Journal of Systematics
JSE and Evolution
Editorial
Systematics and evolution of lycophytes and ferns
Welcome to this special issue of the Journal of Systematics and
Evolution focused on the systematics and evolution of
lycophytes and ferns. It is our hope that it will inspire
innovative research on these plants at this critical time where
human activities are transforming biodiversity as we know it.
Lycophytes and ferns are distinct evolutionary lineages,
with ferns strongly supported as sister to seed plants
and lycophytes, in turn, sister to this large (euphyllophyte)
clade. Notably, however, lycophytes and ferns do share many
features, including vascular plant apomorphies (e.g., vascular
tissue) and land plant plesiomorphies (e.g., spore dispersal
and the absence of seeds). For these reasons, they have long
been collectively treated as “pteridophytes” and it still seems
appropriate to address the biology of both lineages together.
This special issue was initiated in the context of the most
recent international scientific meeting dedicated to ferns and
lycophytes, Next Generation Pteridology, which was organized
by Eric Schuettpelz and colleagues in 2015. Although it is not
designed as the proceedings of that meeting, the theme of
“next generation” is maintained by including a diverse array
of early-career contributors (along with those who are
well-established) and by focusing on a selection of topics
for which data are coming from novel sources enabling
innovative research. These topics may challenge long held
hypotheses or support emerging new hypotheses.
Of course, this special issue cannot cover all aspects of
lycophyte and fern biology, and recent more-comprehensive
compilations already exist (e.g., Ranker & Haufler, 2008;
Mehltreter et al., 2010). Here, we aim to give attention to
contributions that will hopefully be relevant to the research of
the future generations of pteridologists. Topics were selected
to cover a wide range of issues, including some that may, at
first blush, appear a bit old fashioned. At the same time, some
readers may be surprised by the absence of contributions in
some frequently highlighted fields, but some outstanding
reviews have already been published in areas such lycophyte
and fern genomics (e.g., Sessa & Der, 2016).
All contributions to this special issue share a common theme
of aiming to document and understand lycophyte and fern
diversity. Such documentation arguably requires the ability to
classify these plants and we are pleased to publish here the first
installation of a new approach to the classification of
lycophytes and ferns from the Pteridophyte Phylogeny Group
(PPG I, 2016). The initiative towards the establishment of the
Pteridophyte Phylogeny Group was inspired, but not con-
strained, by the successes of the Angiosperm Phylogeny Group.
It involved an open call to anyone working on the systematics
of lycophytes and ferns, with the aim to establish a community-
based classification. The result required considerable compro-
mise and is quite distinct from existing classifications.
Diversity is also the focus of a critical review of our current
understanding of fern and lycophyte systematics in the
November 2016 | Volume 54 | Issue 6 | 561–562
doi: 10.1111/jse.12231
Neotropics (Almeida & Salino, 2016). Although this region
has been intensively studied by many prominent pteridologists,
including John Mickel, Robbin Moran, Alan Smith, and Rolla
Tryon, major gaps in our knowledge remain (e.g., under-
sampled regions and lineages requiring taxonomic revisions).
This review also highlights the need to study these ferns within
a broader framework (i.e., beyond national floras). A similar
message is given in another contribution, which explores the
distinct pteridophyte flora of the East African Biodiversity
Hotspot (Liu et al., 2016). This latter study was enabled by the
outstanding contributions of many pteridologists who ex-
plored Africa, culminating in the work by Jacobous “Koos”
Roux (2009), and further emphasizes the need to identify and
address gaps in our knowledge concerning the distribution and
taxonomy of pteridophytes, in both species-rich and species-
poor regions. Interestingly, these two contributions to this
special issue also serve to collectively highlight an interesting
shift in the study of tropical floras away from the historical
powerhouses of botany based in North America and Europe.
