Nordic Society Oikos

Is 'Life-Form' a Useful Concept in Bryophyte Ecology?

Author(s): J. W. Bates
Source: Oikos, Vol. 82, Fasc. 2 (Jun., 1998), pp. 223-237
Published by: Wiley on behalf of Nordic Society Oikos
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 1998
OIKOS 82: 223-237.Copenhagen

N41NI- Minireviewsprovidesan opportunity

to summarize
ecologicalareas,withspecialemphasis
on current
existing
knowledgeof selected
topicswhererapidand significant
advancesare occurring.Reviewsshouldbe conciseand nottoo wide-ranging.
All key
AddVI Em TT references
shouldbe cited.A summary is required.

a usefulconceptin bryophyte
Is 'life-form' ecology?

J. W. Bates

ecology?- Oikos82:
a usefulconceptin bryophyte
Bates,J.W. 1998.Is 'life-form'
223-237.

Bryophytes are a successfulgroupof non-flowering landplantsthatmostlyavoid

competition withtracheophytes. Many bryophytes growon substratathat are
impenetrable to tracheophyte rootsand toleratefrequent desiccationand poor
conditions. Mostspeciesexhibit clonalor coloniallife-forms. Theserather thanthe
individualshootsorthalliaretheecologically functional units.Approximately eleven
distinct
life-forms can be recognized although theschemes ofno twoauthorsagree.
Bryophyte life-forms can be interpretedas recurring arrangements ofthephotosyn-
tissuesthatminimize
thetic evaporative waterlossandmaximize primary production.
However,otherfactorssuchas prevention of photoinhibition and scavenging of
cloudwater mayalso be important. Manyspeciesshowplasticity oflife-form accord-
ingto environmental conditions. Aerodynamically competent coloniesare probably
fashioned passively becauseprotruding individuals are moreproneto desiccation
thanthoseremaining within thelaminarboundary layer.Alterations ofdensity and
shootand leaforientation appearto be regulated byphytochrome-mediated growth
responsesto shading. Thedifferent life-forms can be arranged in sequences reflecting
wateravailability and lightintensity in different habitatsbut fewercategories are
represented on soilsthanon hardsubstrata becauseof negative interactions(stress
anddisturbance) withtracheophytes. Despitethestrong correlation oflife-forms with
moisture and lightconditions, thereare limitations to theiruse as a framework in
ecologicalstudiesofbryophytes. Different categories of life-form havebeenloosely
appliedand an improvedsystembased on objectivecriteriais required.Also,
life-forms
havenotevolvedindependently ofotherattributes oflife-strategy.
Colonial
life-forms
confer a competitiveadvantage inpre-empting spaceinthepatchy habitats
occupiedbybryophytes; moreover sometypes, like'dendroids', maybe specialists at
capturing space occupiedby otherspecies.Life-forms may sometimes also be
interpreted, especiallyin the morehighlyproductive specieslike Brachythecium
rutabulum, as adaptations forforaging. Internalmovement of nutrients fromfa-
vourably placedrametsmaybe beneficial to thecompetitive fitness of thewhole
colony.Life-forms and breeding systems in bryophytes areunlikely to haveevolved
independently as dioicoustaxatendto formunisexual colonieswithlowchancesof
fertilization.It is concludedthatlife-form is a usefulconceptbecauseof the
exceptionally highdependence of bryophytes on transient external watersupplies;
however, majoradvancesin understanding willdependupon investigation of the
relationships oflife-form to otherattributes ofbryophyte life-strategy.

J. W. Bates,Dept of Biology,ImperialCollegeat SilwoodPark, Ascot,Berkshire,UK

SL5 7PY (j.bates@ac.ic.uk).

Accepted19 January 1998

Copyright?) OIKOS 1998
ISSN 0030-1299
in Ireland- all rights
Printed reserved

OIKOS 82:2 (1998) 223

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 Despite theirlow staturesand limitedcompetitive capa- specifyhow this could be distinguishedfrom'growth
bilities,the bryophytesare a highlysuccessfulgroup form'in the case of bryophytes. Until we have a better
(15 000-20 000 speciesworld-wide)in comparisonwith understanding of therelationships betweengrowth-and
othertypesof non-flowering land plants. Their ability life-forms, it is desirableto have separatetermsforthe
to inhabit impermeablesubstratalike rock surfaces, two supposed levels of morphologicalorganizationof
treebark,leaf cuticlesand permanently frozenground, bryophytes(La Farge-England1996). The definitions
whichare not generallycolonized by vascular species, proposed by Mdgdefrau(1982) are relativelyuncon-
lies at the heartof thissuccess.Bryophytes are poikilo- tentious and unambiguous.Accordingly,life-formis
hydric,the dominantgametophytes mostlylack a wa- used as the appropriatesynonymforcolony organiza-
terproofcuticleand are unable to controltheirwater tion in the followingaccount and regardedas distinct
contentactively(Hosokawa et al. 1964). Whereasmany fromlife-strategy.
speciesare highlytolerantof desiccation,others,mainly Althoughseveralclassification schemesforbryophyte
restrictedto continuouslymoist habitats, are highly life-formshave been published, uncertaintiesexist
desiccation-sensitive (Proctor 1990). Bryophytesare about the relationshipsof these with environmental
also able to grow underverypoor conditions,particu- factors.All earlierwritershave recognisedthe crucial
larlyat low lightlevels,under and betweenthe domi- importanceof moisturesupply.The levelof exposureto
nant vascularplants (Grimeet al. 1990). solar radiationis also identified as an importantfactor
In view of theirextremedependenceon the unpre- in determiningoptimal life-forms(e.g. Birse 1958a).
dictablewatersupplyin precipitation, and the protec- However,a clearmodellinkingthevariouslife-forms to
tion that dense packing of shoots or thalli can give variationsin environmental factorshas not been forth-
against evaporative water loss (Proctor 1981), the coming.Birse(1958b) observedthat,"featuresof com-
colony ratherthan the individualis widelyregardedas munity structurereappear under recurringhabitat
the functionalunitformanybryophytes. Colonies may conditions,irrespectiveof the species present",thus
arise when many gametophoreinitials develop syn- implyinga close concordanceof life-forms with envi-
chronously,or theymay be synthesized by theprolifer- ronment.An inherentdifficulty is that conspicuous
ation of offsets of one or more neighbouring morphologicalfeaturesso farused to define'life-form'
individuals.In many cases the individualsmay be of may easilymisleadthe investigator into erroneouspre-
separatespecies. sumptionsabout functionaldifferences. A secondprob-
The broad matchof colonial architecture to habitat lem is that bryophytelife-forms may vary markedly
has been recognizedfor many decades (Giesenhagen accordingto the capacityof the substratumto supply
1910, Meusel 1935) but it was not until the work of water.Anotheris thatthe selectionpressuresdetermin-
Giminghamand Robertson(1950) and Giminghamand ing life-form may includeless obvious features,such as
Birse (1957) on 'growth-forms' of Britishbryophytes competition,disturbanceand reproduction,that are
that seriouseffortswere made to classifythe range of largelyindependentof the chiefphysicalfactorsdeter-
colony types. Mdgdefrau(1982) and Richards (1984) miningphotosynthetic production.The presentpaper
extendedthe latterclassification to includetropicaland reviews some of the advantages and drawbacks in
otherextra-British species. Migdefrau (1982) also dis- employinga life-form classification as a framework for
tinguishedclearlybetween'growth-form' and 'life-form' investigating the ecological functionof bryophytes.
in bryophytes.Growth-form (followingMeusel 1935)
was definedas themorphologicalcharacterof theplant
as definedby the positions of its growingpoints,its Life-form synthesis and classification
mode of branching,leaf orientation,etc. Life-form, a
higherlevel of organization,combinesthe featuresof Bryophytecolonies are believedto originatein several
the growth-form differentways although the process has rarelybeen
with the assembly of shoots into
observedfrominceptionto maturity.The synchronous
colonies, and modificationof the resultantform by
local environmentalconditions. Thus, Migdefrau's developmentof new gametophoresfrom numerous
growth-form is concernedprincipallywiththe homolo- gametophoreinitialson a singleprotonemaor on sev-
eral neighbouringprotonematais probablyimportant
gies of structures whereaslife-form deals withthe eco-
formanyacrocarpousspecies(e.g. Marino 1991; see La
logical analogies of whole colonies. Etymologically,
thereis littleto choose betweenthese two termsand Farge-England1996 for definitionsof acrocarpyand
althougha fewrecentreviewers(e.g. Proctorand Smith pleurocarpy).Very dense colonies mightbe supposed
1995) have followed Mdgdefrau'sdefinitions,others always to originatein this manner;however,careful
have not done so (e.g. Longton 1988, During 1990, inspectionsometimesshowsthatyoungcolonies(e.g. of
1992, Grace 1995). During (1979) emphasized 'life Tortulamuralis1on a wall) consistof onlyone or a few
form'as the formof 'plants in accordance with their
environment', followingthe originalviewsof Du Rietz I Nomenclature followsCorley et al. (1981) and Corley and
Crundell(1991) formosses and Smith(1990) forliverworts.
(1931), Barkman (1958) and others,but he did not Othertaxonnamesare used in thesenseof thecitedauthors.

