rather rejoiced than otherwise at
the prospect of being delivered from the irksome and incessant
labour of a siege; and we anticipated with satisfaction a friendly
intercourse with the brave men against whom we had so long
fought, without entertaining one rancorous feeling towards them. I
fear, too, that the knowledge of what had passed in Paris caused
some diminution in the watchfulness which we had hitherto
preserved; at least I cannot account upon any other principle for the
complete surprisal of our outposts in the village of St Esprit a few
nights subsequently.
The messenger who conveyed this intelligence to us went on to say
that Sir John Hope had despatched a flag of truce to inform the
Governor of Bayonne that there was no longer war between the
French and British nations. General Thouvenot, however, refused to
credit the statement. He had received, he said, no official
communication from Marshal Soult; and as he considered himself
under the immediate command of that officer, even a despatch from
the capital could have no weight with him unless it came backed by
the authority of his superior. Under these circumstances no
proposals were made on either side to cease hostilities, though on
ours the troops were henceforth exempted from the labour of
erecting batteries, in which it was very little probable that guns
would ever be mounted. In other respects, however, things went on
as they had previously done. The pickets took their stations as
usual; all communications between the garrison and the open
country was still cut off; and several families who sought to pass
through our lines were compelled to return into the town. This last
measure was adopted, as it invariably is when a city is besieged, in
order not to diminish the number of persons who must be fed from
the stores laid up in the public arsenals.
Though there was peace in Paris, there was no peace before
Bayonne. Our brigade having enjoyed its allotted period of rest,
prepared to return to its camp beside Boucaut, for which purpose a
line of march was formed on the morning of the 12th; and we again
moved towards the floating bridge. As yet, however, our services at
the outposts were not required; and as working parties were no
longer in fashion, we spent that and the succeeding day peaceably
in our camp. Not that these days were wholly devoid of interesting
occurrences. During the latter a French officer arrived from the
north, bearing the official account of those mighty transactions
which once more placed his country under the rule of the Bourbons;
and him we sent forward to the city, as the best pledge that could
be offered for the truth of our previous statements, and of our
present amicable intentions. Still General Thouvenot disbelieved, or
affected to disbelieve, the whole affair; but he returned an answer
by the flag of truce which accompanied the aide-de-camp, "that we
should hear from him on the subject before long."
It will be readily believed that the idea of future hostilities was not,
under all these circumstances, entertained by any individual of any
rank throughout the army. For form's sake, it was asserted that the
blockade must still continue, and the sentinels must still keep their
ground; but that any attack would be made upon them, or any blood
uselessly spilled, no man for a moment imagined. The reader may
therefore guess at our astonishment, when, about three o'clock in
the morning of the 14th, we were suddenly awoke by a heavy firing
in front; and found, on starting up, that a desperate sortie had taken
place, and that our pickets were warmly engaged along the whole
line. Instantly the bugles sounded. We hurried on our clothes and
accoutrements. The horses came galloping in from their various
stables; the servants and batmen busied themselves in packing the
baggage; and then hastily taking our places, we marched towards
the point of danger, and were hotly and desperately in action in less
than a quarter of an hour.
The enemy had come on in two columns of attack—one of which
bore down upon the church and street of St Esprit; while the other,
having forced the barricade upon the highroad, pressed forward
towards the chateau where our mortar-battery was in progress of
erection. So skilfully had the sortie been managed, that the sentries
in front of both these posts were almost all surprised ere they had
time, by discharging their pieces, to communicate an alarm to those
behind them. And hence it came to pass that, amid the obscurity of
a dark night, the first intimation of danger which the pickets received
was given by the enemy themselves, who, stealing on to the very
edge of the trench within which our men were stationed, fired down
upon them. In like manner the sergeant's guard which stood beside
the gun in the village was annihilated, and the gun itself captured;
whilst the party in the church were preserved from a similar fate
only in consequence of the care which had been taken to block up
the various doorways and entrances, so that only one man at a time
might make his way into the interior. It was, however, surrounded
and placed in a state of siege, and gallantly defended by Captain
Forster of the 38th Regiment and his men.
Just before the enemy sallied out, a French officer, it appeared, had
deserted. Unfortunately, however, he came in through one of the
more remote pickets, and hence those which were destined to
receive the shock reaped no benefit from the event. His arrival at
headquarters had, however, the effect of putting Sir John Hope on
his guard; so that greater preparations to meet the threatened
danger were going forward than we, on whom it came unexpectedly
and at once, imagined. Five hundred men, for example, who formed
a sort of reserve about a mile in rear of the outposts, were in full
march towards the front when the firing began; and the enemy were
in consequence checked before they had made any considerable
progress, or had reached any of our more important magazines. The
blue house, as we were in the habit of naming the chateau, was
indeed carried; and all the piles of fascines and gabions, which had
cost so much labour to construct, were burned; but besides this,
little real benefit would have accrued to the assailants had the state
of affairs been such as to render a battle at this particular juncture
at all necessary or even justifiable.
