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What Are Rare Species and Why Conserve Them?

Article in Lindbergia · January 2000

DOI: 10.2307/20150038

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What Are Rare Species and Why Conserve Them?

Author(s): Royce E. Longton and Terry A. Hedderson
Reviewed work(s):
Source: Lindbergia, Vol. 25, No. 2/3, The Scientific Basis for Bryophyte Conservation: A
Symposium (2000), pp. 53-61
Published by: Oikos Editorial Office
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LINDBERGIA 25: 53-61. Lund 2000

What are rare species and why conserve them?

Royce E. Longton and Terry A. Hedderson

Longton, R. E. and Hedderson, T. A. 2000. What are rare and why

species
conserve -
them? Lindbergia 25: 53-61.

This paper considers

the concept of rarity in bryophytes from two inter-related
viewpoints: 1) distribution and abundance, and 2) a complex of factors relating
to population biology and extent of genetic variation. A series of questions is
posed, a fundamental one being "why do we wish to conserve rare species?".

Following Rabinowitz, a species may be classed as rare if its distribution fits

one of the three criteria of narrow or highly range,
disjunctive geographical
high habitat or consistently low population size. Some are
R. E. Longton T. A. Hedderson specificity, species
rare on all three counts. Others are classed as rare when they fit two or even
only one of these criteria. Most species would be considered rare on this basis.
There is evidence of correlation between rarity and failure to produce sporo
phytes, among species of both mosses and liverworts. This relationship holds
good only for dioecious taxa, however, and thus it is not clear to what extent

rarity results from

restricted diversity among genotypes consequent upon a
bypassing of meiosis. Some rare bryophytes are long-lived but many
perennials,
are colonist or shuttle species with short-lived Provi
relatively gametophytes.
sion of successional habitats will be essential in their conservation manage
ment. Some rare species may be phylogenetically isolated, while other may be
members of
large assemblages of closely related, possibly currently evolving
species in genera such as Bryum.
Molecular studies, such as our work on the striking pattern of variation within
the cosmopolitan species Bryum argenteum, have shown that many common

bryophyte species exhibit greater degrees of genetic diversity than might be

expected in haploid-dominant organisms. This raises the question whether con

serving races of common

but genetically variable taxa such as B. argenteum

might not be at as valuable

least as conserving some rare but more uniform
species that are closely related to common ones. It also emphasises that our
efforts should be directed towards maintaining the largest possible areas of
natural and semi-natural as reservoirs of genetic at the
vegetation diversity
population level, as well as towards conserving priority species.

R. E. Longton and T. A. Hedderson, Centre for Plant Diversity and Systemat

ics, Dept of Botany, Univ. of Reading, Reading, UK RG6 6AS (present address
ofTAH: Dept. of Botany, Univ. of Cape Town, Private Bag, Rondebosch 7701,
South Africa).

Accepted 7 September 1999

? LINDBERGIA 2000

LINDBERGIA 25:2-3 (2000) 53

Table 1. Types of rarity (Rabinowitz 1981) as exemplified by European mosses. The figures 1-3 indicate how many of
the three criteria of rarity (narrow range, high habitat specificity, consistently low population size) are fulfilled. Species
like Pleurozium schreberi that score 0 are not rare.

Wide geographical range Narrow geographical range*

Low habitat High habitat Low habitat High habitat
specificity specificity specificity specificity

Large populations Pleurozium Schistidium Campylopus Zygodon

schreberi (0) maritimum (1) setifolius (1) gracilis (2)

Small populations Anomobryum Zygodon Weissia Ditrichum

filiforme (1) forsten (2) multicapsularis (2) cornubicum (3)

*
includes endemic and widely disjunct distributions.