224 OIKOS 82:2 (1998)

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shoots, indicatingthat the next method may often forms recognized and many have introduced new
apply. The second mode of formationis by continued classes,oftenbased on conspicuousmorphologicalfea-
productionof offsetsfroma pioneeringshoot and its tures of particularspecies, that have not been sup-
productssuch that the colony graduallyincreasesin a portedby laterauthors.For instance'tails' (Migdefrau
radial manner. The third means of production of 1982) resembleboth pendantsand fans, and it is not
colonies is wherea numberof adjacent pioneershoots clear that this additional categoryis anythingother
produceoffsets(brancheswithan apical cell of indeter- than a particularlystriking'growth-form' contributing
minategrowth)thatgrowinto each other'sorbit.Syn- to a roughmat or pendantlife-form.
thesisof most (but not all) life-formsinvolvesincrease
in depthof the canopy as thisis the main determinant
of water storage capacity (Proctor 1981). The early Life-form and physicalenvironment
stages of the colony are thus at greatestriskfromthe
effectsof severewaterloss (Alpert 1988) and extended Most authorshave interpreted bryophytelife-forms as
periodsof waterdeficiency. adaptations that maximize water use efficiency,but
Depending on the extentto which coalescence of many have also recognized relationshipsbetween
neighbouringgenets occurs, each of these methods colony organizationand lightintensity,and with the
could produce colonies that are geneticallyuniform typeof substratum.These different aspectswerepartic-
(thereis a possibilityof somaticmutationsintroducing ularlythoroughly investigatedin the studiesof Giming-
geneticdiversity;Salomonson 1996) or whichcomprise ham and Birse (1957) and Birse (1957, 1958a, b). In
severaldistinctgenotypes.Furthermore, it is common some cases bryophytecolonies may trap beneficial
to observebryophyte coloniescomposedof mixturesof above-ambientconcentrations of CO2 fromthe respira-
several species (e.g. the bryophytecarpet in Atlantic tion of microorganisms and invertebrates in the under-
woodland; Rieley et al. 1979). Althoughthe observer lying substratum leading to increased photosynthetic
rarelyhas informationabout the dynamicsof these rates(Sveinbjdrnsson and Oechel 1992,Tarnawskiet al.
situations,thereis evidenceat least forsome Sphagnum 1992).
spp. in miresthat association with shoots of another
speciescan be as beneficialto thewaterrelationsof the
individualshoot as forminga monospecificpatch (Ry- Moisturerelations
din 1985, Li et al. 1992). The positive associations
A relativelydense colony of shoots or thalli(cushions,
occurringbetweenmost bryophytespecies in a boreal
turfs;Table 1) has two obvious advantagesoverwidely-
forestfloorcommunityprobablyalso resultfromthe
spaced individuals.It can store water in the capillary
mutualadvantagesto waterrelationsoutweighing nega-
spaces created betweenthe componentsand its rela-
tive interactions(0kland 1994). In other cases where
tivelysolid formis likelyto become enveloped in a
thereis a relativelystable association of species it is laminar boundary layer (Proctor 1981, 1982). Single
probably more correct to interpretthe mixturesas shoots generate turbulencein a moving air stream
consistingof 'dominants'and 'subordinates',the latter whichinducesrapidevaporativewaterloss. In contrast,
life-formfittinginto an environmentdeterminedto the laminar layer surroundingan aerodynamically
some extentby the former.Good examples are pro- smooth colony becomes saturatedwith water vapour,
videdby thethread-like shootsof smallleafyliverworts this reduces the concentrationgradientfor molecular
(e.g. Cephaloziellaspp.) and some mosses (e.g. Callier- diffusion and thusresistsfurther moisturetransfer from
gonstramineum) growingamongdense,spongycolonies leafto freeair. Measurementsof waterloss fromintact
of Leucobryumglaucumand Sphagnumspp. (e.g. Al- bryophyte coloniesheld in a windtunnel(Proctor1981)
binsson 1997). have confirmedthe considerableadvantageof colonial
The major life-forms recognizedby Giminghamand organizationin increasingaerodynamicresistance.With
Robertson(1950), Giminghamand Birse (1957), Gim- increasingwindspeedsthe relativelysmoothcolonies of
inghamand Smith(1971) and Mdgdefrau(1982), with Ceratodonpurpureus(shortturf)and Grimmia pulvinata
modifications by During (1990) and Proctorand Smith (small cushion) gave evaporationrates close to those
(1995), are describedin Table 1. Short-livedbryophytes predictedforthe standardrelationsof a smoothplane,
(e.g. Acaulon,Buxbaumia,Diphyscium, Ephemerum)of whereas rougher cushions (e.g. Mnium hornum,Di-
disturbedsurfacesoftenformopen turfsthatare some- cranummajus) underwent greaterthanpredictedevapo-
timesclassifiedas a separatelife-form (e.g. 'Annuals'; ration.In the lattercases the divergencefromstandard
Migdefrau 1982), but which clearly representlife- laminar flow commencedat a windspeedwhere the
strategies(During 1979, 1992, Frey and Hensen 1995) laminarboundarylayerhad thinnedapproximatelyto
in whichlittleresourceis investedintolong-term colony the dimensionsof surfaceirregularities of the colony.
organization.These have been omittedfromTable 1. Leaf hair-points,responsiblefor the hoary appear-
No two authors agree preciselyin the range of life- ance of colonies of species like G. pulvinata,

