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An introduction to fossil plants

Article  in  Journal of Biological Education · December 2010

DOI: 10.1080/00219266.1976.9654085

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 Repinted from Jounal o{ BiotogicatEducation(1976) 1O (4) 173-.180
An introduction to fossil plants
B. A. THOMAS
I D epartment of B iologicaI Scien ces,
t University ol London Goldsmiths' College, New Cross,
London SE|4 6NW
{
Abstract
This is an introduction to methods of studying fossil
plants and the teaching of palaeobotany. Brief
accounts ale given of different types of preservation
and where to find specimensfor one's own use. An
annotated bibliogaphy is provided to allow tho
readerto pursuethe subjectfurther,
1. Introduction
The study of fossil plants is a subject which is
often completely ignored in school biology
curricula,even though pupils are usually taught
som€thing about palaeontology and animal
evolution. Indeed, on thinking back. I have
memoriesoflearning detailsofthe geat reptilian
dinosau$ and of tlte Ice Age mammoths,but of
their associated plantstheie wasnevera mention.
This situation appearsto remain, with the same
dispadty often persistinginto higher education.
and it will probably become even more pro-
nounced with the reduction in morphology
courseswhich is occurring in many institutions.
However, if botany is to persist as a subject,
some morphologicattraining is still necessary,
and I maintain that some knowledgeof palaeo-
botanycan make plant morphologya little more
stimulating and hopefully a little easier to
understand,
Studiedin isolation,many modem plants can
appearbizare and possibly irrelevant in terms
of generalplant morphology.Such plants must,
however,havesomeinter-relationships with other
more familiar species,and suitablecomparisons
'l should put them irlto better penpectivefor the
student,Any knowledgeof extinct forms. show-
ing ancestral or intermediate characters,can
; naturallyhelpsuchinterpretations.Palaeobotany
should thereforebe thought of as gi\ing an
insight into the history of plants and vegetation,
rather than constituting an abstract lesson in
plant taxonomyor moryhology.
Fossilscan be studiedin a numberof waysfor
various purposes.Anatomical investigationscan
be madeon individual specimens, which can lead
to discussionson taxonomic relationshipswith
other plants and to hypothesesabout the cou$e
of evolution. The identification of groups of
fossil plants can give an id€a of the past vege-
tational cover, and of how drastically plant
ecosystems have changedover the millennia.ThQ
very formation of fossilsis an important process
in itself which can be linked to the wider aspects
of geologyor to the more specialisedprocessof
rock-dating.
This may seemall very well in theory,but how
is fossil botany to be taught?What are the ques-
tions to be asked, and indeed, wher€ are the
facts to be obtained?The main fields of interest
seemto revolvearound preseNation,methodsof
study and interpretation,identification,and the
whereaboutsof specimens.Thesekey aspectsof
fossilplantsare discussed in outline here.and.lor
thosereaderswho uish to enlargeon rhcm,an
annotated biblio$aphy is provided at the end
of this paper.
2. Preservation
The one questionthat is alwaysasked,is 'How
was it made?' and, if this can be answered,the
teacher'sprime difficulty is often over. Thereare
naturally many types of fossils,for dead organ-
isms may be preservedin a variety of ways.
Certain plants, chiefly algae, have exoskeletons
of silica or calciumcarbonatewhich resistdecay
and can be preseNedwith no further change-
rather like animal bonesor shells.However,the
vast majo ty of plant remains have become
altered in some way during fossilisation; the
essentialf€ature in all casesis that total decay
has been prevented.Here the discussioncan be
linked to the related topics of the carbon and
174
nitrogen cycles. If the students are not alr€ady
familiar with these cycles,then this could be a
useful introduction to the subject for enlarge-
ment at someIater date. Such a study will show
that mpid entombment is usually necessaryto
prevent aerobic decay; it is the type and speed
of this entombment that oontrols the ultimate
form of the fossil.
When plant remains are submergedin mineral-
rich water, they can be preservedby infiltration
and inorganic replacement in a similar manner
to the way in which inanimateobjectsare 'tumed
to stone'by tbe water from petrifyingsprings.
Such fossils are naturally called petrifactions and
can be studied anatomically like living plant
matedal (e.g. Fig. 2e) altholugh, of course,
diflarent sectioningtechniqueshav€ to be used.
Cell walls can often be recognisedwith little