In this paper we shall consider the concept of rarity, ble 1 by Zygodon gracilis, a species that locally forms
as applied to bryophyte species, from two interrelated substantial populations on limestone rocks and walls
viewpoints: 1) distribution and abundance, and 2) a in Britain and in parts of southern and central Europe.
complex of factors relating to population biology and This pattern of rarity is shown by a fair number of
extent of genetic variation. This discussion will lead other species such as Barbula maxima, Dicranodon
us to pose a series of questions concerning the origins tium subporodictyon, Grimmia retracta and Plagi
of rarity, and the rationale underlying species conser ochila atl?ntica.
vation. A converse pattern shown by geographically re
stricted species that occupy a range of habitats, but
usually as small populations, appears not to be repre
sented among theNorth American vascular flora con
sidered by Rabinowitz (1981), and yet is shown by
Distribution and abundance several European bryophytes. Weissia multicapsularis
Types of rarity (Table 1) is known only from southern England and
The concept of rarity is a familiar one. In some cases France. It is seldom abundant, but occurs in a range
it is readily defined. A postage stamp is rare if few of non-calcareous habitats including woodland rides,
copies are known to exist. Rarity is then defined by a fields, brick pits, earth banks and sea cliffs (Hill et al.
single parameter, and may even be quantifiable. With 1992). Other examples include Fossombronia fim
biological species the situation ismore complex, and briata and Lejeunea mandonii.
several aspects of distribution and abundance have Campylopus setifolius is an example of a geographi
been used to define rarity. A familiar example is the cally restricted species that is not rare on either of the
scheme of Rabinowitz (1981) who recognised seven two other criteria, being locally abundant in a variety
types of rarity based on combinations of the three cri of acid habitats in the north-west of the British Isles
teria: 1) narrow geographical range; 2) high habitat and in northern Spain. Other species with this pattern
specificity; 3) consistently low population size. Ex include Myurium hochstetteri and Herbertus borealis.
amples from the European moss flora are given in Geographically more widely ranging species can,
Table 1. by definition, meet atmost two of the three Rabinowitz
The most extreme case is exemplified by Ditrichum criteria for rarity, and then only if they are both habi
cornubicum (nomenclature for British bryophytes fol tat specific and occur in consistently small popula
lows Hill et al. 1991, 1992, 1994). This species is tions. An example isZygodon forsteri (Table 1),which
known only from three sites in Cornwall, England. It is widely distributed in Europe, in small populations
is thought to be extinct at one site and occurs in small occupying a narrow range of microhabitats on the
populations apparently restricted to copper-mine waste trunks and branches of trees, usually beech (Proctor
at the other two (Paton 1976, N.G. Hodgetts pers. 1961, Adams 1984, Hill et al. 1994). Buxbaumia viri
comm.). Other European species that, like D. cornu dis and Harpanthus flotovianus are other relatively
bicum, could be regarded as rare on all three of the widespread species that typically occur as small popu
Rabinowitz criteria include D. plumbicola, Weissia lations in a narrow range of habitats.
sterilis and Radula carringtonii. Widely ranging species may be considered rare on
Geographically and ecologically restricted species a single criterion if they either show a high degree of
that tend to be abundant where they occur are rare on habitat specificity, or occur in a range of habitats but
two of the three criteria. They are exemplified in Ta typically in small populations. Thus Schistidium

54LINDBERGIA 25:2-3 (2000)