OIKOS 82:2 (1998) 225

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Racomitriumlanuginosumand Tortularuralis,are im- vertical shoots, than in those of other species with
portantin frictionally increasingthe thicknessof the many horizontal axes. Light penetrationwas also
laminarlayerand thus lengthenthe diffusionpathway greaterin most species upon drying,and it was sug-
further.Colonies of G. pulvinatain which the hair- gestedthat,as theuppermostleaves slowlydehydrated,
points had been cut off showed significantly lower a wave of increasedilluminationand productionwould
aerodynamicresistanceand greaterevaporativewater move downwards through the progressivelydrying
loss than intact ones (Proctor 1981). Hyaline hair- colony therebymaximizingproductionpotential.Al-
points are rare or absent in taxa formingloose com- though, the analysis of Proctor (1982) suggeststhat
pared to dense life-forms. hydraulicconductivityof bryophytetissues is suffi-
Although densely-packedcushions and short turfs cientlyhigh to keep water potential approximately
predominatein the most xeric habitats, even these equal throughoutthe gametophore,it is common to
life-forms have stronglimitationswhencomparedwith observeplantsin theprocessof dryingwithsome upper
thecuticularizedand stoma-regulated leaf of a vascular leaves foldingand more shaded lower ones still fully
plant.Thus Proctorand Smith(1995) foundthatcush- expanded.
ions of the xerophyticmoss G. pulvinataon a wall-top Fully exposed shoots carrya highriskof photoinhi-
in south-westEnglandremainedmoistforonlyan hour bition throughgenerationof free radicals especially
or two afterprecipitation in summer,but in the cooler when,because of desiccationor freezing, the excitation
and cloudierautumnand earlywintertheywerecontin- energyof the chlorophyllcannot be quenched by the
uously hydratedfor long periods. The contrastingar- dark reactions of photosynthesis(Oquist and Fork
rangementsof the shoots, radial in cushions and 1982, Kappen et al. 1989). Severalbiochemicalinvesti-
verticalin turfs,may have a functionalsignificance. A gations (Dhindsa and Matowe 1981, Dhindsa 1982,
plausibleinterpretation is thatthe verticalarrangement 1987a, b, 1991, Seel et al. 1991, 1992a, b) have shown
characterizing turfson soil produces the shortestdis- thathighlydesiccation-tolerant taxa possess a rangeof
tances forconductionof moisturefromthe soil to the biochemicalfeaturessuch as anti-oxidantenzymesand
growingapices. Many short-turf bryophytes are endo- enhancedcarotenoid:chlorophyll ratios that help com-
hydricor mixohydric whichsupportsthisview.Alterna- bat oxidativedamage. It is quite possiblethatself-shad-
tively,radial expansionof protonemamay be inhibited ing in dense colonies also plays a part in alleviating
earlieron hard surfacesby droughtso thatincreasein photoinhibitionunder high irradiances.Leaf-folding
densityis mainlyby productionof offsetsfromthe few movements,which are perhaps best developed in spe-
colonizingindividuals.It is also possiblethatthe radial cies withcushionand turflife-forms, probablyplay an
shoot arrangement in small cushionshas advantagesin importantpart too as the tissuesdehydrate.Measure-
colony water storage capacity and in providingan mentsmade by Seel et al. (1992c) showed significant
aerodynamically smoothcushion.Available data, how- reductionsin lightabsorptionby dryingTortularurali-
ever,(e.g. Mankiewicz1987, Smith1988) do not allow formisand Bryumpseudotriquetrum whichwerebelieved
a firmconclusion. to be partlydue to leaf foldingand partlyto a reduc-
tion in tissueabsorptivity.
Where desiccation stress is low, the need to trap
capillarywaterand immersethecolonywithinan envel-
Lightcaptureand photoinhibition
ope of moistair becomesmuch less important.In full
The densely-packedcushion and turflife-forms have lightcushionand turflife-forms stilltend to predomi-
usually been interpreted in termsof theirwater rela- nate (e.g. Giminghamand Birse 1957), probably be-
tions but it is also importantto consider theirlight cause thereis always a danger of dryingin exposed
capturingproperties.Whilst it is obvious that dense habitatsin even the wettestclimates,and photoinhibi-
packingmust resultin shoots receivingless lightthan tion in high irradiancesmay also be importantand
widely-spacedindividuals due to self-shading,few favourlife-forms withself-shading features.In contrast,
quantitativedata are available on this topic. Notable more open life-forms such as weftsand pendantsusu-
exceptionsare the studiesof Skr6et al. (1983) and Van ally occupyshadierenvironments wheretheycan retain
der Hoeven et al. (1993), whichdemonstratethat pho- a hydratedstate for long periods (Birse 1958b, Dilks
tosynthetically activeradiationbecomesattenuatedto a and Proctor1979). Weftsare frequently the dominant
small percentageof the incidentvalue duringpenetra- life-form on the groundin moistboreal-coniferous and
tion of only a few centimetresof a dense bryophyte Atlantic-broadleafforest.Pendants, the life-formin
canopy.Gametophoreorientationwithinthecoloniesis which the photosynthetic cells appear to be the most
also important.In a study of ten Antarcticmosses, stronglyexposed to sunlight,almost always appear to
Davey and Ellis-Evans (1996) found that light pene- require some degree of shading by tree canopies or
trateddeeperinto turflife-forms (Polytrichum alpestre, topography.The pendantlife-form is most evidentin
P. alpinum, Tortula saxicola) and cushions of various typesof tropicaland sub-tropicalcloud forest
Brachythecium austro-salebrosum, each composed of (Richards 1984) where a high atmospherichumidity