dimculty and, very occasionally, preseryation is
so fine that some sub-cellular structures such as
nuclei or starch graius can be identified.
Not all fossil plants are so pedectly preseryed
as petdfactions. Many more plant remains
becomedepositedalong with normal sediments
in watery anvironmentssuch as lakes,estuaries,
and deltas.Theseplant remainsundergopadial
decomposition and gradually become flattened
by the weight of the accumulating sediments.In
time these deposits may be converted into clays,
shales, or sandstones still retaining their en-
tombed, and now fossilised,plants. Here the
fossils will lie concealeduntil the rock is split
open, unlessthey are fust destroyedby erosion,
oxidation by water percolation,or by someother
geological activity. These fossils are called.com-
pressions arrd, as their nams implies, consist of
compressed plant material coDtainingvarying
amounts of the original organic matter (Figs Ia
and d). Sometimes,however,the decomposition
of the matter may be complet€ or nearly so,
leaving only an outline of the plart as marks on
the rock matrix. These are called.impressions
(Figs 2c and,d).'then again, large rigid structures
such as stems or roots may decay before com-
pactionofthe surroundingsedimentshasfinished.
The cavity which results can become filled with
mud or sand which later hardens to folm a cast.
Occasionally the internal decay may be
incomplete and perhapslimited to the central
medulla or pith, thus producing a pith cast. If
compaction of the surrounding sedimentsoccurs
beforethe entombedtissues decayakind ofthree-
dimensional impression is formed called a
mould. ^lherc
is often some confusion about the
terrns cast and mould but it is relatively easy to
Journal
of Biological (1976)10 (4)
Education
rememberthat a cast is a replica whilst a mould
is a negativeof the original,
Under certain conditions,decompositioncan
be very limited, allowing more of the original
plant material to r€main, and under these con-
ditions vast amounts of organic matter can
accumulate.This is how the coal seamswere
formed, as they are really compressedpeats or
swamp deposits. Conversely, if coal seams are
massivecompressions, singleplant compressions
tt
can be thought of asminiaturecoal seams.
Whatever the state of preseryation, ooe must t
alwaysremember that it is not detelmiqed by the
ageofthe rocks.Oncefossilised,organic remains
stay unchanged unless further aflected by later
geological phenomena.Thus a 300 million-yeax-
old fossil may be better preservedthan one less
than a million yearsold. The teacheris, therefore,
well advised to have available several similar
specimensof widely different ages, with prefer-
ably evena living examplefor funher comparison.
The student should be made awaxe of the
typesoffossilisationprocesses that canoccur,but
should not think that this is somethingwhich
happenedonly in the dim and distart past.Plants
are being fossilisedtoday in a yariety of habitats,
just as they alwayshaye been,Shells on the beach
are ready examplesof durable remains,eren
though they are animals; leavesand twigs being
washed by rivers into lakes may eventually form
compressionsor impressions,while the many
swamps and active peat bogs clearly illustrate
large-scale accumulation of organic matter. It
should become apparent that plarts ftom some
habitats are more likely to becomefossilisedthan
plants from others, and that some axeasof the
world are more favourable for fossilisation pro-
cesses.The plants being fossilisedtoday will
certainly not be a complete record of the present-
day vegetationofthe wholeworld. soit shouldbe
appreciated that fossils can never be expectedto
tell us everything about the many t'?es of plants
which haveexistedin the past.
One other feature of plant fossilisation which
should also be clarified is the usual dismember-
ment of plants prior to their preservation.
Fragmentation nearly always occurs, so it is the
isolatedplant organswhichlie entombedawaiting I
discovery. We therefore find leaves,stems,roots, (
seeds,etc., but only rarely do we find whole
plants, This should explain even further why our Ii
knowledgeis often so incomplete.The person
studying living plants ca.nusually obtain maoy
specimens,ilustrating all stages of their life
history, but the comparable worker dealing with
 i Fossilplants
Thomas
fossils may have only one or a very few badly
prese ed fragments. The fossil botanist tries
hard to interpret his specimensbut is more often
than not hampered by lack of suitable material.
3, Methods of study
(a) Petrifactiotrs
t Tho type of prese ation naturally detcrmin€sthe
way in which fossils can be studied, P€trifactions
are often consideredto be the best type for study,