maritimum (Table 1) and Nardia compressa are widely able habitat patches, while restricted extent of suit
distributed and sometimes abundant, but are largely able habitats could enhance problems of dispersibility.
confined to coastal rocks and to rocks and wet soil by Consistently low population size could result from
streams respectively. Conversely, Anomobryum fili small size or low carrying capacity of suitable habitat
forme (Table 1) is widely distributed in a range of patches, the latter perhaps reflecting low competitive
moist, montane habitats, but usually occurs in small ability, while high habitat specificity may result in few
quantity. This pattern is also shown by Harpanthus suitable habitat patches being available.
scutatus and Haplomitrium hookeri. Acceptance of different criteria by which rarity may
be judged is useful in emphasising that different spe
cies may be rare for different reasons, and may there
fore require different forms of conservation manage
Implications for conservation ment. Occurrence of a species as relatively large popu
Recognition that several criteria can contribute to lations but in a restricted suite of habitats is likely to
wards a definition of rarity may assist in comparing have an entirely different explanation from occurrence
vulnerability among species. It seems reasonable to in small populations in a wide range of habitats. As
argue that species such as Ditrichum cornubicum and Rabinowitz stresses, however, similar patterns of rar
Radula carringtonii that are rare on all three of the itymay also have very different explanations. A widely
Rabinowitz (1981) criteria are among the most vul disjunct distribution, e.g. Dicranodontium subporo
nerable, but are those species rare on the basis of two dictyon, may often be relictual, but a narrowly en
criteria generally more at risk than those that meet demic range could reflect either a species that is on
only one? the verge of becoming extinct, or one that has newly
It must be recognised, however, that while these evolved.
three criteria are simple in concept the limits are im This raises a philosophical question. Millions of spe
possible to define objectively, even if distributions cies, and countless higher groups of organisms, have
were perfectly documented which usually they are not. become extinct throughout biological time. Extinc
Ditrichum cornubicum is now known from only two tion is a natural process: it is an inevitable consequence
sites a few kilometres apart in Cornwall. There can be of continuing, sometime cataclysmic environmental
no argument that it has a narrow geographical range, change, and of organic evolution which inexorably
at least as currently recorded. But where does one draw replaces older taxa with newly evolving ones that
the line? Should narrow geographical range be taken prove competitive in the environments within which
as
meaning endemic to, say, Cornwall, to Great Brit they arise. Should we attempt to restrict natural ex
ain, to western or to western Eurasia? How tinction, or should we concentrate our efforts on lim
Europe,
do we treat disjunct ranges? Would there be universal iting extinction resulting from human activities? To
agreement that Dicranodontium subporodictyon what extent can we distinguish between the two?
should be as rare on the basis of narrow geo
regarded
graphical range occurring, as it does, in a few locali
ties in western Scotland, in Yunnan, China, and in
western Canada?
Similar problems arise in attempting to define habi ofMossSpeciesinDifferent
Numbers Numbers
ofSquares
of theBritish
National
Grid
tat specificity and population size. Myurium hochstet
teri occurs on a variety of substrata, but should it be
regarded as showing high habitat specificity because
it is predominantly coastal? Moreover, the three cri
teria may to a degree be inter-related. Restricted geo
graphical range could be a result of inefficiency in
dispersal. However, it could also be indicative of a
narrow of climatic tolerance, and thus repre
degree
sent a form of habitat specificity, as may well be the
case with Dicranodontium subporodictyon and other
"Atlantic" species.
Itwill be evident that the classic parameters of meta
Number
ofSquares
population dynamics (S?derstr?m and Herben 1997) 1. Distribution of mosses in Great Britain
will have different, possibly interacting relationships Fig. expressed
as number of species recorded in different numbers of 10
with these three spatial aspects of rarity. Thus narrow km squares of theNational Grid since 1950 during the Brit
geographical range could result from poor ish Bryological Society Mapping Project. Data from Hill
dispersibility, and/or the restricted occurrence of suit etal. 1991, 1992, 1994.

LINDBERGIA 25:2-3 (2000) 55

How many rare species?
401
It is another fundamental fact of biology that, quite Dioecious Rare

