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Journal of Arthropod Identification
ARTHROPOD BIODIVERSITY PROJECTS BUILDING A FACTUAL FOUNDATION
Biological Survey Of Canada (Terrestrial Arthropods)
Document Series No. 7 (2000)
|Guidelines for conducting
studies of arthropod biodiversity properly are reinforced using results from selected
recent studies in Canada. The costs for doing such work are also given explicitly. The
necessary components of a biodiversity study, and selected examples, are briefly tabulated
for ready reference. Careful advance planning should include explicit scientific
objectives and ways to ensure that the work proceeds to completion. Work on more than one
taxon is necessary, because neither patterns of species richness nor relevant ecosystem
involvements can be extrapolated from one taxon to another. Plans for identification,
normally to species, are especially important, requiring specific collaboration with
systematists. Protocols for sampling, sorting, specimen preservation and data management
should be clearly defined and costed. Curation and retention of specimens and ongoing
scientific and other publications are also essential if projects are to have real
long-term value. Examples and references illustrate how these components can be developed.
Proper support for studies of biodiversity, as opposed to superficial promotion of its
importance, therefore requires mechanisms to provide stable long-term funding.
Projets sur la biodiversité
des arthropodes terrestres établissement dune base de travail
Des exemples détudes canadiennes récentes sur la
biodiversité des arthropodes viennent valider les directives suggérées précédemment
pour ce genre de travail. Les coûts reliés à ce type détude sont évalués de
façon détaillée. Les composantes essentielles dune étude de la biodiversité, de
même que des exemples choisis, sont présentés ici sous forme de tableaux de référence
simples et commodes. Une planification soignée suppose lidentification des
objectifs scientifiques précis et des méthodologies qui assureront la bonne marche du
projet jusquà la fin. Létude de plusieurs taxons à la fois simpose,
car ni les patterns de la richesse en espèces, ni linfluence des variables
écologiques ne peuvent être extrapolés dun taxon à un autre.
Lidentification des organismes, normalement jusquà lespèce, est un
aspect particulièrement important et requiert le concours de taxonomistes reconnus. Les
protocoles déchantillonnage, de tri, de conservation du matériel et de traitement
des données doivent être définis clairement et leurs coûts évalués. La mise en
collection des spécimens et la planification des publications scientifiques et autres
sont aussi essentielles pour garantir lintérêt à long terme des projets. Des
exemples et des références viennent illustrer comment ces composantes peuvent être
agencées. La réalisation détudes de la biodiversité qui soient plus que de
simples exercices de relations publiques suppose donc la mise en place de mécanismes
propres à assurer un financement continu sur une longue période.
Table of contents
Scope and scale of the work
Steps in biodiversity assessment
Sampling and sorting
Results and completion
Costs for sampling and sorting
Costs for identification
Costs for curation and publication
Appendix: Recommended procedures and
Interest in studying biodiversity, including that of
arthropods and other invertebrates, increased following the International Convention on
Biodiversity that was signed following the landmark meeting in Rio de Janeiro in 1992.
This convention was followed up in Canada (e.g. Canadian Biodiversity Strategy 1995), and
as a result increasing numbers of studies on biodiversity began. However, not all of them
were based on detailed knowledge about the true, very great, extent of arthropod diversity
nor on realistic plans for its assessment. To foster more fruitful approaches, the
Biological Survey of Canada produced two briefs offering prescriptions for how to carry
out work aimed at assessing and interpreting the composition of the fauna (Marshall et al.
1994; Danks 1996).