becausethey show internal anatomy. Originally,
such specimens were treated like rocks or
minerals, being cut, ground, and mounted in a
similar manner. More recently though, other
techniques have been evolved which are easier
and sometimesgive better results.They rely on
carbonaceous matedal remaining in the cell
walls, a feature which was not so important
whenusingthe older sectioningmethods.Among
the most well-known petrifactions are the 'coal
balls' which occur in somecoal seams.Theseare
limestonenodules containing plant remains which
wereprobably formed by mineral depositiondudng
the rarg marine incursions over the Carboniferous
swamps.Such petrifactions should be fiIst cut,
smoothed with carbotundum powder, and then
etched with dilute (about 2%) hydrochloric acid.
No specific time can b€ given for this etching, as
the reaction time naturally dependson both the
mineral content of the coal ball and the concen-
tration of the acid. Personal expedmentation is
clearly needed at this point. The etched surface
then needsto be washed and speedily dried with
a warm air drier. Acetone should then be flooded
over the surface, and a thin film of cellulose
acetate sheet (0.05mm) laid gently aqoss it.
When the solvent has evaporated, the acetate
shect can be gently peeled ofi taking with it the
now embedded cell walls. Warm air blowing
gently over the sudace of the acetate sheet will
again speed up this part of the procedure. The
'peel' can be cleanedwith more dilute acid, to
remove any adhering mineral matter, before
being finally washed, pressed&y, trimmed, ard
mounted (Frg. 2e).
This is a straightforward technique which gives
good results once the etching time has been
mastercdand providing that the etched surfaceis
never touched. Any pressure on the suface will
destroythe exposed,upstandingwalls. The next
'peel' car be madeby simply smoothingthe coal
ball surface and going through the etching and
peeling procedues once again. Compared with
the older technique this is simpler arld quicker,
175
gives a much thinner preparation, and there is
virtually no wastage of material due to cutting
and grinding. This meansthat even small delicate
organs, such as seeds aud pollen sacs, can be
serially 'sectioned'enabling detail€d studies to
be made. It is a techniquewhich can also be
used on oth€r tlpes of petdfactions. There are
examples of limestone preseryation other than
coal ba.lls and there are also many examples of
plants preseryedin silica or py,rite. Silica is
attacked by hydrofluoric acid so the procedure is
virtually identical except for the far greater
safety precautions which are essantial,This acid
is of course extremely dangerous and needs to
be kept in polythene containers, In fact, the
safety requirements will prevent such prepara-
tions ftom being attempted in the general
laboratory. Pyrite specimens,however, cannot be
etched, so sectioning is the procedwe usually
undertaken for their study. Much work has been
done recently on rather fragile plritised speci-
mens; this has necessitatedthe deyelopment of
specialisedtechdques such as embedding speci-
mens in plastic before sectioning. Thesemethods
are clearlyoutsidethe scopeof the non-specialist;
so they ffe not consideredhere. Readerswishing
to persuc this subject are recommendedto read
the methods section in the paper by Grierson and
Banks 'Lycopods of the Devonian of New York
State' (Palaeo togrctphica Americana, lY, 31,
1963).
(b) Compressions
Cedain tlpes of compressions may alsobe studied
in the laboratory using straightforward prepara-
tofy methods. The technique which lends itself to
general laboratory classesis one allowing the
epidermisto be examinedin more detail than
can be seenby direct magnification. At first sight
this may seamrather trivial, but it is really far
from it. Epidermal characters are important
features used in classification,especiallywhen
one is dealingwith isolatedorgans.Indeed,mary
leavesand shootswould be indistinguishableif
it were Dor for their epidermisesshowing varia-
tions in the arrangement and type of stomata,
epidermal cells, and trichomes.
The method relies on the important fact that
cuticles,as opposedto other plant constituents,
may sometimesbe little affectedby fossilisation.
Oliginally the cuticle lay over and merged into
the anticlinal walls of the epidermalcells. Thus,
if isolated, it will reveal much of the odginal
structwe of the epidermis. The problem is
simply to remove the rest of the altered plant
176
tissues.Fragments of the compressionshould be
removed from the specimen and immersed in
Schultze'ssolution, a mixture of potassiumor
sodiumchloratein concentratednitricacid (about
0.3 mg in 10 cm8).The reagentoxidisesthe plant