apart from the effects of man, there are far more rare
than common species, if rarity is defined on the basis 20H
of distribution relating to any of the accepted criteria.
There are likely to be few abundant species compared
with rare ones within any community. Most species C F O R VR UB UA
80
occupy a narrow range of habitats, with rela
relatively
Dioecious Common
tively few displaying little habitat preference. Most
species have a narrow range, or are sparsely distrib 60 H
uted, within a given area or globally.
This truism is commonly expressed in the form of
40
"hollow curve" distributions such as that in Fig. 1,
which shows the distribution of moss species within
10-km grid squares inGreat Britain, as recorded since 2CH
1950 during the British Bryological Society Mapping
Project (Hill et al 1991,1992,1994). While a few spe
cies are widely distributed throughout many of the C F 0 R VR UB UA
more than 2400 grid squares, 50% of some 700 spe 40
cies were recorded in fewer than 100 squares. Indeed Monoecious Rare
40% were recorded in fewer than 50 squares, and 25% 20
in fewer than 15. Similar examples are presented in
e.g. Urmi and Schnyder 2000). ~L
O
Given that so many are rare, how does one F
species C O R VR UB UA
select those most meriting conservation? In this con
text it is important to distinguish between global and 100 Monoecious Common
local rarity, as a species may be widely distributed
and perhaps abundant in some areas, but of extremely
local occurrence elsewhere, often towards the edge of 80
its range. Many of the species with a highly restricted
distribution within Great Britain are widespread in 60
other parts of the world. Examples include Arctic
montane such as Timmia austr?aca and Medi
species 40
terranean species like Pottia commutata. Such locally
rare species are not considered in the scheme in Table
1. 20
Is it sensible that amajor conservation effort be ex
pended on species that are rare only within a particu
lar, perhaps politically defined, region, if this means C F O R VR UB UA
diverting resources away from globally threatened FREQUENCY OF SPOROPHYTES
species? Or should one argue that locally rare species Fig. 2. Contrasting relationships between number of spe
contribute to local diversity and must be respected if cies and frequency of sporophytes among mosses that in
we are to harness local conservation effort to maxi Britain are dioecious and rare, dioecious and common,
mum overall benefit? Moreover, theoretical consid monoecious and rare or monoecious and common. Fre
of sporophytes decreases from left to right: C, com
erations suggest that peripheral populations of a spe quency
mon; F. frequent; O, occasional; R, rare; VR, very rare;
cies may differ genetically from those in the centre of
UB, unknown in Great Britain; UA, unknown anywhere.
the range (Furlow and Armijo-Prewitt 1995, Lesica from Longton
Reproduced (1992) with permission from
and Allendorf 1995). If so, may not such populations the Editor of Biological Conservation.
merit conservation in their own right, possibly as in
cipient new species evolving in reproductive isola
tion or as populations in which could evolve geno
types tolerant of global environmental change (Safriel Population biology
etal. 1994)? Reproduction and rarity
It is widely accepted that potentially important rela
tionships exist between reproductive biology and rar
ity, particularly in bryophytes where some species

56LINDBERGIA 25:2-3 (2000)

 seldom, if ever, produce spores. Growth and branch esis that lack of genetic recombination is important in
ing of existing gametophytes may be sufficient to pro leading some species to become rare by limiting ge
vide for colony maintenance and expansion in many netic variability and therefore capacity for adaptation,
perennials. However, production of some form of that monoecious species thus tend to become rare if
-
diaspore spores, specialised asexual propagules,
or self-fertilization becomes essentially obligate, and that
- is essential for the estab dioecious species tend to become rare if they fail to
gametophyte fragments
lishment of new colonies in existing populations, and produce sporophytes possibly through limitations on
for the establishment of new populations enabling a both variability and dispersibility. A second hypoth
species to expand its range or simply to replace popu esis is that species become rare for reasons unrelated
lations that become extinct. to the frequency of meiosis. Increasing rarity would
Moreover, spore production following sexual repro not affect fertility in monoecious species, but might
duction and meiosis could be vital inmaintaining and decrease the frequency of sporophyte production in
enhancing variation among genotypes and thus facili dioecious species by reducing the frequency with
tating adaptation to a range of, possibly changing, which male and female plants become established side
environments, i.e. to reducing the chance of rarity aris by side.
ing through high habitat specificity. Conversely, spe The occurrence of substantial numbers of common
cies reproducing principally asexually may be abun but rarely, or non-fruiting dioecious species (Fig. 2)
dant in terms of cover or biomass, but rare in respect tends to support the first hypothesis, by which failure
of the number of genetic individuals present. This to produce spores precedes rarity rather than being a
pattern has been documented in Pteridium aquilinum consequence of rarity. It will be noted that the first
(Harper 1981), and a similar situation may exist among hypothesis involves the assumption that, among mo
bryophytes, for example in liverworts of formerly gla noecious mosses, self-fertilisation will be more preva
ciated areas (Wyatt 1994). lent in rare than in common species, whereas no such
At the first European Conference on the Conserva assumption is implicit in the second hypothesis. Com
tion of Bryophytes, a comparison was presented of paring the incidence of self-fertilization in rare as
certain features of reproductive biology in rare and opposed to common monoecious bryophytes would
common British mosses (Longton 1992), rare species thus be of great interest. Itwould also be of interest to
being defined on the objective, but arbitrary basis of establish how far the relationships between reproduc
records in 15 or fewer 10-km2 grid squares since 1950. tion and rarity evident in Fig. 2 apply to the different
Itwas shown that a significantly higher proportion of patterns of rarity summarised in Table 1, as a contri
monoecious species is rare than of dioecious species. bution towards understanding these various forms of
It was also shown that a significantly higher propor rarity.
tion of species not known to produce sporophytes in
Britain is rare than of fruiting species, but this rela
tionship clearly differs between monoecious and
dioecious taxa (Fig. 2). Among dioecious mosses, most Table 2. Distribution of British mosses among life-history
strategies 1992). Figures are percentages of spe
species with sporophytes unknown, either in Britain
(During
cies in each strategy. Only two British mosses are regarded
or throughout their range, are rare in Britain, while
as fugitives: neither is rare. Species with un
sporophytes
most species known to produce sporophytes in Brit known are omitted. Data from Longton *
(1992,1997). Sig
ain, even are more common. monoe different from value for all species =
rarely, Among nificantly (%2 9.57; p
cious mosses, most sporo < 0.01). No other values for rare species are significantly
species normally produce
phytes, and a substantial proportion of freely fruiting different from values for all species (p >0.05).
monoecious species is rare.
All species Species rare
The 177 species considered rare in this analysis in
inGreat inGreat Globally
cluded some that are globally rare, and others that are Britain Britain rare species
rare in Britain but common elsewhere. However, the