Links between evolutionary and ecological processes are at
the core of the research on elevational diversity patterns
summarized by Kessler et al. (2016). This study provides an
outstanding example of the power that can be achieved by
integrating observations of local diversity gradients into a
global hypothesis. A somewhat more local perspective is
provided by focusing on the pteridophyte diversity of Hawai’i
(Ranker, 2016). Oceanic islands are known to have exception-
ally high proportions of pteridophyte diversity (Kreft et al.,
2010), but are also widely recognized as “evolutionary
laboratories” in which to explore the accumulation of
biodiversity. The Hawai’ian islands are unique in combining
geographic isolation with extensive habitats that are suitable
for ferns and lycophytes. Future studies may attempt to
compare this oceanic archipelago with other islands that are
rich in pteridophyte diversity, such as the Mascarene Islands
(Hennequin et al., 2014) and to employ pteridophyte diversity as
an indicator for the impact of climate change on the biodiversity
on oceanic islands with high-elevations (Pouteau et al., 2016).
Two contributors to this special issue were willing to
assemble perspectives on topics closely associated with a pair
of the most influential pteridologists of the mid-20th-century:
Irene Manton and Edward Klekowski. The two reviews
explore state-of-the-art of research on hybridization (Sigel,
2016) and apomictic reproduction (Grusz, 2016). Both reviews
comprehensively summarize our current understanding and
point out promising research directions that take advantage
of new genomic tools (the access to genomic data has
historically been one of the main limitations of research on
these topics). Progress in our understanding of these aspects
of pteridophyte biology has always been closely linked to the
establishment of new methods, as well as the careful
application of groundbreaking concepts (Haufler, 2014).
© 2016 Institute of Botany, Chinese Academy of Sciences
562 Editorial
The combination of old and new approaches will likely also
lead to a renaissance in our understanding of the interactions of
lycophytes and ferns with other organisms. Pressel et al. (2016)
provide a critical look at pteridophyte-fungal associations that
is inspired by the authors’ research on these associations across
all seed-free land plants. Fungal associations, including
mycorrhizal associations, have been studied for a long time,
but, as the authors carefully document, many key aspects are
still in need of clarification. This contribution will hopefully
inspire similar critical investigations into other associations with
pteridophytes, such as that of phytophagous insects utilizing
ferns as their core food supply (Schneider, 2016). Much like
fern-fungus associations, fern-insect associations have been
long-studied but careful exploration of existing evidence,
combined with new evidence, will certainly lead to additional
insights (Schneider, 2016).
A final contribution to this issue is provided by Brummitt et al.
(2016), who provide an important look to the current threats to
pteridophyte diversity, as assessed using IUCN Red List criteria.
This is the first comprehensive analysis of available data for
these plants, and covers about 8% of extant pteridophyte
diversity. The results of this study may be interpreted as
evidence supporting the hypothesis that pteridophyte diversity
will experience a major decline in coming decades. In this
context, the subsequent analyses defining spatial aspects of
the threats to pteridophyte diversity will hopefully encourage
pteridologists to examine changes in diversity focusing on both
global threats (as in Brummitt et al., 2016) and local changes
(Pouteau et al., 2016). Progress may not only require further
inventories of fern and lycophyte diversity (see Almeida &
Salino, 2016; Liu et al., 2016) but also the integration of different
types of evidence to obtain reliable predictions based on
understanding ecological-evolutionary processes (Kessler et al.
2016), the impact of interactions (Pressel et al., 2016), and the
role of reproductive biology (Bystriakova et al., 2014). We hope
that this special issue will inspire pteridologists to consider their
contributions to ongoing efforts to protect the diversity of
these plants within established frameworks to conserve plant
diversity (Corlett, 2016).
In closing, we would like to express our gratitude to the
Editors-in-Chief, Song Ge and Jun Wen, of the Journal of
Systematics and Evolution for providing us with the opportu-
nity to compile this special issue and we also want to express
our thanks for all the patience and support provided by the
editorial team, especially by Yan Liang. And, of course, we
want to thank all of the contributing authors, whose inspiring
efforts enabled us to assemble this special issue that
demonstrates the diversity of expertise in a truly global
community carrying out research on ferns and lycophytes.