tissues,eventuallyturning them light brown and
translucent in appearance. The time necessary
for complete oxidation may vary considerably
from less than an hour to several days, but as a
rule overnight imme$ion is normally suitable.
The fragments should then be carefully washedto
remove all tracas of the acid, and afterwards
immersedindilute amrnoniasolution.This alkali
dissolvesthe oxidised non-cuticular plant sub-
stancesleaving the cuticle to be finally washed
and mounted. Caxe must be exercised at all
stages,as the cuticle may be delicateand easily
broken or lost in the washings.If damageor loss
is persistently encountered the whole procedure
can be caried out on a microscopeslide with
the matedal safely held under a cover-slip. The
various solutions can easily be changed by care-
fully drawing them through with filter paper.
Reproductiye organs (Frg. Ia) can be studied
in a similar manner, often with spectacular
results, Apart from fragments of the cuticle,
evidenceis often obtained of the organ's spore
content (Fk. 1r). This is possibly because
maceration leavesthe outer spore coat relatively
unharmed as it is made of a very resistant
cutinJike substancecalled sporopollenin.Such
spores and pollen grains are genemlly very
varied in sizeand surfaceornamentation,show-
ing a great variety of granules.lumps. spines,
flanges, or even bladders. Normally, however,
plant speciesproduce their own individually
recognisabletypes of sporesso it has been pos-
sible for researchworkers to gradually build up
an overall picture of plant-spore relationships.
Such work can also demonstrateheterospory
when two markedly different spore types are
consistcntlyobtainedfrom the sameparentplant.
Clearly this oxidation proceduregiles us a
method of showing that compressionsare not
merely black smearson thg rock surface, but are
the actual remains of once-livingplant tissues.
There are naturally difrculties which should be
pointed out to the potential user. Not all com-
pressionscontain cuticle; the plants may never
have had any in life, or it may have been des-
troyed during compression or during subse-
quent geological activity. Reproductive organs
may be empty giving only cuticle flagments.
However, the function of such an organ is to
reproducaand to do so it must shedits spores.
Journal
of Biological ('1976)
Education 10 (4)
Remembering that coal seams are really
massivecompressions, one can extendthe useof
this techniqueto give many more usefulresults.
Coal when maceratedusually gives sporesand
fragments of cuticle, demonstratingthat coal
was formed by large-scaleaccumulationof plant
matedal. More important is the fact that the
huge vadety of sporesfound shows how vaded
was the vegetationwhich formed the coal, Many *
of these isolated sporescltn be identified with
their parentplantsasa resultof previouswork on
fructifications.By suchwork it hasbeenpossible
to build up a much more accuratepicture of the
vegetationalcovfl at the time of deposition.We
now know that the Carboniferousswampswere
not dominatedby a homogeneous vegetationbut
were covered by a changing series of plants,
ranging from forest in standing water to drier
moorland-like communities.Such an oxidative
macerationof coal is similar to the lessdrastic
procedureused for studying the much younger
Quatemary peat. This related subject has already
been dealt with by F. M. Slater in this
journal; his paper and other generaltcxts cover-
ing this topic are listed in the bibliography.
4. Where to f ind fossils
The paramount problem for anyone attempting
to teach palaeobotanyis usually the dificulty of
obtaining sufficient suitable material. Pictures,
or colour slides, are useful but are really no
substitutefor real specimens.Studentsmust be
able to seeand handle some fossilsbefore they
can develop any real insight into the subject.
Most museumshave someform of collection on
show, and thesecan be very instructivein illus-
trating the rangesof form and presentationand
the varietiesof vegetationwhich have existedin
the past, This can be even more valuable if the
museumhas collectionswhich can be borrowed
through a schools'loan service.However, for
long term usage,it is obviouslybestto haveone's
own specimens for laboratory classes.This
usuallymeansbuyingthem,or prefemblycollect-
ing them oneself.
Fossil plants can usually be bought ftom
reputablegeologicaldealen, although the avail-
ableselectionis nearly alwayslimited to the most
gasily obtainablecoal measurespecimens.Most
parts of the Bdtish Isles contain fossililerous
Iocks, so neaxly everyonehas accessto some
collectinglocalities.Therquestionis alwayswhere
to go. Quanies, cliffs, pits, road and river
 T h o m a s : F o s s i lp l a n t s

