Biodiversity and its relationship to ecosystem health and

integrity1
by J.P. (Hamish)~ i m m i n s ~

Introduction biological diversity and var-

The human population has increased from about 1 billion ious concepts of ecosystem
to its present 5.8 billion in the time taken to grow a mature for- "health" and "integrity".
est stand in most of Canada. Over the period required to
grow another mature stand, the population is expected to Origins of Biological
grow to between 12 and 16 billion. With the present per Diversity
capita wood consumption and the present rate of population The term biological diver-
growth, the annual increase in demand for wood is greater than sity refers to the variety of the
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the present allowable annual harvest in British Columbia. living components of ecosys-
Although both the area of forest and/or the volume of wood tems. This encompasses
are increasing in some countries (e.g., Scandinavia,Canada), many levels of variation,
the global area of wood-producing forest is declining, and the from the genetic variation
pressure on the remaining forest continues to increase. within a local population of
The increasing demands on the world's forests have raised pine trees or deer, for exam-
serious concerns about loss of biological diversity. This con- ple, to the variety - of species
-
cem was originally focused on the biodiversity cdnsequences within a biotic community, and the variety of biotic commu-
of logging in species-rich,primary, humid, tropical forest. In nities across the landscape.
this type of forest, it is expected that timber Biological diversity has a variety of
harvesting will result in the loss of some plant origins which must be understood if particular
and animal species from the affected area. There is no desired levels of diversity are to be achieved
Because many ~ o p i c aspecies
l have rather of health that and sustained in managed forests.
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restricted geographical ranges, such local loss

of individuals of a species can result in the applies to all I. Ecological Diversity
loss of the entire species if adequate unhar- One of the major determinants of diver-
vested forest reserves are not established. In stages of ecosystem sity across a forested landscape is the vari-
recent years, this concern has been re- ety of climatic and soil conditions. Variations
focused on temperate and boreal forests; in slope, aspect, elevation, latitude, dis-
ecosystems in which well-regulated forest tance to the oceans or large lake, local and
harvesting does not threaten the loss of plant species, and in regional topography as it influences climate, geology and soil
which most other species have sufficiently broad ranges that conditions create the environmental framework for biologi-
species extinction is not a threat. cal diversity. Differences in plant species adaptations result
The greatest threat to biological diversity results from per- in different plant communities along climatic and soil gradi-
manent deforestation: not sustainable forest management ents, and the variations in both plant communities and phys-
and forest harvesting. Land use change from forest cover to ical environmental factors result in different animal and
agriculture, urban area, water storage, transportation corridors microbial communities. These in turn affect the diversity of
and some types of recreation is generally the major cause, but the plant communities.
repeated burning, grazing andlor firewood cutting by local peo- This physical-chemical environmental framework sets the
ple following timber harvesting is the major loss in some devel- continental, regional and landscape potential for biological diver-
oping countries. However, forest management does have the sity. Within this environmental framework, which is largely
potential to change biological diversity, and it is important that unaffected by forest management, the actual biodiversity
this potential impact be considered for each type of forest. The that one can observe is the result of evolution, population and
implications of any changes for "forest health" and "ecosys- community processes, and processes of natural and human-
tern integrity7'also needs to be evaluated. These are fundamental caused disturbance.
issues for forest certification, and must be understood and eval-
uated on a site-specific and landscape-specificbasis if certi- 2. Evolution
fication is to succeed as a mechanism for promoting sustain- Species evolve as a result of physical factors and biolog-
able forest management. ical processes inter-acting with genetic variation through the
In this paper, I review the origins of biological diversity, the process of natural selection. This leads to progressive spe-
various measures used to describe it, and our current under- cialization of the ecological roles of species Over time,
standing of the relationship between various measures of which allows new species to develop by dividing up the ''eco-
- -
logical iobs" between a larger number of species. Generally
<

speaking, all other things being equal, the longer evolution

'Reprinted from Policy Options, November 1996.
2Professor, Department of Forest Sciences, Faculty of Forestry, University has continued without interruption (e.g., by events such as
of British Columbia, Vancouver, BC ice ages or major climatic change), the greater the degree of