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and frequentprecipitationis maintainedfor much of observethattall turfsand mats of bryophytes in mod-
the year. In these situationsit is probable that the erateshade giveway to tiered'fans' of thesame species
narrow,featherystems of these bryophytesscavenge in thedeep shade of wet rock walls and cave entrances.
aerosol dropletsor 'cloudwater'withconsiderableeffi- A similar observation was made for protonemal
ciency(Proctorand Smith 1995). In industrializedre- coloniesof the'coppermoss' Scopelophilacataractaeby
gions this behaviourcould lead to efficient uptake of Takenaka and Satake (1991). Colonies withan unusual
atmosphericpollutantswhich become concentratedin wave-like form recalling the fan life-formwere re-
cloud dropletsparticularly over hill ranges(e.g. Fowler strictedto deeplyshaded verticalstonewalls of a ditch
et al. 1988). A highdepositionof acidic pollutantsin an where incominglight was largelyunidirectional.The
aerosol format an upland oak forestin the English waves had amplitudesof 2-12 mm and wereseparated
Lake Districtwas deducedin thestudyof Farmeret al. by similardistances.Only the upper surfaceof each
(1991) and implicatedin the impoverishment of the wave consisted of living cells with chloroplasts,the
epiphyticLobarion alliance at thissite.The disappear- lowersurfacesbeingpredominantly of dead cells. Mea-
ance of apparently'ultra-sensitive'epiphyticspecieslike surementsand calculations showed that protonemal
Antitrichia curtipendulafrompartsof Europe may have cells on the surfacesof the wave laminae were under
a similarexplanation.Most similarto the pendantsin much higherirradiancesthan if theyhad grownflatly
generalaspect are the aquatic life-forms of bryophytes on the stone surface.
of deep clear lakes such as Sphagnumsubsecundum (Riis As the above examplesshow, many bryophytespe-
and Sand-Jensen1997) wherewater is neverin short cies are extremelyplastic in theirresponsesto condi-
supply,but light levels (and perhaps CO2 concentra- tionsand the balance betweengrowthof the main axis
tion) are extremely low. The life-forms of aquatic spe- and lateral branches,if disturbed,can lead to great
cies in streamsare ratherdifferent, beingdetermined in varietyin the life-forms(During 1990). On the basis of
largepartby drag and abrasionin movingwater(Proc- fieldobservationsand simpleexperiments, Birse(1958b)
tor 1984). concludedthat the ecological amplitudesof bryophyte
In contrastto the foregoing,the noticeablyflattened species were extendedby modificationsto their life-
life-formsof fans and smooth mats show a strong formsinduced by environment. This is borne out by
association with deeply shaded habitats (Birse 1957). some more recent experimentalstudies. A study of
This observationsuggeststhat,as lightis attenuatedby growthof Polytrichum communeunderdifferent condi-
a forestcanopy or by steeptopography,the placement tions of humidityand lightproducedsome contrasting
of the stemsand leaves of bryophytes to maximizelight colony life-formsthat are encountered in nature
capture becomes a major force shaping colony (Sarafis1971). In the weft-forming pleurocarpousmoss
organization. Rhytidiadelphus the spacing of individuals,
triquetrus,
In dry,deep shade, smoothmats are the commonest thedegreeof elongationof determinate lateralbranches
life-formon both soil and hard substrata.Familiar and thefrequencyof offsetproductionare major deter-
examples in Europe are the coveringsof Pseudotaxi- minantsof life-form. Experimentalstudieswith artifi-
phyllumelegans on forestbanks and of Lophocolea cial coloniesstronglysuggestedthattheseattributesare
heterophylla on treebases. They can be interpreted as determinedby a simple trade-offbetween moisture
essentiallysinglelayeredcanopies thatmaintaina posi- conservation(favoursclose spacing) and lightcapture
tive carbon balance by possessinglittlenon-photosyn- (favourswide spacing) in each situation(Bates 1988).
theticbiomass. Their groundhuggingformmaximizes Somewhatsimilarrelationshipswerefoundfornatural
eithercapillarymoistureentrapmentagainst,or water clones of Hylocomiumsplendensin Norwegianspruce
conductionfrom,the underlyingsurface,but it may forests(0kland and 0kland 1996).
also reflectan absence of non-photosythetic stemsor
othersupportingtissues.Numerousthalloidand leafy
liverworts, withrelatively poor toleranceof desiccation,
Regulationof colonyform
have adopted this arrangementin deeply shaded and
even relativelymoist foresthabitats. The absence of What is the physiologicalbasis of the variationsin
bryophytes frommanyforestfloorareas can be related life-formcaused by differentmoisture regimes and
to burialunderleaflitter(Duringand Verschuren1988, irradiances?
0kland 1995). In the case of moisture,the stronggrowth-retarding
In constantlyhumid conditions,close contact with influenceof desiccationon any shoots that protrude
the substratumis less importantfor maintenanceof fromthe main canopy is probably a major force in
hydration,and it is undermoist,deeplyshaded condi- ensuringthat colonies conservea reasonablyaerody-
tions that fans become important.Although certain namically smooth surface. For instance,in artificial
taxa (see Table 1) onlyeveroccur as fans,manyothers coloniesof themoss Rhytidiadelphus triquetrus
themar-
are more plastic and may assume a fan formwhen ginal shootsgrewless thanthecentralones due to their
growingin deep shade. In forest,it is common to more frequentdehydration(Bates 1988). The typically