#'

Frg. 1. (a) A Lower Carboniferouslycopod conecompression(Lepiclostrobus

col osus),x2. (b) An isolatedspore
preparedfrom the samelycopod cone, )i 1000.(c) An isolated spore preparedby macerating Upper Carboni-
l-erouscoal, X300. (d) An Upper Carbonjferouslycopod shoot compression(Lepido.lendtullsp.), X2. (e) An
Upper Carboniferouslycopod rhizophorecast (Stigmaia sp.), )<A.6.
 Journalof Biological (1976)t0 (4)
Education

:ij

Frt. 2. (a) Cuticle preparedby nacerating a Jurassicpteridospermleaf compression(Pdchypterispapillosa), x3.

(b) An enlargedportion ofthe leafcuticleshowinga stomaand the outlinesof epidemal cells, X 300.(c) A Lower
Carboniferouslycopod stem impression(Eskdaliasp.), x 1. (d) An impressionof a Lower Carboniferousmoss
(Muscitessp.), t 4: ftofi Annalsof Botany,36 (1972),pl. 1, p. 160.(e) A celluloseacetatepeel lrom an Upper
Carboniferousgymnospermwood petrifaction (Codaites sp.), x45,
 Thomas: plants
Fossil
cuttings,and coal tips are all potential collecting
localities; indeed,any exposedsedimentaryrock
is possibly fossiliferous,although, as a general
rule, the bett€r fossil plants are confined to finer
glained strata. Local museums or natutal
history societiesmay be able to help and to offer
advice about local sites.Alternatively, one can
tuur to the literaturg for records. The Memoirs
-t! of the Geological Survey are the most useful
referencasourceand containd€tail€ddescriDtions
of si(es and lists of recordedfossils.Of similar
t value is the D ectory of FossiliferousLocalities
edited and publishedby the Palaeontogaphical
Society (1954). This is, however, somewhat
limited in content becauseit covem the whole
country in a single volume. Perhapsone word
of caution should be introduced here: because
many of these sites were fust described many
yeaxsago, there is some risk that a particular
locality may no longer be accessible, especiallyif
it is within or neax a large conurbation. Some
sites are destroyed by rubbish tips and new
buildings,while otherssimply seemto disappear
under encroachingvegetation.Thus it is really
essentialto make a preliminary visit to a placa
before taking a party there, to avoid any possible
disappointment or embalrassment. Permission
should also be sought beforehandfor any visits
onto private land, especiallyif the site is being
worked in any way.
Firally, I must stress the need for adequate
safety precautionsat all times when collecting,
as cliffs, old quard€s, and tips are potentially
dangerousplacesto visit. Many are really quite
unsuitable for parties, especially if there are
many younger members in them. However, I
should not like to make the readertoo despon-
dent before he even starts collecting. There are
hundreds of good collecting localities, and all
that is necessaxyto build up a collection is
perseyeranceand, sometimes,hard work.
In conclusion,may I make a plea for reason.
Recognise the point when you have enough
material for your needs or for youf mode of
transport. Always collect sensibly-sites can be
ruined by senselessrock smashing-and remem-
ber that others will come after you. There is a
'Code for geological fieldwork' issued
by the
Geologists' Association (c/o Geology Depart-
ment, UniversityCollege,Gower Street,London
{ WC1E 68T) which succinctly lists the points