Immediately on the alarm being given, Sir John Hope, attended by a
single staff-officer, rode to the front. Thither also flew Generals Hay,
Stopford, and Bradford; whilst the various brigades hurried after
them at as quick a pace as the pitchy darkness of the night and the
rugged and broken nature of the ground would permit. Behind them,
and on either hand, as they moved, the deepest and most
impervious gloom prevailed; but the horizon before them was one
blaze of light. I have listened to a good deal of heavy firing in my
day, but a more uninterrupted roar of artillery and musketry than
was now going on I hardly recollect to have encountered.
As the attacking party amounted to five or six thousand men, and
the force opposed to them fell somewhat short of one thousand, the
latter were, of course, losing ground rapidly. The blue house was
carried—the highroad, and several lanes that ran parallel with it,
were in possession of the enemy—the village of St Esprit swarmed
with them,—when Sir John Hope arrived at the entrance of a hollow
road, for the defence of which a strong party had been allotted. The
defenders were in full retreat. "Why do you move in that direction?"
cried he, as he rode up. "The enemy are yonder, sir," was the reply.
"Well, then, we must drive them back—come on." So saying, the
General spurred his horse. A dense mass of French soldiers was
before him; they fired, and his horse fell dead. The British picket,
alarmed at the fall of the General, fled; and Sir John being a heavy
man—being, besides, severely wounded in two places, and having
one of his legs crushed beneath his horse—lay powerless, and at the
mercy of the assailants. Captain Herries, his personal attendant on
this occasion, did his best to extricate Sir John from his dilemma. He
sprang from his horse, strove to drag the dead animal aside, and
would not listen to the entreaties of his chief, who besought him to
look to his own safety; and he paid the penalty of his devotion. The
French continued to advance. They fired another volley at the
retreating Guardsmen, a shot from which broke the bone of Captain
Herries' leg close to the knee-joint. The young man fell, and was
removed, together with his wounded General, a prisoner into the
city.
Of this sad catastrophe none of the troops were at all aware, except
those in whose immediate presence it occurred. The rest found
ample employment, both for head and hand, in driving back the
enemy from their conquests and bringing succour to their comrades,
whose unceasing fire gave evidence that they still held out in the
church of St Esprit. Towards that point a determined rush was made.
The French thronged the street and churchyard, and plied our
people with grape and canister from their own captured gun. But the
struggle soon became more close and more ferocious. Bayonets,
sabres, the butts of muskets, were in full play; and the street was
again cleared, the barricade recovered, and the gun retaken. But
they were not long retained. A fresh charge was made by increased
numbers from the citadel, and our men were again driven back.
Numbers threw themselves into the church as they passed, among
whom was General Hay; whilst the rest gradually retired till
reinforcements should come up, when they resumed the offensive,
and with the most perfect success. Thus was the street of St Esprit,
and the field-piece at its extremity, alternately in possession of the
French and the English—the gun being taken and retaken not fewer
than nine times between the hours of three and seven in the
morning.
Nor was the action less sanguinary in other parts of the field. Along
the sides of the various glens, in the hollow ways, through the
trenches, and over the barricades, a deadly strife went on. At one
moment the enemy appeared to carry everything before them; at
another, they were checked, broken, and dispersed; but the
darkness was so great that confusion everywhere prevailed: nor
could it be ascertained, with any degree of accuracy, how matters
would terminate.
At last day began to dawn, and a scene was presented of great
disorder and horrible carnage. Not only were the various regiments
of each brigade separated, but the regiments themselves were split
up into little parties, each of which was warmly and closely engaged
with a similar party of the enemy. In almost every direction, too, our
men were gaining ground. The French had gradually retrograded; till
now they maintained a broken and irregular line through the
churchyard, and along the ridge of a hill which formed a sort of
natural crest to the glacis. One battalion of Guards, which had
retained its order, perceiving this, made ready to complete the
defeat. They pushed forward in fine array with the bayonet; and
dreadful was the slaughter which took place ere the confused mass
of fugitives were sheltered within their own gates. In like manner a
dash was made against those who still maintained themselves
behind the churchyard wall; and they, too, with difficulty escaped
into the redoubt.