relationships evident in Fig. 2 were shown even more Perennial

strikingly in a preliminary list of 76 British mosses stayer 25 14 20

considered to be rare at a world level. Thus most rare Colonist 51 60 55
Dominant 3 05
mosses in Great Britain are either 1) dioecious and
Long-lived
fail to produce sporophytes in Britain, or 2) monoe shuttle 13 10 14
cious and produce sporophytes freely. More recently Annual or
we have demonstrated similar relationships between short-lived

reproduction and rarity among British liverworts shuttle 7 18* 8

Total number
(Laaka-Lindberg et al. 2000).
These relationships are consistent with the hypoth of species 680 152
51

LINDBERGIA 25:2-3 (2000) 57

 sequences such as dubious to
taxonomically attempts
Rarity and life-history strategies maintain locally variant populations at specific rank,
Several authors have commented that annual species, rather than include them within more widespread spe
and other taxa associated with disturbed habitats, are cies, in order to increase the justification for preserv
particularly susceptible to becoming rare or endan ing those particular populations. This is an example
gered. Watkinson (1981) noted that weeds of arable of "taxonomy as destiny" discussed by May (1990).
land are among themost severely threatened vascular So, is there any particular property of the category
plants in the British flora. Orb?n (1992) analysed the species that justifies our viewing species as the basic
Hungarian bryoflora in terms of During's (1979) life unit for conservation, even at the organismal level?