The results of recent work in Canada is now becoming
available. General information from elsewhere is also accumulating (compare Groombridge
1992; Stork 1994; Stork et al. 1997; New 1998; Redford and Richter 1999; some of the
papers on methods cited below). This brief therefore takes stock of the guidelines
proposed earlier. Such an examination has proved to reinforce most of the original
prescriptions, because we are able to outline the positive rewards of following nearly all
of them. Here, we emphasize Canadian work familiar to us, especially work in ancient
forests in western Canada, although some of the same points could have been made using
The examination reinforces earlier contentions that doing
things properly pays off, whereas dabbling in biodiversity is a waste of resources. In
other words, effective studies to establish a firm factual foundation for understanding
biodiversity demand scientific focus and detailed planning for all aspects of the work.
scale of the work
Because arthropods are so diverse, effective study of their
diversity requires a relatively large scope and a relatively long time frame. The main
focus here, therefore, is on major projects. However, studies of smaller scope (such as
discrete graduate student projects) can make key contributions, especially if they address
key themes, questions or taxa within the context of a larger investigation. For example,
investigations of biodiversity have been made in relation to forest age for a series of
taxa in Newfoundland (Arulnayagam 1995, Puvanendran et al. 1997 for springtails; Dwyer
1995, Dwyer et al. 1997, 1998 for mites; McCarthy 1996 for beetles), and for different
forest practices in Alberta (Niemelä et al. 1992, 1993, 1994; Langor et al. 1994; Spence
et al. 1996, 1997; see also http://www.biology.ualberta.ca/emend/emend.html)
and in British Columbia (Lavallee 1999, Craig 1995, McDowell 1998, Lemieux 1998 for
carabid beetles; Brumwell 1996 for spiders). These investigations illustrate the
importance of working within a defined conceptual context, which can involve a focus on
patterns of diversity (such as species richness) or a focus on processes relevant to
ecosystem integrity (such as the effects of disturbance) which function through species
and their interactions. Broad assessments of diversity have been initiated in Quebec
(Paquin and Coderre 1997a, b), Ontario (e.g. Blades and Marshall 1994), Alberta (e.g.
Finnamore 1994; Hammond 1997) and B.C. (e.g. Blades and Meier 1996; other studies cited
below). Studies of insects and other groups on the Brooks Peninsula, BC, were carried out
in the context of the possible existence of a glacial refugium in the area (Cannings and
Cannings 1997). All of these investigations confirm the need to plan for the long term and
especially for the inclusion of a wide range of taxonomic expertise.
Information on biodiversity is linked closely to scale.
Biological relationships differ at different levels, on a range from microhabitats, sites,
landscapes and regions to global patterns. For example, characteristic taxonomic patterns
change with latitude (Danks 1981, 1992, 1993). The predominance of particular functional
groups or the dominance of particular taxa changes according to the spatial context (cf.
Anderson 1997; Niemelä 1997). Therefore, typical studies should be designed to address
several different scales, because it is not always clear beforehand at what scale
particular processes or patterns operate. Moreover, results for one scale cannot simply be
extrapolated to apply at a different scale.
An adequately planned scientific study of
biodiversity requires attention to a large number of steps, from initial objectives and
planning to the disposition of specimens and the publication of results (Danks 1996,
1997). These steps, and their potential benefits, are outlined in the first two columns of
the Appendix. The final two columns of the Appendix
outline selected specific studies that confirm these benefits, with references for the
examples. The sections below provide further details and examples of these essential
The scientific objectives of a biodiversity study should be made
clear from the outset, so that work can be targeted at both longer-term and shorter-term
results of interest.
Establishing the richness and relative abundance of species
in specific habitats makes data available as a long-term reference for future
use. For example, individuals of the predator guild, composed primarily of arachnid
species, are numerically dominant in various sites in the ancient forest canopy. Such
numerical dominance (e.g., Voegtlin 1982; Winchester 1997a, 1998; Winchester and Ring
1999) helps to characterize these systems for current and future comparison. The need for
detailed work to establish a reliable basis for comparison is confirmed by the fact that
the number of species in different sites is similar, but the actual spider communities are
dissimilar (e.g. Halaj et al. 1998). For example, only five of the many spider species
were shared by all of the three ancient forest sites (Mt. Cain, Carmanah, Rocky Point)
studied in this respect in British Columbia (Winchester and Ring 1999).