MARCWAPRIL 1997, VOL. 73, NO. 2, THE FORESTRY CHRONICLE 229

 ecological specialization, and the greater the number of To this can be added climbing plants and epiphytes. In addi-
species. tion to the structure of the living plant community, forests vary
in the number, size and condition of standing dead trees
3. Population and Community Processes (snags)and of decaying logs on the ground (coarse woody debris
The ecological processes that determine the abundance of or CWD). Some forests have a relatively unchanging vertical
any one species, and the community processes that determine structure as one walks through a stand; others are rather
species interactions and the response of communities to biot- patchy, with small gaps caused by the death of individual large
ic and physical disturbances, all play an important role in deter- canopy trees or small groups of trees, interspersed through a
mining biological diversity at the local stand-level. Because matrix of closed-canopy forest. Structural diversity can be as
of the change in these processes over time as ecosystems recov- important for animal species diversity as is the diversity of plant
er following natural or human-caused disturbances, there is a species in the forest plant community.
continued change in biological diversity in a forest stand
over time. 4. Functional Diversity
Management of forests to achieve specific biodiversity Forests vary in the functional characteristics of their tree and
objectives requires that we understand the overriding influence other plant species. This aspect of biodiversity encompasses
of ecological diversity, the history of evolution, the ecolog- evergreen vs. deciduous, broadleaved vs. needle-leaved,
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ical role of disturbance, and the ecological process of ecosys- shade tolerant vs. light demanding, deeprooted vs. shallow-
tem recovery from disturbance in the forest area in question. rooted, nutrient demanding vs. non-nutrient demanding, etc.
Because these all vary fiom one forest area to another, the issue
of biodiversity and the effect of forest management thereon Spatial and Temporal Scales of Biological
must be region-specific, landscape-specific and site-specific. Diversity
The measures of biological diversity defined above can be
Measures of Biological Diversity applied at various different spatial scales, and all of them vary
Few people would disagree with the premise that we should over time as ecological succession causes ecosystem change
manage to conserve and sustain desired levels of "biodiver- following ecosystem disturbance.
sity". However, it is not sufficient simply to say that foresters
should "manage for biodiversity". Because there are so many 1. Alpha Diversity
measures of biodiversity, such a broad, general statement of Measures of biological diversity at the local ecosystem or
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intent provides no basis for deciding how a particular forest stand level. Much of the debate about biodiversity has been
should be managed. Specific, desired levels of different mea- about alpha species richness diversity, a focus that is gener-
sures of biodiversity, and how these should vary over time, should ally less important for species conservation than landscape mea-
be identified. Only then can we decide which forest practices sures of diversity. Alpha species richness may sometimes be
are consistent with particular biodiversity objectives and less important in conservation than alpha species eveness, struc-
which are not. tural or functional diversity.
1. Genetic Diversity 2. Beta Diversity
The variation in the genetic make-up of organisms. This usu- Variation in the various measures of alpha diversity as
ally refers to the genetic variation within a population or a species, one moves across local landscapes. Beta diversity reflects both
but it can also refer to the variety of species, genera, families local soil and topographic diversity, and variations in forest
or other taxonomic groupings. These different taxonomic age and sera1 stage that have resulted from variations in dis-
levels reflect a gradient from the most genetically similar to turbance regimes and disturbance history.
the least genetically similar groupings of organisms.
3. Gamma Diversity
2. Species diversity Variation in alpha and beta measures of diversity as one moves
The number of species in a particular area or community, across regional landscapes and along major environmental gra-
or the relative abundance of the species therein. The number dients (e.g., latitude, altitude and other climatic grad~ents).Gamma
of species is measured as species richness;this is the most com- diversity is largely a reflection of ecological diversity.
monly used measure of biological diversity. The relative
abundance of the species is referred to as species eveness.For 4. Temporal Diversity
a given number of species, a community will have hi&mqwies All other measures of diversity change over time. Sometimes
diversity if all species are represented by the same number of this change is quite rapid as a result of disturbance and eco-
individuals,than if W ?of all individualsare h m a s q l e species, logical succession; sometimes the change is slow as a result
and the other species account for only 10% of the individu- of climate change, evolution, or changes in species distribu-
als. tions. Temporal diversity is seen most at the alpha or local stand
scale, and least at the gamma or regional scale. Beta tempo-
3. Structural Diversity ral diversity is usually intermediate.
Variation in how plant communities are structured, both hor- Much of the focus in discussions about biodiversity has been
izontally and vertically. Some forests have a single tree at the alpha or local ecosystem level. However, thislevel exhibits
canopy layer, no herbs and shrubs, and just a moss layer on the greatest temporal diversity. Sustaining natural patterns of
the ground. In contrast, some forests have several tree canopy change in other measures of alpha diversity to achieve natural
layers, one or more shrub layers, a herb layer and a moss layer. alpha temporal biodiversity means that one cannot sustain any

