How to interpret the chart
The chart depicts a recently developed phylogeny of all living ferns.
The disks represent families of ferns. Groups of related families are indicated by a hierarchy
of regions and some of these groups are named.
For more than 250 years, families of plants and relationships between families were
determined mainly by morphology (the structure, shape and number of plant parts). Since the
1990s, genetic information has become available and now the affinity of species, genera and families
is determined primarily by this molecular evidence. The previous morphological classifications
were, to some extent, supported by the molecular analysis. However, major changes have been
necessary both to the circumscription of some families and to the placement of families relative to each
other and to the corresponding higher groupings.
The chart is based on the recent work aimed at resolving the molecular
phylogeny of ferns (see the bibliography below).
The chart depicts a phylogenetic tree in terms of a hierarchical grouping of families.
Many of these placements are now well-established, but uncertainties remain - these
we have tried to indicate on the chart
(see below for details of how uncertain placements are depicted). The chart will be updated
as new results become available.
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Phylogenetic trees
The chart corresponds to a phylogenetic tree. Regions on the chart,
delimited by boundaries or shaded areas, correspond to nodes in a
tree, and families, as disks on the chart, are leaves of the tree. In trees,
proximity of two families is represented by the depth of the root of the
smallest subtree containing the two families. Here it is represented by
a shared region. Proximity does not cross boundaries: families close
on the chart but separated by a boundary are no closer than their
common region indicates. Boundaries are depicted with varying styles
and colours - this is merely to aid the eye and has no phylogenetic
significance.
Uncertainties
Many of the placements of families are now well-established
and much work has been done to resolve remaining issues.
However, uncertainties remain. The chart is not intended
to be a resolution of these. Where evidence is poor, we have either (a)
left relationships unresolved (as a `polytomy'), or (b) chosen the best
supported on current evidence, or (c) indicated alternative placements.
On the whole, we have been conservative in the depiction of family relationships.
As new evidence appears, new versions of the chart will be published.
Nomenclature
Common names: Common, or vernacular, names (in English) are used to describe some familiar
ferns. These names vary from language to language. This is
particularly a problem when only local names are available. In addition, the
same common name may be used for several (sometimes unrelated) species.
Occasionally, these ambiguous names have been used but accompanied by
the relevant genus. In all, common names on the chart should be treated with caution.
Ages
Some extant fern families appear to originate as far back as the Carboniferous period. For example,
there are Carboniferous fossil ferns with marattioid (Marattiaceae) characters.
However, there have been several more recent periods of fern
development giving rise to other extant families, with some families in the
Polypodiales, for example, appearing in the late Mesozoic. Of course, the
vast majority of fern species are extinct, and with them are lost many aspects of fern diversity.
One of the curiosities of
the fossil record of ferns is that some extant species seem to have remained
unchanged for very long periods. For example, there are fossils from ~75Mya which
appear to be identical to Osmundastrum cinnamomeum (= Osmunda cinnamomea) and
fossils from ~57Mya which appear to be Onoclea sensibilis.
For recent summaries of fern paleontology
(and the relationship to phylogeny), see:
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Phylogeny and evolution of ferns: a paleontological perspective, G.W. Rothwell and R.A. Stockey, in Biology and Evolution of Ferns and Lycophytes. T.A. Ranker and C.H. Haufler (Eds.), C.U.P. 2008.
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Evidence for a Cenozoic radiation of ferns in an angiosperm-dominated canopy. E. Schuettpelz and K.M. Pryer. Proc. National Acad. Sci. USA. 106: 11200-11205. 2009.
Chart details
The chart is designed to be viewed at A3 size (29.7cm × 42.0cm, 11.75in × 16.5in) or larger.
If viewing in PDF, you may change the magnification in order to view and navigate the chart.
Personal note
This series of charts was prepared with the personal objective of creating attractive visual aids and, to some
extent, informative displays for my own use. Early versions were prepared before the advent of
molecular systematics and used morphological classifications and a display inspired by that of Stebbins (Flowering Plants: Evolution above the Species Level, G.L. Stebbins. The Belknap Press, Harvard University Press, Cambridge, Mass. 1974) extended to the family level. With the development of molecular phylogenies, considerable redesign was required,
including now much 'fine-structure' in the relationship of families.
I have decided to make the current versions generally available. I hope that these are found
as useful and as attractive as I intended. The charts will be updated as new results become available.
Printed versions of
this fern chart (A3 size) are available. For a chart of flowering plant families, visit the Angiosperm page.
David Rydeheard (2011)
Source papers
Recent resources for fern phylogeny include:
- Phylogeny and evolution of ferns (Monilophytes) with a focus on the early Leptosporangiate divergences, K.M. Pryer, E. Schuettpelz, P.G. Wolf, H. Schneider, A.R. Smith and R. Cranfill. American Journal of Botany 91(10): 1582-1598. 2004.
- A classification for extant ferns, A.R. Smith, K.M. Pryer, E. Schuettpelz, P. Korall, H. Schneider and P.G. Wolf. Taxon 55(3) 705-731. 2006.
- Ferns (Monilophyta), K.M. Pryer and E. Schuettpelz, in The Timetree of Life, S.B. Hedges and S. Kumar, Eds. O.U.P. 2009.
- Is Morphology Really at Odds with Molecules in Estimating Fern Phylogeny?, H. Schneider, A.R. Smith and K.M. Pryer. Systematic Botany 34(3):455-475. 2009.
- Utility of a large, multigene plastid data set in inferring higher-order relationships in ferns and relatives (Monilophytes), H.S. Rai and S.W. Graham. American Journal of Botany 97(9) 1444-1456. 2010.
- A linear sequence of extant families and genera of lycophytes and ferns,
M.J.M. Christenhusz, Xian-Chun Zhang and H. Schneider. Phytotaxa 19:7-54. 2011
- First insights into fern matK phylogeny, Li-Yaung Kuo, Fay-Wei Li, Wen-Liang Chiou and Chun-Neng Wang. Molecular Phylogenetics and Evolution 59: 556-566. 2011.
A useful dictionary of families and genera is:
- Mabberley's Plant-book: A Portable Dictionary of Plants, their Classifications, and Uses,
3rd Edition, David J. Mabberley. Cambridge Univ. Press, 2008.
A large bibliography (concentrating on flowering plants) can be found on the Angiosperm Phylogeny Website.
Related websites
- A valuable resource, with considerable detail and discussion, as well as
incorporating the latest results, is the Angiosperm Phylogeny Website: A website detailing molecular phylogeny of the
flowering plants, constantly updated and including discussions on placements,
detailed phylogenies, characters of groups, common names, etc. This
has been extended to include ferns and other non-flowering vascular plants.
Developed and maintained by Peter Stevens (Missouri Botanical Gardens and the University of Missouri) and Hilary Davis.
Contacts
The chart and website were developed by David Rydeheard. To contact me with comments, corrections or any other matters concerning this site, please email me
at BotanicalChart@gmail.com.
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