Harald Schneider
Institute of Ecology and Evolution, School of Life Sciences,
Sun Yat-Sen University & Department of Life Sciences,
Natural History Museum, London
Eric Schuettpelz
Department of Botany, National Museum of Natural History,
Smithsonian Institution
J. Syst. Evol. 54 (6): 561–562, 2016
References
Almeida TE, Salino A. 2016. State of the art and perspectives on
neotropical fern and lycophyte systematics. Journal of Systemat-
ics and Evolution 54: 679–690.
Brummitt N, Aletrari E, Syfert MM, Mulligan M. 2016. Where are
threatened ferns found? Global conservation priorities for
pteridophytes. Journal of Systematics and Evolution 54: 604–616.
Bystriakova N, Ansell SW, Russell SJ, Grundmann M, Vogel JC,
Schneider H. 2014. Present, past and future of the European rock
fern Asplenium fontanum: combining distribution modelling
and population genetics to study the effect of climate change
on geographic range and genetic diversity. Annals of Botany
113: 453–465.
Corlett RT. 2016. Plant diversity in a changing world: status, tends, and
conservation needs. Plant Diversity 38: 10–16.
Grusz AL. 2016. A current perspective on apomixis in ferns. Journal of
Systematics and Evolution 54: 656–665.
Haufler CH. 2014. Ever since Klekowski: testing a set of radical
hypotheses revives the genetics of ferns and lycophytes.
American Journal of Botany 101: 2036–2042.
Hennequin S, Kessler M, Lindsay S, Schneider H. 2014. Evolutionary
patterns in the assembly of fern diversity on oceanic Mascarene
Islands. Journal of Biogeography 41: 1651–1663.
Kessler M, Karger DN, Kluge J. 2016. Elevational diversity patterns as
an example for evolutionary and ecological dynamics in ferns and
lycophytes. Journal of Systematics and Evolution 54: 617–625.
Kreft H, Jetz W, Mutke J, Barthlott W. 2010. Contrasting environmen-
tal and regional effects on global pteridophyte and seed plnat
diversity. Ecography 33: 408–419.
Liu HM, Zhang SZ, Wan T, Kamau PW, Wang ZW, Grall A, Hemp A,
Schneider H. 2016. Exploring the pteridophyte flora of the Eastern
Afromontane biodiversity hotspot. Journal of Systematics and
Evolution 54: 691–705.
Mehltreter K, Walker LR, Sharpe JM. 2010. Fern ecology. Cambridge:
Cambridge University Press.
PPG I. 2016. A community-derived classification for extant lycophytes
and ferns. Journal of Systematics and Evolution 54: 563–603.
Pouteau R, Meyer J-Y, Blanachard P, Nitta JH, Terorotua M, Taputuarai
R. 2016. Fern species richness and abundance are indicators of
climate change on high-elevation island: evidence from an
elevatinal gradient on Tahiti (French Polynesia). Climatic Change
138: 143–156.
Pressel S, Bidartondo MI, Field KJ, Rimington WR, Duckett JG. 2016.
Pteridophyte fungal associations: Current knowledge and future
perspectives. Journal of Systematics and Evolution 54: 666–678.
Ranker TA. 2016. What do we know about Hawaiian ferns and
lycophytes? Journal of Systematics and Evolution 54: 626–637.
Ranker TA, Haufler CH. 2008. Biology and evolution of ferns and
lycophytes. Cambridge: Cambridge University Press.
Roux JP. 2009. Synopsis of the Lycopodophyta and Pteridophyte of
Africa, Madagascar, and neighbouring islands. Strelitzia 23: 1–296.
Schneider H. 2016. The ghost of the Cretaceous terrestrial revolution
in the evolution of fern-sawfly associations. Journal of Systematics
and Evolution 54: 93–103.
Sessa EB, Der JP. 2016. Chapter Seven—Evolutionary genomics of
ferns and lycophytes. Advances in Botanical Research 78: 215–254.
Sigel EM. 2016. Genetic and genomic aspects of hybridization in ferns.
Journal of Systematics and Evolution 54: 638–655.
www.jse.ac.cn