history strategies. Colonists were the largest group, Theoretical considerations lead to the prediction that
comprising 44% of all species, with this proportion rare species may tend to display a narrow range of
rising to 60-80% among various classes of endangered genetic diversity due to factors such as genetic drift
species. Hodgetts (1996) showed for British bryo and inbreeding among small groups of individuals.
phytes that among threatened species most perennial Electrophoretic studies reviewed by Wyatt (1992,
stayers are vulnerable, with relatively few endangered 1994) have shown that moss species may contain as
or critically endangered, while the reverse is true for much internal genetic variation as species of diploid
short-lived shuttle species. organisms. The extent of intra-specific variation dif
The distribution of life-history strategies sensu Dur fers between species, however, and a careful study of
ing among British mosses is shown in Table 2. Spe species inPlagiomnium section Rosulata showed that,
cies not known to produce sporophytes are omitted as predicted, values of percentage loci polymorphic,
due to the difficulty in assigning such species to strat mean number of al?eles per locus and mean expected
egies in the absence of data on spore size. Colonists, heterozygosity were substantially greater in widely
at 51%, form the largest group, as in the Hungarian distributed species such as P. ellipticum than in
bryoflora, and the proportion of colonists rises slightly endemics like the northern European P. elatum. Wyatt
but not significantly among species considered rare (1992) discussed the implications of such findings for
in Great Britain or globally. The only significant dif conservation management.
ference here is the substantially higher percentage of If we accept the view that conservation should aim
annual or short-lived shuttle species among the mosses tomaximise the preservation of information contained
considered globally rare (18%) than in the British flora in the DNA of living organisms (Wilson 1992, Cro
as a whole (7%), a result that could be influenced by zier 1997), then species would have high conserva
the ephemeral nature of shuttle species creating a false tion value, even if within-species variation is low, if
impression of rarity. they contain unique genetic information. In other
The high proportions of colonists and shuttle spe words, the worth of a species is dependent on its de
cies among rare bryophytes have implications for con gree of phylogenetic distinctiveness. However, the
servation whether or not these propor same argument to populations, and
management, applies equally
tions are higher than in the flora as a whole. Harper again raises the question of where conservation effort

(1981) has noted that such species are likely to show is to be expended.
variation with time in their degree of rarity, and Dur In Fig. 3 we present a preliminary unrooted phyl
ing (1992) has pointed out that some may be rare above ogeny for various members of the Bryaceae (as rede
ground but common in the soil propagule bank. A fined by Cox 1998), based on maximum-likelihood
continuous availability of successional habitats will analysis of nucleotide sequence variation in the inter
clearly be required in reserves designed to accommo nal transcribed spacer (ITS) region of the nuclear-en
date such species, with climax vegetation perhaps coded rRNA cistron. The data set includes individual
being favourable for some types of rare species, e.g. isolates of 15 species, including some very rare taxa
epixylic taxa, but not for others such as soil-inhabit (e.g. Mielichhoferia macrocarpa: see Brassard and
ing colonist and shuttle species. Hedderson 1983) and 18 geographically diverse iso
lates of the weedy and cosmopolitan Bryum argen
teum. Topologies in some regions of the tree are rather
poorly supported (information not shown), but for the
present discussion we are interested primarily in
What to conserve? Species or branch lengths which are not strongly affected by topo
populations? logical change. Branch lengths in the tree are propor
Intra-specific variation tional to the number of nucleotide substitution events
It is commonly accepted, even among biologists, that which occur along them, and thus provide ameasure
extinction of species should be avoided wherever rea of "distinctiveness" for each observation in the data
sonably possible. This sometimes has unfortunate con set.