230 MARSIAVRIL 1997, VOL. 73, NO. 2, THE FORESTRY CHRONICLE

 of these other measures at a fxed constant level. Conversely, humans varies from babies, to teenagers, to adults and to old
sustaining any other alpha measure at a constant level means people. There is no "norm" of health that applies to all stages
that one cannot have natural temporal diversity for that mea- of human health. Similarly,there is no "norm" of health that
sure. applies to all stages of ecosystem development. A healthy human
Forest landscapes generally represent a mosaic of different population might be defined as one that has a "normal" bal-
forest ages and seral stages. Each subcomponent (each stand ance of all age classes; not all young and not all old. A
or local ecosystem) of this mosaic changes with time, but the healthy forest landscape can similarly be defined as one that
overall beta diversity may remain relatively constant, except has all the different ecosystem conditions represented in it: not
in landscapes that are periodically affected by large-scale, stand- all "young" and not all "old".
replacing disturbance, such as fire in boreal forests. In such Ecosystem "health" is clearly different when considered at
forests, beta diversity may change considerably over time. the landscape and stand levels. Both are important in conservation
Particular levels of various measures of beta diversity can be and forest management.
sustained at rather constant levels (i.e. low beta temooral
diversity) even though V ~ ~ ~ Oalpha
U S IkaSUres of diversik are Relationship bemeen Biodiversity and
changing for each of the components of the landscape mosa- Ecosystem Health and Integrity
ic (i.e. high alpha temporal diversity).
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Most of the measures of biolo~icaldiversity differ between

different stages of ecosystem development. ~ i h l a r ldifferent
~,
Ecosystem Health and Integrity sera1 stages are associated with different "health" conditions.
Having defined the diversity of measures of biological seral stag- exhibit ecosystem integrity. Scientists have smg-
diversity, 1will consider eCOSyStem "health" and "integrity", gled for some time to establish reliable relationships between
and their relationship to biological diversity. these different ecological concepts, but the diversity of nature
An ecosystem is any biological-~h~sical system that exhibits has defeated this human quest to generalize and order nature.
the athibutes of structm, hction, complexity, No clear scientific relationships between
interactions/interco~ectionof the sub- diversity and ecosystem health, &versity and
components, and change over time. By def- Disrespect for ecovstem inte@ty, diversity and ecosys-
inition, a system has integrity. Therefore, any tem productivity, and between diversity
system that is an ecosystem also has integri- nature means and ecosystem stabilityhave been identified
ty. To suggest that an ecosystem lacks applying the same The conclusion has been reached by eco-
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integrity is to require a change in the mean- logical scientists that the quest for such
ing of these words. TO say that an ecosys- approach to relationships is fruitless; that while rela-
tem has lost its integrity implies that it has tionships may exist, their validity will be lirn-
lost the attributes of the system. forests that vary ited to particular forest types and sera1
@& a mcan in a in their ecological stages.
ety of conditions. These different condi- Unmanaged forests exist in a wide range
tions are called sera1stages: different stages characteristics of biodiversity conditions. There are many
in the development or change of an ecosys- "natural"forests that have low levels of many
tem over time from some condition-altering measures of biological diversity, but are pro-
disturbance back towards some original condition, or to a rel- ductive, healthy and have ecosystem integrity (e.g., pine
atively stable, self-replacing end state or condition (the seral forests, eucalypt forests). Similarly, one can find forests with
climax). Each individual condition or seral stage will have its low values of several measures of diversity that have low pro-
own integrity, which is lost as natural processes or disturbance ductivity and stability (e.g., climax temperate rain forest in coastal
replace that condition or seral stage with a new condition or BC). Conversely, there are forests that are stable, productive
seral stage, which itself has integrity. Thus, the integrity of the and moderately diverse (e.g., eastern temperate hardwood forests
forest as a ecosystem remains undiminished as the ecosystem in the US). And, of course, there are forests with high diver-
develops throughvarious conditions or seral stages, but the integri- sity but low productivity and low stability (e.g., tropical rain
ty of each stage is transitory. There is only a loss of ecosys- forests growing on nutrient-poor tropical white sand soils). All
tem integrity if the ecosystem processes are altered to beyond combinations of measures of diversity, stability and ecosys-
the range that is characteristic for one of the seral stages of that tem health can be observed in nature.
ecosystem. A major problem with measurement of ecosystem health is
This can occur followingvery severe ecosystem disturbance. that it is not exclusively a scientific concept. It is defined as
It persists only until the processes of ecological succession have much by the social and economic values people want from their
re-established an early sera1 stage. forests as it is by a scientific assessment of forest health. This
Ecosystem health is a metaphor borrowed from human results in conflicting definitions of what people think a
health. With humans, it is easier to define the "health" of indi- healthy forest is, which in turn renders the job of establishing
vidual people than of populations or entire human societies. relationships between measures of biodiversity and mea-
With forests, it is easier to define the health of individual trees sures of ecosystem health even more difficult.
or animals than of populations of those species, and easier to Because of the complex mix of science and social values
define the health of populations than of biological communities, in defining these ecosystem attributes, a lot of mythology has
local ecosystems or landscapes. developed about them. These have failed under carell scientific
Each stage of forest ecosystem development is character- scrutiny yet have persisted in the public debate about forestry.
ized by specific "health" conditions, just as the health of In his book Discordant Harmonies, Daniel Botkin warned that