Specific objectives also are very important for the
relevance and continuity of a study. For example, the impact of changes can be assessed by
properly designed studies, provided the objectives are considered at the outset. Thus,
comparison among forest sites of different ages and histories can provide information to
resolve conservation and other issues: ancient forests differ in guild structure and other
elements even from relatively old second-growth ones, and apparently certain species
persist only in the ancient forests (Battigelli et al. 1994; Arulnayagam 1995; McCarthy
1996; Spence et al. 1996, 1997; Brumwell et al. 1998; Carey 1998; Winchester 1998; Fagan
and Winchester 1999). However, the prevalence of species characteristic of older stands
varies among taxonomic groups (see below).
A shorter-term objective of particular current interest
concerns the impact of introduced species, which can be evaluated using standard
protocols. An example is the tracking of bark beetles (Scolytidae) in western forests
using specific sampling procedures, long-term monitoring, and an extensive reference
collection (L. Humble, pers. comm.)
Work focussed on key groups can provide valuable information
about specific habitats after a relatively short period of time. For example, some species
characteristic of the canopy of ancient forests are rare and potentially endangered
(Campbell and Winchester 1994; Winchester and Ring 1996; Winchester 1997b; Behan-Pelletier
and Winchester 1998; Marshall and Winchester 1999; Walter and Behan-Pelletier 1999;
Behan-Pelletier 2000; Humble et al. 2000).
Proper general planning also profits from assembling
previous knowledge and experience. Such aspects are treated by Danks (1996) and
Winchester (1999b); see under Sampling methods for more details.
Coupled with these plans in the context of the project
objectives, a well constructed overall plan is required to define the focus and
the availability of resources for the project. A focussed study asks specific questions
from a scientific perspective, and so is more valuable than an inventory alone. For
example, some species and habitats are especially sensitive to change from human activity.
The canopy of ancient forests has suspended soil habitats that are easily
disrupted (e.g. by timber harvesting) and all but impossible to recreate, because
second-growth stands, even those 80-120 years old, do not have well developed suspended
soils. Certain species of oribatid mites are characteristic of the suspended soils in
these ancient forests (Behan-Pelletier and Winchester 1998; Winchester et al. 1999), and
would be especially sensitive to environmental impacts. Other taxa, such as mites of the
family Zerconidae, and certain springtails (Fjellberg 1992) also have distinct arboreal
A major requirement of the overall plan is collaboration with
systematists and ecologists, before the project begins, to help select key taxa, design
the sampling, and ensure identifications. The importance of systematics in this context
cannot be overemphasized (cf. Danks 1996; Vane-Wright 1996). Also, the flow of
resources must be adequate to complete the project. Without explicit attention to
longer-term resource needs, there will probably be little pay-off from the work in the
form of curated specimens, reliable and accessible databases, publications, and
conclusions useful to support further action. An analysis of various resource trade-offs
of this sort was made for a major project on forest biodiversity monitoring in Alberta
(Schneider et al. 1999; Winchester 1999b).
Overall planning also requires statistical expertise
early in study design, because methods of analysis must be appropriate to test the
objectives of the study. Because any particular statistical program or analysis requires
that data be entered and stored appropriately, initial datafiles have to be set up
properly to favour comparisons. For example, including taxa from all sites and treatments
in a single database (so that absence as well as presence is explicit for a given site)
allows statistical analysis for differences, and avoids the need to transcribe data from
separate presence only lists of taxa. Thus most statistical analyses of
species richness and abundance require that sites (treatments) form the columns of data
and that species (items) form the rows: zeros show absence and numbers of specimens show
abundance. Statistical programs normally treat such issues in their documentation (e.g.
Within the framework of a general plan, important planning
decisions have to be made in more detail, especially the level of identifications, the
selection of sites, the target taxa, and the duration of the study.