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roundedprofilesof manybryophyte coloniesmay arise thelightbeam would occurand explainthetendencyto
in thisway. Similarly,in coloniesof Hylocomium splen- grow away fromthe substratum.
dens the mean size of segmentsincreasedwithincreas- stemsand thalliof gametophores
In themulticellular
ingbryophyte cover,a reflectionof themorefavourable theeventsthatcause growthcurvaturesdifferconsider-
moistureconditions that a large colony size brings ably fromthosein singlecells of theprotonemasystem.
(Okland and 0kland 1996). The 'dryingpower' of the However,theystillappear to be primarily governedby
air depends on atmospherichumidity,net incoming the ratio of red:farred light and mediated by phy-
radiation and windspeed,and it varies considerably tochrome.Fredericqand De Greef(citedby Hartmann
between adjacent microenvironments (Monteith and and Jenkins 1984) showed that, whereas thalli of
Unsworth 1990). Such variationsprobably determine Marchantiapolymorphagrewnearlyhorizontallyunder
whetheran epiphytelike Frullaniatamarisciremainsas whitelight,a briefexposureto far-redlightcaused the
a smoothmat or, as it does in extremely humidhabitats apices to reorientateto a nearlyverticalgrowthdirec-
in Europe and Macaronesia,developspendantregions. tion. Simpleresponsessuch as thiscould easilyexplain
Commonly,the two life-forms of F. tamariscimay be the adoption of a fan life-formin deep shade and,
observed in contrastedconditionsa few centimetres togetherwithotheretiolationresponses(Hoddinottand
apart. However,more complex responsesto moisture Bain 1979),accountformanyapparentlyadvantageous
stressalso exist. Birse (1957) found that a numberof alterationsof life-form observedunderdifferent condi-
speciesrespondedto low humidity, at least underhigh tions. As Birse's (1957) studyof Climaciumdendroides
light,by growingin close contactwiththe substratum. suggests,significantadjustmentsof the life-form may
The dendroidClimaciumdendroidesproduces'stolons' accompanyseasonal fluctuations in conditions.This is
that at firstgrow horizontally(plagiotropic)but then perhapsalso reflected in the verydifferent lightextinc-
turnupwards(orthotropicgrowth)and branchto form tion coefficients found for colonies of chalk grassland
the characteristic shoots of thismoss. In sand mosses in December and September(Van der Hoeven
'fir-tree'
cultures,Birse (1957) was able to prolong the pla- et al. 1993) and is a possibilitythat meritsfurther
giotropicphase indefinitely by maintaininga low hu- investigation.
midityin the atmosphereabove the moistsubstratum.
It was hypothesized that,in drydune habitats,seasonal
increasesin air humiditytriggerthe formationof new
A simplelife-form/environment model
ascending shoots each year. In another dendroid,
Thamnobryum alopecurum,growth was very limited As many studieshave emphasizedthe close match of
even when the plants were maintainedconstantlyat bryophyte life-formswithparticularenvironmental con-
100%/orelativehumidity.In thisspecies,frequentspray- ditions,it is surprising to discoverthatno formalmodel
ing with liquid water appeared to be crucial for full of thisrelationshiphas ever been proposed. Construc-
developmentof thedendroidlife-form. This is in agree- tion of a hypotheticalmodel is a logical step on the
mentwiththe observationthatbryophytes, in contrast pathway to improvedunderstandingas it makes the
to some algae and lichens,requireliquid waterrather topic more readilyaccessibleto experimental investiga-
than highhumidityto become photosynthetically com- tion.A simplegraphicalrepresentation of thelife-form/
petentfollowingdesiccation(Rundel and Lange 1980). environment relationshipsbased on existingknowledge
The effectsof shadingon developmentof bryophytes is therefore attemptedhere.From the foregoingreview
are not well known but thereis some evidence of a it is evidentthatvariationsin moistureavailabilityand
varied responsedependenton ecology(Hoddinottand lightintensity are themajorenvironmental factorsto be
Bain 1979, Bates 1988) as in floweringplants (Grime considered.However,a formidablecomplicationis in-
1966). 0kland and 0kland (1996) concludedthatvaria- troducedby the arrayof different substrataon which
tions in regenerationfrequencyof Hylocomiumsplen- bryophytes grow.
densweftswererelatedto density-dependent regulation
of budding.In this case shadingat high densitieswas
believedto inhibitbranching.Controlof offsetprolifer- Influenceof substratum
ation and lateral branchingis almost certainlyunder
the controlof phytochromein bryophytes(e.g. Hart- Considerationof a wealthof publisheddata on substra-
mann and Jenkins1984), with far-red-enriched 'shade tum requirements(e.g. Barkman 1958, Smith 1982,
light'inhibitingbud developmentand red lightpromot- Palmer 1986, Bates 1992a, b, and referencestherein)
ing the process. In the case of the protonemataof shows that, apart from a small number of extreme
Scopelophilacataractae growingin waves, Takenaka specialists(e.g. mossesthatgrowon dung or metallifer-
and Satake (1991) suggestedthat,by analogywiththe ous rocks; see Brown 1982), bryophytesare rarely
growthof the caulonemata of Physcomitrella patens exclusivelytiedto a specificsurface.For instance,most
(Jenkinsand Cove 1983),underthelow intensity unidi- epiphytescannot be considered 'obligate epiphytes'.
rectionallightin thishabitat,growthat rightanglesto Indeed, 'climax' communitiesof epiphyticbryophytes

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are invariablyformedby 'facultativeepiphytes',species dominant tracheophytecanopy. Thus, whereas on
that also commonlyoccur on other substrata(Smith 'hard' substrataa range of bryophytelife-forms has
1982,Bates et al. 1997). Instead,successon any reason- evolvedin responseto variouslevelsof moisturesupply
ably stable substratumdepends on two main factors: and irradiance,on soils bryophytes are largelyrestricted
(1) whetheror not the bryophyteneeds to exploitany to habitat'windows'characterizedby poor conditions,
water store that the substratumholds, (2) whetheror or for briefperiodsin 'gaps' (During 1979), and fewer
not the surfacesatisfiesany chemicalrequirements of formsare represented.'Poor conditions'('stress' sensu
thebryophyte, withthepH rangebeingnotablyimpor- Grimeet al. 1990) mainlyincludesdroughted(continu-
tant for many taxa (Bates 1992b). Often the set of ously or seasonally) and/or shaded habitats, but in
requirements of a particularspecies can be met by a other ecological dimensionshighlyacid, wet environ-
range of different surfaceswith, for instance,occur- ments (Sphagnum mires), extremelyhot (volcanic
renceson both rock and bark beingcommon.A third springs) and extremelycold (tundra) habitats all
factor,the period forwhichthe surfaceis available, is provide niches that are largely unoccupied by tra-
of great importance ecologically (e.g. Herben and cheophytes.Bryophytes usuallyonlybecomeimportant
Soderstrbm1992), but is arguablyof more importance on soils under good conditionsfor short periods by
to overalllife-strategy(During 1979, 1992) thanit is to occupyingtemporary gaps formedin thevascularplant
thesuccessof a givenlife-form. thepersis- canopy, althoughpersistencein continuallydisturbed
Nevertheless,
tent occurrenceof short turfsand smooth mats may habitats is also frequent.The latter include heavily
sometimesbe related to frequentsmall-scale distur- selectively-grazed (e.g. thegroundfloraof manyupland
bances ratherthan physicalfactors(During and Ter oak-woods and Chalk grasslandin Britain)or herbi-
Horst 1985). There is littleevidence to suggestthat cide-treated(fruitfarmsand moss gardens)situations.
substratumchemistryis an importantfactorin deter- The 'fugitive','annual shuttle'and 'colonist'life-strate-
miningbryophytelife-form. Whetheror not the sub- gies of During (1979, 1992) includemost of the plants
stratummay hold a storeof waterthatcan be exploited capable of growingin gaps and the majorityhave an
by plants,however,appears to be of considerableim- open turf or thalloid mat life-form.As their main
portance.A fairlyclear distinctionexistsbetweenrela- period of growthis short and generallycorresponds
tivelyimpermeablesurfaceslike rock, bark and leaf withthe moistseason, the selectionforcolonial assem-
cuticleson the one hand, and soils withtheirstoresof blages is relativelyweak (During 1979). A widerrange
capillarywateron the other.This importantdistinction of perenniallife-forms occurs in continuallydisturbed
has long been recognizedin Buch's (1945, 1947) classifi- habitats, depending on the physical conditions
cation of bryophytesinto ectohydricand endohydric experienced.
types,theformeroftenexploitingimpermeablesurfaces
(or having poor contact with the surface),the latter
exploitingsoil waterto varyingextents.Rocks and tree
The model
barkshold relativelysmallquantitiesof available water
which, nevertheless,in some cases appear to be of The ideas discussed in this and the precedingsection
importancein determiningwhich species may grow are summarizedin Fig. 1 whichshowsthemajor peren-
(Smith 1982), but few data on this aspect have been nial bryophytelife-forms of Table 1 positionedwith
published. The rather differentbryophytefloras of respectto notional gradientsof moisturesupply and
rocksand barkscan probablybe attributedpartlyto a irradiance.Separate diagramsare given for hard sub-
tendencyforbarks to have the higherwaterabsorben- strataand soils in recognitionof the significant mois-
cies, although climatic conditions may determine turereservoirpotentiallyprovidedby thelatterand the
whetheror not this factorassumes importance.Thus, excludingeffecton soils of vascular plants. Life-forms
the main effectof the substratumfactorin relationto in the greyarea of the 'soils' diagramoccur onlywhere
life-forms is the abilityof soils to contributeto the thereare special factorsoperating(e.g. extremeacidity
moisture relationshipsof bryophytesin contrast to in mires)to reducethesuccessof tracheophytes. Similar
impermeablesurfaceswhichgenerallydo not do so. life-forms
do not necessarilyoccupyidenticalpositions
One other substratum-related factor has probably in the two diagramsowing to the moisture-enhancing
also influencedthe evolutionof bryophytelife-forms, effectof soils comparedto hard substrata.For simplic-
theirexclusionby vascular plants on soils but not on itythe environmental axes have been representedwith
hard (and generallywater-impermeable) substratathat just threelevels of intensity.The life-forms have been
are impenetrableto roots. Bryophytesare effectively positionedon thebasis of theexistingliterature and the
excluded fromparticipatingin a major way in plant author'sfieldexperience.Giminghamand Birse (1957)
communitieson soil whereresourcesupplyis good and and Birse (1957, 1958a, b) providedthe most compre-
disturbanceinfrequent(Grime et al. 1990) because of hensiveset of examples by studyingthe variationsin
the impositionof reducedradiation,loweredmoisture life-formalong transectsaligned with environmental
inputsand increaseddisturbancesfromlitterfallby the gradients.An 'experimental'element has also been