A battle such as that which I have just described is always attended
by a greater proportionate slaughter on both sides than one more
regularly entered into and more scientifically fought. On our part,
nine hundred men had fallen; on the part of the enemy, upwards of
a thousand: and the arena within which they fell was so narrow that
even a veteran would have guessed the number of dead at
something greatly beyond this. The street of St Esprit, in particular,
was covered with killed and wounded; and round the six-pounder
they lay in heaps. A French artilleryman had fallen across it, with a
fuze in his hand; there he lay, his head cloven asunder, and the
remains of the handle of the fuze in his grasp. The muzzle and
breech of the gun were smeared with blood and brains; and beside
them were several soldiers of both nations, whose heads had
evidently been dashed to pieces by the butts of muskets. Arms of all
sorts, broken and entire, were strewed about. Among the number of
killed on our side was General Hay; he was shot through one of the
loopholes in the interior of the church. The wounded, too, were far
more than ordinarily numerous: in a word, it was one of the most
hard fought and unsatisfactory affairs that had occurred since the
commencement of the war. Brave men fell when their fall was no
longer of use to their country; and much blood was wantonly shed
during a period of national peace.
A truce being concluded between General Colville, who succeeded to
the command of the besieging army, and the Governor of Bayonne,
the whole of the 15th was spent in burying the dead. Holes were
dug for them in various places; and they were thrown in not without
sorrow, but with very little ceremony. In collecting them together
various living men were found, sadly mangled, and hardly
distinguishable from their slaughtered comrades. These were of
course removed to the hospitals, where every care was taken of
them; but not a few perished from loss of blood ere assistance
arrived. It was remarked, likewise, by the medical attendants, that a
greater proportion of incurable wounds were inflicted this night than
they remembered to have seen. Many had received bayonet-thrusts
in vital parts: one man, I recollect, whose eyes were both torn from
the sockets and hung over his cheeks; whilst several were cut in two
by round-shot, which had passed through, yet left them breathing.
The hospitals, accordingly, presented sad spectacles; and the shrieks
and groans of the inmates acted with no more cheering effect upon
the sense of hearing than their disfigured countenances and
mangled forms acted upon the sense of sight.
It is unnecessary to remind the reader that, whilst our column of the
army was thus engaged before Bayonne, Lord Wellington, following
up his success at Orthes, had gained the splendid victory of
Toulouse. As an immediate consequence upon that event, the
important city of Bourdeaux was taken possession of by Lord
Dalhousie, and declared for Louis XVIII.; and farther conquests were
prevented only by the arrival of Colonels Cook and St Simon—the
one at the headquarters of Lord Wellington, the other at those of
Marshal Soult. By them, official information was conveyed of the
great change which had occurred in the French capital. An armistice
between the two generals immediately followed; and such an order
being conveyed to General Thouvenot, as he considered himself
bound to obey, a similar treaty was entered into on our side of the
theatre of war. By the terms of that treaty all hostilities were to
cease. The two armies were still, indeed, kept apart; nor was any
one from our camp allowed to enter Bayonne without a written pass
from the adjutant-general. Foraging parties only were permitted to
come forth from the place at stated periods, to which limits were
assigned, beyond which they were prevented from penetrating. Yet
the truce was regarded by both parties as an armed one. After so
recent an instance of treachery, we felt no disposition to trust to the
word or honour of the French governor; and the enemy guessing,
perhaps, what our feelings were, did not pretend to trust us. On
each side, therefore, a system of perfect watchfulness continued. We
established our pickets and planted our sentinels with the same
caution and strictness as before; nor was any other difference
distinguishable between the nature of those duties now and what it
had been a week ago, except that the enemy suffered us to show
ourselves without firing upon us. So passed several days, till, on the
20th, the war was formally declared to be at an end.
 CONCLUSION.
Little now remains for me to add. My tale of war and its attendant
dangers and enjoyments in this part of the world is told; and I have
nothing left to notice except a few of the most prominent of the
adventures which befell between the period of my quitting one
scene of hostile operations and my arrival at another. These are soon
narrated.
Early on the morning of the 28th of April 1814, the whole of the
Allied troops encamped about Bayonne drew up, in various lines, to
witness the hoisting of the white flag upon the ramparts of that city.
The standards of Britain, Spain, Portugal, and of the Bourbons,
already waved together from the summit of every eminence in our
camp. Up to this date, however, the tricolor still kept its place upon
the flagstaff of the citadel; to-day it was to be torn down, and the
"drapeau-blanc" substituted in its room. To us, no doubt, the
spectacle promised to be one of triumph and rejoicing; for we
thought of the gigantic efforts of our country, which alone, of all the
nations in Europe, had uniformly refused to acknowledge the
sovereignty of the usurper; but by the French it was very differently
regarded. Even among the country people not a spark of enthusiasm
could be traced; whilst by the garrison no secret was made of their
abhorrence of the new state of things, and their undiminished
attachment to their former master. But there was no help for it. "La
fortune de la guerre," said a French officer to me one day, as we
talked of these matters; but he shrugged his shoulders as he spoke,
and gave no proof that he was satisfied with its results.
We had stood in our ranks about an hour, dressed in our best attire,
and having our muskets loaded with powder only, when a signal-gun
was fired from one of the batteries of the town, and a magnificent
tricolored flag, which had hitherto waved proudly in the breeze, was
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