Detailed information of most value for the project normally
comes from identification to species. In general terms, of course, species names
are used to access all biological information (e.g. Danks 1988), so that identification to
species allows much wider information to be brought to bear on the results. In particular,
attributes that determine the meaning or importance of the results, such as habitat
specificity, range, or biological interactions typically are visible only at the species
level but not at higher taxonomic levels. For example, many species of arthropods occur
only in the high forest canopy, but such arboreal specificity cannot be detected at the
family level, as for oribatid mites (Walter and Behan-Pelletier 1999). Differences among
spider communities from similar habitats in different geographical regions are visible
only from differences in species composition, and not from general measures such as
species richness or distinctions at higher taxonomic levels (see above; Winchester and
Ring 1999). In this and other contexts, the most striking conclusion about biodiversity
studies is that there is very limited resolution without species identifications (Cross
and Winchester 2000; Fagan 1999). Of course, such interspecific differences occur on a
variety of spatial scales (see above), requiring deliberate decisions about the scales on
which sampling will be carried out.
Material is analysed in a sequence, formalized into a series
of 7 steps by Winchester (1999b). Each successive step adds complexity (and cost) to the
analysis, but at the same time increases sensitivity. The first three steps require
arthropod sorting to ordinal or family level, but give only coarse-grained resolution.
Subsequent steps require identification to species for the designated target taxa, but
provide fine-grained sensitivity to answer biodiversity questions. Useful information
about trophic interactions can sometimes be obtained by analysis of guilds at the family
level, for example, but typically higher-level sorting is only a staging point in the
resolution of pivotal questions using species-level identifications.
An ideal site for study would be accessible,
discrete, stable, well characterized and representative (Danks et al. 1987; Danks 1996).
Such a site makes sampling efficient. Of course, the sites should also be relevant and
adequate for assessing the target taxa. In practice the choice of a site will be a
trade-off between such things as ready access to a close-by area where a complete sampling
program can be carried out relatively cheaply, and a unique, unexplored area of great
scientific and conservation value but involving a large cost because of its relative
inaccessibility. For example, work in the Carmanah Valley, British Columbia, was led by a
high conservation concern for a unique and previously unsampled habitat that includes the
worlds largest Sitka spruce trees. Therefore, residues (such as the Diptera sorted
to family) from multiple samples from this unique site were stored for future benefit (see
The choice of taxa for the initial work depends on
the scientific objectives, but may be modified by the relative ease of trapping or
identification. Nevertheless, using feasibility alone to select taxa is not desirable. In
several studies carabid beetles have been chosen almost by default because they can be
identified relatively easily, and not necessarily for biological clarity in addressing the
Because detailed study is costly, many workers have tried to
identify indicators characteristic or surrogate species or sets of
species that could serve as a magic bullet to describe and evaluate
ecological conditions (see McKenzie et al. 1990; McGeoch 1998). Valid indicators would
greatly reduce the need to cope with the true diversity. Unfortunately, the criteria used
to select indicators are seldom based on objective information. Specific indicator choice
generally has not been supported by scientific findings or methodology, although useful
procedural steps for justifying the choice are given by McGeoch and Chown (1998) and
Dufrêne and Legendre (1997). At present there is little evidence to suggest that any
taxon or group of organisms qualifies as a universal biodiversity indicator. This is true
from a broad faunal viewpoint, but also for more specific questions. For example, in the
study by Neave (1996), species confined to old-growth forests were present, but not in
this study among the carabid beetles, the group most often studied in such habitats in
Canada and for which old-growth specialists have been claimed elsewhere (e.g. Spence et
al. 1996). Results for one taxon do not predict results for another over different spatial
scales (e.g. Lawton et al. 1998; Prance 1994; Williams and Gaston 1994; Niemelä 1997).
This finding is not surprising, because ecological patterns and processes depend on scale
(see above), and different groups differ in local species richness, endemism, habitat
association or specificity, vagility and so on (e.g. Prendergast et al. 1993). Some
insects are relatively insensitive to environmental factors of potential interest (e.g.
Whitford et al. 1999). Faunal composition is dynamic over time (e.g. Ellis et al. 1999).
In other words, ecosystems are complex, so that multitaxa approaches (e.g. Hammond 1994)
are more realistic.
A second type of short cut in work on biodiversity is the use
of summed statistics, often based on supraspecific taxa and usually called biotic indexes.