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 Hardsubstrata Soils
Fans a)Pnats c)Talltufs a) Fans Dend
b) Tallturfs

5 ~~~~~~~~~~(Large
cushions)
a)
Dendroidss Rough
b) Wefts mats
tia

co

Thalloid& Rough Small Thalloid& cushions)Shortturfs

smoothmats mtcuhossmooth mats d) Tallturfs rohmt

Deepshade Fullsunlight Deepshade Fullsunlight

Irradiance
Fig.1. Approximate ofbryophyte
relationships life-forms ofmoisture
to gradients andirradiance
availability on hardsubstrata
andsoils.'Wet'refers
tohabitatswhere desiccationstress
is rarely In theright-hand
encountered. graph,thegreyarearepresents
habitatsfromwhichbryophytes are normally excludedby negative withvascularplants.Key: Arrowsdenote
interactions
extending uptoandsometimes beyond thenextcategory; inparentheses
life-forms a) Life-form
occurinfrequently: onveryfreely
draining,inclined b) life-form
surfaces, on levelsurfaces,
c) extremelydenseorrichlybranched e.g.Sphagnum,
versions, d) open
or little-branched e.g.Mnium
variants, hornum.

providedby a description of the changesin the inconsistencies in theapplication of theexisting life-

bryophyte life-formsofvarioushabitatsaccompanyingform classifications,(2) thefactthatthelife-form repre-
thefelling oftreesina woodedravine(Gimingham and sentsonly one aspectof a widerrangeof traits
Birse1957). characterizing 'life-strategy'.
Severaldilemmas werefacedinplacingthelife-forms As alreadydetermined, thereis no overallagreement
in Fig. 1. For instance,weftsare shownas one ofthe in therangeof existing life-forms norin theirprecise
mostcharacteristic life-formsoccurringundermoist definitions. In practice, theallocationof life-forms to
and moderately shadedconditions. Nevertheless,Birse thedifferent bryophyte taxa growing in a particular
(1958a) foundwefts(Hyiocomiumsplendens,Rhytidi- community can be a problematic exerciseand names
adelphusspp.) in an unshadedScottishdune slack may oftenbe wrongly,or at least approximately, ap-
wherethe watertable providedsignificant seasonal plied.It is clearfromtheliterature thateventhemain
moisture inputs.Thus,althoughthe arrangement in proponents oflife-forms haveexperienced of
difficulties
Fig. 1 fitsmanyactualcircumstances, it willprobably thissort.For instance, Migdefrau(1982) provided,
notbe difficultto findexampleswherethelife-forms do undertheheading'lifeforms', illustrationsdepicting,
not respondto lightand moisture gradientsas indi- strictly
speaking, a mixture of growth- and life-forms.
cated.Reasonsfortheoccasionally unpredictable be- Thefailure ofmostearlier writersto distinguish clearly
haviourofbryophyte life-forms
arediscussedbelow. between thesetwolevelsoforganization has ledto the
presentunsatisfactory position. The situation is made
moredifficult by the tendencies of manyspeciesto
adoptdifferent life-forms indifferent conditions.Exam-
of thelife-form
Limitations concept ples include the familiarmoss Climaciumdendroides,
dendroid
characteristically onshadedandmoiststream-
Inadequaciesof existingclassifications sidesbutforming tallturfon
a kindof Sphagnum-like
Life-formshave particular
importance to bryophytessand dunes in some regions,and Grim ia antarctici,
becauseof theirhighrelianceon transientwatersup- whichforms looseturfsonmoistground andnearlakes
in thepracticaluse of thelife-formand meltwater
plies.Difficulties streamsbut existsas densely-packed
approachcan be ascribedto two main causes: (1) cushionsin driersites(Kappenet al. 1989,Tarnawski