which should be observedwhen collecting.In
particulax it stresses the points which I have
made, that is to take care and to avoid ruining
natural localities by over-collecting.
179
Annotated bibliography
Palaeobotanical lirerature is largely \aritien for
advanced botanical study or for research and is
therefore not suitable for direct classroom use.
Some books are useful as rcfercnce material. however.
so I havelisted the best of the English languagetexts.
Andrews, H. N. Ji (1961) Studies in paleobotany.
487 pp. New York : John Wiley.
A general introductory textbook, now a little out of
date in places.
Banks, H, P. (1910) Evolution and Plants of the past.
170pp. Belmont, California: Wadsworth.
A good itrtroductory book, although one concen-
trating mainly orl the ea.rliestevidence of plant life
on the earth, the grcen plant itrvasion of dry land,
and the development of the pine cone. It also con-
tains a useful glossary.
Chaloner, W. c. and ColliNon, M. E. (1975) An
illustrated key to the commotret British UDDer
Carboniferouspiant compressiotr| os,ils. Proiietl-
ings of the Geologists' Association, 86, 1,
1-44.
A very useful up-to-date key.
Crookall, R. (1929) Coal Measure Plants. BO DD.
Lotrdoo:FdwardArDold.
A basic identification book with descriDtions.
figlres. and photogaphs of the commoneir coai
measurefossil plants. Out of pint but occasionally
obtainable,
Delevoryas, T. (1962) Moryhology afid Evolatiotr of
Fossil Plahts. New York: Holt, Rinehart and
Winstotr.
A useful intrcductory book although a little dated
rn palts.
Harris, T. M. (1961-) I:he yorkshire Jurassic Flora,
London: British Museum (Natual Hislory).
Staxted in 1961, there will eventually be five parts.
These ale again very detailed, but expeosive,so
they are rccommended solely for rcference and plant
ideorificatioD.
Hughes, N. F. (197Q Palaeobiology of Angiosperm
Oriei s. U2 pp. Loodon: Camb dge University
Prcss.
An advanced, but rcadable account which stresses
palaeobotany rathet than comparative morphology
ofliving plants.
Lacey, W. S. (1963) Palaeobotanicaltechdques. In
yiewpoints in Biology, yol. 2, pp. 202-243. London:.
Butterwodh.
A very useful rcview article giving a1l necessary
practical details.
Pennington, W. (1969) The Ektor! of Btitish Vegeta-
tion. 152 pp. Londot English UniveNities Press.
An introductoly book linking Tertiary and Quatem-
axy palaeobotany with the present-dayB tish flora,
includiDgmaD'seffecrson rhe environmeDt.
Slater, F. M. (1972) Quatema-ry pollen analysis ir
secondary school ecology. Ioarxal of Biological
&lucatiak, 6, 109-117.
A concise introductory account suitable for
advatrced school work.
 180 Journslof BiologicalEducation(1976) 10 (4)

Sporne, K. R. (1971) Ihe Mysterioas Origin of
Floweihg Plants. Oxford Biology Readers No. 3,
16 pp. Londo[: Oxford Univ€rsity Press.
Strongly recornmended for this topic.
Walker, D. and West, R, G., Eds. (1970) .Strd,'4 r',
the yegetational History of the British ltles, 6pp.
Londoa: Cambridge University Press.
A sedes of specialist essays on the PleistoceDe
history of the British flora.
West, R. G. (1971)Stub,ttc' the Past by Pollen Analysis.
Oxford Biology Readers No. 10, 16 pp. Londolr:
Oxford University Press.
Strcngly recommetrded as an introductory text for
this tooic.

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