These indexes are supposed to allow simplified comparisons with normal
conditions or with other sites. They are best developed for rapid
assessment of aquatic systems (e.g. Washington 1984; Hilsenhoff 1988; Klemm et al.
1990; Rosenberg and Resh 1993; Cao et al. 1996; Pflakin et al. 1989). Some of them merely
give numerical scores to specific indicator organisms (see above). However,
others rely on more or less extensive calculations from the taxonomic composition or
abundance at a given place (compare McGurran 1988 for the calculation of various indexes;
Colwell 1999b chiefly for estimating species richness). Typically, they confirm obvious
differences, but their validity for assessing subtle differences is far from clear.
Indeed, different and more useful results are obtained when working at the species level
(which takes account of individual species biologies) than by using a particular biotic
index. For example, recommendations to prevent the decline of a rare species dependent on
a particular foodplant or on a key microhabitat would not emerge from a statistic based on
insect community structure.
Given these difficulties, there is no clear consensus as to
which groups or species best allow environmental changes to be assessed. Few attempts have
been made to contrast attributes that might confer susceptibility or resilience to habitat
change. Individual species responses are the key to understanding and measuring these
impacts, but species so far examined in biodiversity studies in northern temperate forests
might be atypical of the majority of invertebrates because they are taxonomically
convenient (e.g., carabid beetles: Huhta 1971; Holliday 1991; Niemelä et al. 1993; Spence
and Niemelä 1994; Spence et al. 1997), conspicuous (e.g., butterflies: see Davidar et al.
1994), broadly distributed geographically (see Platnick 1991; Danks 1993, 1994), or have
specific habitat requirements (cf. Danks 1979). Moreover, experiments have not always been
designed to resolve the effects on local biodiversity of factors such as habitat loss,
degree of spatial or temporal isolation, and dispersal capabilities, which influence
different taxa to widely different degrees. In other words, ecosystems are heterogeneous
(cf. Niemelä 1997), the species richness of different taxa is not well correlated, and
limited or arbitrary choices of the study taxa are unwise.
These complexities confirm that it is necessary to sample and
assess more than one taxon, and to include the diversity associated with different
habitats at different spatial scales. Focussed collecting by collaborating systematists in
addition to the standardized trapping is very helpful for such assessments. By the same
token, biological features of the taxa should also be considered explicitly in the context
of the study. For example, groups characterized by many species with wide distribution and
high ecological valency are less likely to point to key microhabitats than are groups with
many distinctive stenotypic species.
Finally, because most research on arthropod biodiversity
relies on multiple traps, many arthropods will be trapped that do not belong to the
initial target taxa. However, these residues are likely to hold the answer to many
questions, because we do not necessarily know in advance which group might be suitable to
seek the answers, especially because biodiversity concepts, questions and analytical tools
evolve over time. Assuring proper curation and availability of the residues therefore is
of great future service (see curation below).
The duration of a study depends on its objectives.
Valuable information, for example about habitat associations, can be gained by short-term
inventories, but long-term events cannot be assessed from data for a single season. Even
simply assessing the richness of a particular group requires sampling at least throughout
the active season. From a practical point of view, whether duration (and sampling methods)
are adequate to document richness can be assessed by plotting species accumulation curves:
the increase in number of species and its seasonal pattern help to show how likely it is
that the samples give a valid estimate of the real diversity (see Coddington et al. 1991;
Soberon and Llorente 1993; Colwell and Coddington 1994; Longino 1994; Longino and Colwell
1997; Colwell 1999a).
Multiple rather than single sampling methods usually
are desirable. Using a variety of trapping techniques increases the numbers of species
sampled, covers more habitats at a particular study site, and samples a range of species
with different vagility. These findings apply to insect orders, and apparently also to
families and species (Southwood 1968; Blades and Maier 1996; Winchester and Ring 1996;
Winchester 1999b). For example, adding Lindgren traps to pan or pitfall trap sampling
increases the number of beetles caught, and catches additional species (L. Humble, pers.