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et al. 1992). A furtherapparentobstacleto understand- An alternativeand probably more promisingap-
ing is thatin manycommunitiesthecommonestspecies proach would be to describethecanopyin termsof the
may exhibitquite different e.g. the tall turfs
life-forms, volume of livingtissue supportedabove unit area of
of Dicranummajus oftenassociate withweftsof Rhy- substratum. Biomass is imprecise because of the
tidiadelphus loreusin Atlanticoak woodland. difficultyin bryophytesof knowingwhetherolder tis-
In reality,quite differentlife-forms
may provideap- sues are livingor dead and, althoughthe extentof the
proximately the same netphotosynthetic benefitin par- 'green zone' appears to be a reasonable predictorof
ticularzones of the conditionsrepresentedin Fig. 1. favourableenvironmental conditions(Plantefol1927),
The concept of life-formin the foregoingtreatment chlorophyllcontentis known to vary markedlyin re-
impliesthe possessionof analogous structures resulting sponse to seasonal alterationsof the light level (e.g.
from convergentevolution. However, perceptionsof Kershaw and Webber 1986, Martinez-Abaigaret al.
life-form made by a human observerare also likelyto 1994). Assays of anothermarkerchemicalwhose levels
be influencedstronglyby the evolutionaryhomologies remain relativelystable withinliving cells might be
of taxa. In otherwords, the human observermay be developed to determinebiomass indirectly.Investiga-
blindedto the functionalsimilarities thatexistbetween, tions of the quantitiesof living tissues that can be
say, a Dicranumturfand a Rhytidiadelphus weftbe- supportedin each situationthroughthe medium of
cause of theirgreatlydifferent growth-forms,the result differentlife-formscould give valuable insights.
of wide taxonomicseparation.Clearly,the adoption of
more objectivelydefineddescriptionsof the various
life-forms is desirable.
Life-form
and competitive
status
A serious objection could be raised to the life-form/
Improvements
of life-form
classification
environment model if it could be shown that life-form
How could the life-form schemebe refinedto improve was strongly linkedto otherattributes of life-strategy
in
the correlationwith environmentalconditions?There bryophytes.Besides the stressesimposed by limited
would appear to be two possibilities.The firstinvolves lightand water supply,other selectionpressuresthat
improvingthe definitions of the differentcategoriesby mightbe expectedto influencelife-form includecom-
quantifyingdimensions,shoot orientationsand densi- petitive relations, foraging for resources and
ties. An improvedlife-form schememightbe preceded reproduction.
withadvantageby a classification of growth-forms that Competition has been relativelylittle studied in
emphasized the differencesand inter-relationships of bryophytes whichare largelystress-tolerators and rud-
the two levels of organization.The second and prefer- erals (Grime et al. 1990). Even so, quite large differ-
able possibilitywould be to adopt a more rigorously ences exist in the productionpotentialsof different
functionalapproach whilstretainingthe existingap- species (e.g. Rincon and Grime 1989) and evidenceis
proximateframework alongside.Thus life-forms would accumulatingthat interspecificcompetitionmay be
be definedby the natureof the photosynthetic canopy, stronglyrelevantin many situations(Scandrettand
especiallyits depthand density. Gimingham1989, Marino 1991, Li et al. 1992, Litke
For tracheophytes the leaf area index (L) has long Twenhdven1992,Rydin 1993a, Li and Vitt 1994, 1995,
been employedto aid descriptionof the complexities Wilson et al. 1995), althoughin boreal forestsrecent
and efficienciesof contrastingvegetation canopies evidencesuggestscompetitionis unimportant(0kland
(Kvet and Marshall 1971, Fitterand Hay 1981, Nor- 1994, 1995, 1997). In comparisonwith the extensive
man and Campbell 1989). Applyingthesame approach tracts of relativelyuniformland colonized by many
to bryophytes is problematicand it has been attempted tracheophytes,bryophytesare predominantlyoccu-
only rarely (e.g. Proctor 1981, Simon 1987). Most pants of patchy habitats (Herben 1994). Colonial
bryophytes have leaves thatare onlyone cell thickover growthhas been shownto confera competitiveadvan-
most of the lamina, but thalloid species and some tage to encrusting marinealgae (Olson and Lubchenco
mosses (e.g. Polytrichum) have severallayersof photo- 1990, Paine 1990) and invertebrates (Jackson1977) on
syntheticcells. Other difficulties are provided by the hard substrata by pre-emptingthe available space
complexleaves of mosses like Leucobryumand Sphag- patches. In an investigationof competitionbetween
num where the photosynthetic cells are separated by Splachnaceae on the dung of large mammals,Marino
water-storing hyalocysts,and by the factthatshootsin (1991) argued that pre-emptionof patches,by clonal
leafykinds also oftenhave significant photosynthetic expansion,may be the primarymechanismby which
capacity.Thus directmeasurements of L, whichare in the uprightstems of many acrocarpous mosses com-
any case difficultto make accuratelybecause of the pete. Rydin (1993a, b) also considered that clonal
small sizes of bryophytes, mightbe misleadingif they growth was important in pre-emptivecompetition
involvenon-comparableunits. among Sphagnumspeciesin mires.

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 The dendroidlife-form can be picked out as one in training leachates fromthe tree canopy (Bates 1992b).
which selectionfor competitiveabilitymay have been The highlyproductiveBrachythecium rutabulumap-
as importantas fitnessforthephysicalenvironment. In pears to forageactivelyfornutrientswhichit exploits,
temperatecountriesdendroidsare probably the least in dead plant matter,by means of rhizoids (Rincon
common life-form (Birse 1957), yet where theyoccur 1988). This pleurocarpousmoss has a highlyplastic
dendroidsmay predominate.In Europe examples are formwith almost any lateral branch able to develop
provided by Isothecium alopecuroides and L into a main axis of indeterminate growthas foraging
myosuroides on moistforesttreebases, and by Thamno- proceeds. In contrast,the slower growingforestand
bryumalopecurumand Isotheciumholtii,respectively on grasslandmoss Pseudoscleropodium purum,which ap-
calcareous and acid rocks by waterfalls.In North pears to be relianton wetdepositionof nutrients alone,
America ClimaciumamericanumBrid. is characteristic retainsrelatively constantgrowth-and life-forms under
of similarlytightly circumscribed habitatson verymoist varied conditions and appears better able than B.
and well shaded, but litter-free, forestsoils (Sholl and rutabulum to redistribute
nutrientresourcesfromold to
Ives 1973). Each of these species commonlyformsan new tissues(Bates 1994, 1997).
extensivemonoculture.The horizontalshootsproduced The sharingof nutrientresourcesby different parts
by dendroid bryophytesmay be as much concerned of a branching,colonial shoot systemis likelyto occur
withcolonizingstrategy, allowingpenetration and over- in bryophytesby analogy with vascular plants (e.g.
topping of existingbryophytepatches as they are in Alpert1996). For instance,in theclonal herbGlechoma
underpinningan optimal photosynthetic canopy. An hederacea thereis strongevidencethat the abilityof
experimentalinvestigationof growthplasticityin rela- some rametsto exploit locally favourableconditions
tion to light interceptionby six mosses found T. increasesthe fitnessand competitiveness of the whole
alopecurum, the onlydendroidstudied,to be unable to clone (Slade and Hutchings1987a, b). Moreover,the
concentrateits biomass in small areas withpredictably life-formof Glechomahederacea is highlyplastic in
higher light input than the dimly-litsurroundings response to patchinessin the availabilityof mineral
(Rincon and Grime 1989). However, growth of T. nutrients (Slade and Hutchings1987c),light(Slade and
alopecurum and another 'shade' moss, Fissidens Hutchings1987d)or both(Slade and Hutchings1987b).
cristatus,were significantly stimulatedwhen a random There is evidence for internalmovementsof carbon
'sunfleck'componentof higherlightintensitywas su- compounds and phosphorus within bryophytesthat
perimposedon the low backgroundillumination.This lack obvious internal conduction pathways (Alpert
suggeststhat physiologicaladaptations to low irradi- 1989, Rydin and Clymo 1989). It has recentlybeen
ances may be more importantthan foragingfor re- shownthatmosses (but not liverworts) possess unique,
sourcesin dendroids.Nevertheless, theirinherentmode polarised 'food conducting'cells that may effectsuch
of growth,formingdense clones and expandingout- transport(Ligrone and Duckett 1994, 1996) although
wards by runners, combines both 'phalanx' and thishas not yetbeen demonstrated forinorganicnutri-
'guerilla'tactics(see LovettDoust 1981,Bell 1984),and ents.The possessionof systemsfornutrientmovement
may be vitallyimportantin preemptingand capturing could be an importantfactorin the selectionof differ-
space. ent bryophytelife-forms.
Life-forms have receivedsome considerationin clas-
sificationsof bryophyte In his influential
life-strategies.
paper on bryophytelife-strategies, During (1979) al-
Life-form and foragingforresources
luded to a strongcorrelationbetweenlife-form (called
The capacity to forage for resourcesis a recognized 'growthform')and life-strategy. For example,the pro-
attributeof successfulcompetitors.Rincon and Grime ductionof dense turfsobviouslyaccompanieslong per-
(1989) obtainedstrongconcentrations of growthin the sistence,whilstthe capacityfor horizontalspread was
better-litpatches for the highly productive mosses seen as enhancingcompetitiveability.During (1979)
Brachythecium rutabulum and Thuidiumtamariscinum in listedthe life-forms foundwithineach of the six main
their experiments.They concluded, contraryto the life-strategies of bryophytesrecognized. Thus: den-
hypothesisof Birse (1957, 1958a, b), Giminghamand droids,largecushionsand weftsare restricted to 'peren-
Birse (1957), Gimingham and Smith (1971) and nial stayers';shortturfsare foundamongst'colonists',
Schofield(1981) that the observed growthplasticity 'short-livedshuttlespecies' and, rarely,'annual shuttle
represented'foragingbehaviour' rather than adjust- species';cushionsare represented in the 'perennialshut-
ment to maintainwater status. Acquisitionof scarce tle species' and 'perennialstayers'categoriesonly. In-
nutrientresourcesmay be another factorinfluencing formationon characteristiclife-forms was omittedin
life-form.The richlybranched surfacesof the forest the revisedversionof thisscheme(During 1992) but it
floor bryophytes in boreal-coniferand Atlantic- has been reinstatedin a variantof During's classifica-
broadleaf forestmay have significancein capturing tion generalizedto include all plant groups (Frey and
nutrients retainedin precipitation and throughfall con- Hensen 1995). The latterauthorsassociate well-devel-
OIKOS 82:2 (1998) 233