58 LINDBERGIA 25:2-3 (2000)

 /Galapagos from the Churchill population, although field-grown
Galapagos 2
plants at Churchill had a long apiculus, and the three
populations differ in other morphological features
(Longton 1981). Extensive RAPD diversity has re
Churchill cently been reported within and between populations
Ellesmere of B. argenteum from Australia, New Zealand and
Ross Island 1
Ross Island2 Antarctica (Skotnicki et al. 1998).
Assuming that the DNA region analysed here pro
vides at least a vague reflection of overall genomic
differences, the phylogeny has clear implications for
Bryum funkii +B caespiticium etc (7 spp.)
Mielichhoferia macrocarpa conserving genetic information in the Bryaceae. Per
Bryum alpinum haps the most important finding is that individual
Bryum cellulare of B. have a conserva
populations argenteum higher
Bryummuehlenbeckii
tion "worth" than do several species. For example,
Anomobryum julaceum
the rareMielichhoferia macrocarpa is virtually indis
tinguishable from a set of species including several
common and taxa caespiti
widespread (e.g. Bryum
Bryum blandum cium), and two geographically isolates of M.
disjunct
macrocarpa proved uniform in the present analysis.
These results, if confirmed for additional populations,
suggest that loss ofM. macrocarpa, lamentable though
such an event would be, would not result in any ma
Bryum wrightii
jor loss of genetic information. Thus, however con
3. Preliminary, unrooted maximum-liklihood troversial itmay appear, it could be argued that in a
Fig. topology
for Internal Transcribed Spacer (ITS) regions of the nu world of limited conservation resources the species
clear-encoded rRNA Cistron of Bryaceae (Hedderson et M. macrocarpa should take lower priority than some
al. unpubl.). Species names refer to one, or two or three sensu
populations of B. argenteum lato.
uniform isolates from a range of taxa. Geographical names
The studies to date thus suggest that there may be
refer to isolates of Bryum argenteum. Branch lengths are
to the number events
more genetic information within a single common and
proportional of functional substitution
which occur them. variable species such as Bryum argenteum or Plagi
along
omnium ellipticum than there is within several rarer,
more such as P. elatum or M.
st?notypie species
macrocarpa. Furthermore, individual populations of
A notable feature of the tree is thatmuch of the to some
widespread taxa may harbour more
unique ge
tal variation resides within Bryum argenteum. Dis netic information than some rare and cherished spe
tances between populations of this species are much cies. Indeed, consider the effects of widespread ex
greater than between some species traditionally as tinction of populations in a common, variable species.
signed to different genera. It could be argued that some One result might be the creation of discontinuities in
of this apparent diversity arises from inadequate spe the pattern of variation such that several less common
cies delimitation, and that B. argenteum actually rep and less variable species may legitimately be recog
resents a series of sibling species. Indeed there may nised among the remaining populations! This scenario
be justification for recognising the clade comprising offers one realistic explanation for the origin of rela
the Churchill (Sub-Arctic), Ellesmere Island (Arctic) tively uniform species, but are such species of greater
and Ross Island (Antarctic) populations at the species conservation value, even if rare, than those popula
level as this group, so far, emerges as a tions of the more common, whose
monophyletic original, species
group sister to rest of the isolates. However, such an extinction led to the new species meriting specific
argument could not be used for the equally distinct rank?
Galapagos Islands populations which are clearly
nested with a series of less divergent isolates. The Ross
Island populations have variously been regarded as
conserve
representing B. argenteum (e.g. Greene 1967) or a dis Why biodiversity?
tinct species (e.g. Seppelt and Green 1998). The Given that common species may contain a
greater in
Ellesmere Island and Ross Island populations resem formation content than rare species, is a preoccupa
ble each other in showing an unusually short leaf tion with conserving rare species justified? Is it rea
apiculus, in the field and in cultivation. This feature sonable to expend resources in seeking to conserve
was also exhibited by a clone isolated from a spore the rarest species thatmay already have fallen below

LINDBERGIA 25:2-3 (2000) 59

aminimum viable population size (Soul? 1987) nec suring that a reasonable number of species
are not