MARCHJAPRIL 1997, VOL. 73, NO. 2 , THE FORESTRY CHRONICLE 231

 attempts at conservation and sustainable resource management acter of the forest. Similarly,management of naturally ~ a n n c s
have consistently failed when based on paradigms founded on driven forests may appear to be destructive and "disrespect
mythology about nature rather than scientifically-vigorousobser- nature". However, it is simply respecting nature in the sense
vation and analysis of the form and fimctions of nature, and of respecting the natural role of disturbance in those forests.
how this varies from place to place and time to time. Only when Respect for nature in the context of biodiversity and ecosys-
nature itself is the paradigm for nature will we be successful tem health means understanding the ecology of particular val-
in achieving our biodiversity and resource sustainability ues in particular local ecosystems or landscapes. It means under-
objectives. standing how the ecology of values varies from place to
Much has been said in the environmental debate about place and time to time.
"Respect for Nature". However, there is frequent confusion Disrespect for nature means applying a single approach to
about what this term means, and how we should conduct forest management and forest harvesting over forests that vary
forestry in order to respect nature. This is because there are in their ecological characteristics. It means basing our man-
two major different meanings of thisword. One is to have "due agement on a paradigm of nature that is inconsistent with the
regard for" or "to take particular notice" of the object of character of nature. These errors have been common in
respect. Another is to "esteem" or "venerate" the object. forestry in the past. It cannot be allowed to continue. Equally,
Many of the mythologies about ecosystem diversity, health and groups in society who are trying to replace this forestry error
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integrity are based on the latter definition. Successll sustainable by their own error of applying their own disrespect for nature
resource management must be based on the former. cannot be permitted.
Respect for nature in forest management requires that we
understand the functioning of the forest ecosystem, the eco- Conclusions
logical role of disturbance, and the processes of ecosystem recov- Biodiversity is part of our biological inheritance. We have
ery from disturbance. It means that as we alter natural process- the obligation to pass it on to future generations as unimpared
es through management, we must replace the effects of those as possible. However, we will fail to do so unless we respect
natural processes that we have altered by the ecological nature by "taking due notice" of what it is and how it varies.
effects of management practices. In those forests where nat- If we do not understand nature, we are unlikely to sustain it.
ural disturbance that is typical for the desired sera1stage is char- Similarly, we cannot use the concepts of biological diversi-
acteristically small scale and fkquent, forest management should ty and ecosystem health and integrity in the development of
emulate this. Where natural disturbance is severe, large scale sustainable forest management (SFM) systems unless we
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and infrequent, forest management should have the same understand the meaning of these concepts. These ecosystem
characteristics if it is the objective to sustain the historical con- attributes cannot be used as criteria in the certification of SFM
dition of the forest. In the former type of forest, management unless we understand them and apply them with scientificrigor.
may appear to "revere" nature by a "soft-footprint" approach; Using them only in terms of environmentalmythology will not
in fact it simply "takes particular notice" of the ecological char- help us to fulfill our intergenerational equity obligations.

232 MARSIAVRIL 1997, VOL. 73, NO. 2, THE FORESTRY CHRONICLE