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oped life-forms withthelongerlived'stresstolerance' and Longton's (1995) analysis, for the dioicous
life-strategiesbut do not significantly advance our mosses to have a greatertendencyto produce asexual
knowledge of thesignificance of particularcolonyar- propagulescould also be relatedto thisfactor.
rangements. The correlationwith life-strategy ob- The life-formof the gametophyte,usually con-
served by the foregoingauthors suggeststhat cerned with the maintenanceof an envelope of still
life-form is by no meansa matterof fitness to mois- air duringwindy conditions,is oftenin strongcon-
tureand lightconditions only,butthatit is probably trast to the functionof the sporophytewhose main
determined by trade-offs involving severalotheras- job is to deposit spores in the turbulentair above the
pectsof the lifecycle.A moreobjectiveanalysisof boundary layer. It is probably not an accident that
thepatterns of lifehistory variationin theimportantthe characteristicvascular plant features,cuticle and
moss ordersFunariales,Polytrichales and Pottiales stomata, are predominantlysporophytefeatures in
was recently undertaken by Heddersonand Longton bryophytes.Proctor (1984) has drawn attentionto
(1995). Multivariate analysesweremadebased on 13 correlations between gametophyte life-formsand
lifehistory traits,including 'growthform',scoredfor sporophytemorphologies,such as the tendencyof
357 species.'Growthform'was simplified into four cushion-forming plants in exposed habitats to have
categories('scatteredindividuals'; 'gregarious'; 'loose erect capsules, shortersetae and simplerperistomes
turves';'denseturvesor cushions')ratherthanapply- than weft species in more shelteredhabitats. Repro-
ing one of therecognized systems (e.g. Table 1). Al- ductivefrequencyis now knownto be a key factorin
thoughthe resultssuggestthat simplelife-strategy abundance and persistenceof bryophytes (Herben and
classificationslike that of During(1979, 1992) are Sbderstrdm1992, Longton 1992, Herben 1994). The
realistic,
therewas littleexplicitindicationthatlife- extentto which the evolutionof reproductivemecha-
formcontributed significantlyto the maincategories nisms has become interwovenwith that determining
or thatit correlated withotherattributes (like leaf gametophytelife-forms offersa difficultbut fascinat-
papillosity) concerned withwaterrelations. This may ing fieldforfurtherstudy.
be a reflection of theapproachadopted,or indicatea
relativeuniformity of life-form in the threeorders
selectedforstudy.Clearlythistopicdeservesfurther
investigation. Conclusions
The foregoinganalysis leads to an equivocal answer
to the question posed in the title of this paper. The
Life-form and reproduction life-formsof bryophytes correlateremarkablystrongly
Despite the inconclusive resultsof Heddersonand withgradientsof moistureavailabilityand lightinten-
Longton's(1995)studywithrespectto life-form there sity, and their study at an advanced or elementary
is good reasonto supposethatcolonytypewillshow level can continueto provide valuable ecological in-
correlations withmodeof reproduction and evenwith sights (Grace 1995). Moreover, Proctor (1984) ob-
particularbreedingsystems.Sexual reproduction in served that each life-formoffersdifferentecological
bryophytes occursbya primitive mechanism involving problems. Potential studies range from the survival
externalwaterdroplets, 'rainsplash',to ef- and demographyof whole colonies in different
especially habi-
fectspermtransfer fromantheridium to archegonium.tats, to the rhythmof mortalityand ramet replace-
Althoughabout 40% of bryophytes are monoicous ment in maintenanceof colony integrity(e.g. 0kland
(roughlyequivalentto 'monoecious'in angiosperms,1995, 1997, 0kland and 0kland 1996). Knowledge of
see Wyattand Anderson1984), rathermore are these areas for bryophytesis very sparse and much
dioicous (equivalentto 'dioecious'). In the latter furtherwork is needed. Nevertheless,life-formsare
cases, wherethereare strongselectionpressures for undeniablylinkedwithotheraspects of plant function
clonalproliferation, maybe producedthat and improvedunderstandingin the futurewill be de-
life-forms
are not onlymonospecific, but also oftencomposed pendenton the adoption of an holisticplant-strategy
of one sex. The limitations of the spermtransferapproach. Hedderson and Longton (1995) found life
mechanism inevitablymeanthatthiswillrestrict suc- historyvariationin the mosses thattheystudiedto be
cessfulfertilization.
This mayhave led to subsequent stronglyinfluencedby phylogeny.This suggestion
adaptations of thereproductive
mechanism itself.The raises interestingquestions about the relationshipsof
dwarfepiphytic male shootsof Leucobryum and Di- life-formsto other traits such as the possession of
cranumspp. could be the productof a selection desiccation-tolerance, and about the extentto which
trade-offallowingboththesynthesis of large,compet- these are co-evolved phenomena or whetherposses-
itive,water-entrapping coloniesand the retention of sion of one attributeprecededand dictatedthe evolu-
sexual reproduction. The observation, in Hedderson tion of others.

234 OIKOS 82:2 (1998)

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Acknowledgements- I am indebtedto H. J. During, R. H. Dhindsa,R. S. 1987b.Protein
synthesis
duringrehydrationof
0klandand M. C. F. Proctor
fortheircritical
comments and rapidlydriedTortula Evidence
ruralis. foroxidationinjury.
suggestions
forimprovements of themanuscript. - PlantPhysiol.85: 1094-1098.
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