essary for their long-term survival? Should we not forced into extinction. Is it not really our responsibil
concentrate on the infinitely more challenging task of ity to disrupt the lives of other organisms, particu
limiting the loss of individual populations, "an insidi larly perhaps sentient animals, as little as we reason
ous form of biodiversity erosion" (Ehrlich 1995), that ably can?
will "tend to reduce the taxonomic, genetic and func Yes, of course people derive satisfaction from bio
tional diversity of sites, and perhaps the performance - from
diversity observing, studying, being stimulated
of ecosystems" (Gaston and Spicer 1998)? by or even simply by being aware of an infinite range
The answers to some of these questions will inevi of other organisms. Distinctive species contribute
tably be conditioned by our reasons for seeking to con greatly to our appreciation of nature, but would not
serve
biodiversity,
reasons that are commonly con that appreciation be sadly diminished if the rare spe
sidered under four headings. First, it can be argued cies were confined to reserves, zoos and test-tubes,
that there are moral considerations impelling us to and we seldom actually encountered any but those few,
wards conservation, that we are simply not justified common plants and animals that are best adapted to
in making some species extinct in order to improve urban and intensive agricultural landscapes?
the quality of human life. Second, there are aesthetic Yes, of course we should seek to preserve as much
reasons: biodiversity is unquestionably one of factors as possible of the variation in genetic information
that enhances the richness of the human experience. content present within the biotic community because
Many people derive immense pleasure from observ of its potential for yielding compounds of value in
ing birds and large mammals, rather fewer, perhaps, agriculture, medicine and other vital human activi
from flowers, and some even from mosses and liver ties. But where does that variation reside? Itmay re
worts. Third, there are
practical and economic rea side as much in populations of common and variable
sons, since loss of genetic information means loss of species as in rare species, as Fig. 3 so clearly indi
information about compounds potentially beneficial cates.
to man and therefore marketable. Fourth, there is con Yes, of course we should seek to preserve biotic di
cern that ecosystem stability may be threatened if too versity because of its potential role inmaintaining the
many species become extinct too quickly. effective functioning and stability of the global eco
These issues, and their implications for conserva system. That role may not yet have been established
tion management, have recently been discussed by beyond doubt, but does not the precautionary princi
Crozier (1997). He notes that economic considerations, ple demand that we jealously guard our remaining
by which we may best hope to sell the need for con store of biodiversity, at the community as well as the
servation politically, and aesthetic considerations that species and population levels, as long as we suspect
may be paramount for many people, would both rate that such diversity may be essential to maintain the
distinctiveness as a criterion in assessing con ecosystem upon which we, as animals are absolutely
major
servation value. Bisang and Heden?s (2000) describe dependent. The problems we are about to face through
a phylogenetic approach to identifying the most dis climatic change would have been infinitely less se
tinctive species in terms of genetic information con vere if the world community had adopted the precau
tent, thus providing a rational basis for selecting pri tionary principle when scientists first began to warn
ority species for conservation. Other contributors have of the probable effects of greenhouse gas emissions
stressed the importance of identifying which ecosys as long ago as the 1960s.
tems, habitats or sites should be safeguarded in order The notion of preserving distinctive species because
to preserve in situ as many as possible of any list of of the potential value of their genetic information con
such priority species (Sergio et al. 2000), while tent in yielding marketable compounds may be the
Jackson et al. (unpubl.) describe a developing strat conservation message most readily accepted by our
egy for conserving bryophyte species ex situ. all too economically orientated political leaders. As
We contend, however, that this priority species ap biologists, however, we surely realise that preserving
proach, while vitally important as part of a twin-track a list of priority species will not, in itself, achieve any
strategy, does not effectively address any of the four of themajor conservation objectives, and that we shall
broad reasons for seeking to preserve biodiversity. We be abrogating our responsibility to the world commu
consider all four reasons valid, and from this it fol nity if we pretend that it will. All of us who under
lows that we agree with Ehrlich (1995) that we must stand the broader issues have a clear responsibility to
also seek to limit the ongoing loss of biotic popula challenge, forcibly and incessantly, the wisdom of
tions consequent upon the present global wave of habi continuing global habitat destruction, and the twenti
tat destruction. eth century obsession with economic growth for
-
Yes, we do have a moral obligation towards other growth's sake rather than economic development
-
organisms, but this surely extends beyond merely en geared to the real needs of people that is making

60LINDBERGIA 25:2-3 (2000)

 that destruction as inevitable as it may eventually Longton, R. E. 1981. Interpopulation variation in morphol
ogy and
physiology in the cosmopolitan moss Bryum
prove disastrous.
-
argenteum Hedw. J. Bryol. 11: 501-520.
- -
- 1992. Reproduction and rarity in British mosses. Biol.
Acknowledgements We are grateful to the Natural Envi
Conserv. 59: 89-98.
ronment Research Council (UK) for
financial support to -
1997. Reproductive biology and life-history strategies.
TAH through the Taxonomy Initiative funding awarded to -
Adv. Bryol. 6: 65-101.
The Univ. of Reading. We also thank Emma Pharo for help -
May, R. M. 1990. Taxonomy as destiny. Nature 347: 129
ful comments on the manuscript.
130.
Orb?n, S. 1992. Life strategies in endangered bryophytes
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in Hungary. Biol. Conserv. 59: 109-112.
Paton, J. A. 1976. Ditrichum cornubicum, a new moss from
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Cornwall. J. Bryol. 9:171-175.
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