Making the sampling methods standard is very
important to allow meaningful, including statistical, comparisons. For example,
standardization allows comparisons of sites, forest ages, and so on (e.g. Spence et al.
1997; Trofymow and Porter 1998). Trapping specifications, including trap type, trap
construction and sample design were tested in several large-scale biodiversity studies in
Canada and the results (although typically they were not formally published) were used to
improve sampling recommendations summarized in Marshall et al. (1994), Winchester and
Scudder (1994), Behan-Pelletier et al. (1996), Finnamore et al. (1998) and Winchester
(1999b) (see also Paquin and Coderre 1996; for a tropical site see also http://viceroy.eeb.uconn.edu/ALAS/ALAS.html).
The progression and development of these standard protocols, and the evolution of the
ideas from 1994 until the present day, was expedited through rigorous field work,
collaborations among researchers, and the ability to make firm comparisons based on
properly curated reference collections and secure permanent sample sites. Aspects of these
standards, covered in detail in Winchester (1999b), include the sampling protocols and
logistics appropriate for monitoring, detailed costing for time, equipment, processing and
storage of data, integration of data, archiving target groups and residues and the
infrastructure needed to complete the entire field and laboratory segments of the project,
as well as data management considerations.
Because biodiversity sampling of arthropods produces so many
specimens, procedures must be as cost-effective as possible. Specific analyses of
cost are a relatively recent development (Marshall et al. 1994; Scudder 1996; Winchester
1999b; see section on costs below). Cost-effective sampling also comes from fine-tuning
procedures for access, sampling and trap placement over time (Winchester, pers. obs.).
The significance of differences to answer ecological
questions or test hypotheses can be assessed only with adequate sample design, including replication
(cf. Krebs 1989, chap. 8). Some ways exist to assess the numbers of replicates needed (see
references under Duration of a study, above; Didham et al. 1996). Unfortunately, explicit
statistical tests (e.g. ANOVA) are still relatively rare in studies of arthropod
biodiversity and the high cost of sampling and sorting misleads many workers into using an
inadequate number of replicates.
Sampling must be done carefully. The only way to ensure quality
control for a large-scale project is to formalize the relevant protocols. For
example, the field crew for a pilot project on Alberta forest biodiversity was trained
prior to the start of the pilot project in all aspects of the program, according to
established procedures for trap installation and other details (D. Farr, pers. comm.; see http://fmf.ab.ca/pro.html).
Sorting is more labour-intensive than sampling: sufficient
time for sorting must be planned for, as well as its substantial costs (see section
on costs below). Note that the efficiency of individual sorters also differs. In all
cases, formal protocols for sorting (to avoid cross-contamination of samples, for
example), and preservation and curation are needed (see below). Providing continuing
positions for trained people, such as parataxonomists and biodiversity technicians,
increases the efficiency of sorting and ensures that only as much material as necessary
will be prepared further for identification.
Specific procedures for mounting specimens should be
obtained from the taxonomists who are collaborating in the project, because each taxon
(and even each scientist) has particular requirements. For example, sphaerocerid flies are
best prepared by critical-point drying, pompilid wasps require male genitalia to be
exposed to aid identification, braconid wasps can be prepared using amyl acetate, and
oribatid mites must be correctly mounted on microscope slides to avoid damage and leave
key characteristics visible.
Sorting procedures depend especially on the level of sorting,
but a hierarchical process is most efficient. For example: 1. Target all Coleoptera from
all traps (storing residues properly after Coleoptera have been removed), and enter
Coleoptera data into a database; 2. Target specified families from the Coleoptera sets,
and store the rest of the Coleoptera as residues; 3. Mount and enumerate all specified
families, involving recognition of morphospecies and association with data labels; 4.
Proceed to identification by group as required.
It is also essential to develop a system for tracking each
specimen and data on its trap, site, date of collection, etc., so that the ecological
information associated with each specimen (some of which may be retained by specialists)
can be retrieved for the project analysis (see also data management below).
Identification to species is especially important to interpret
results, but requires particular care in planning because there are so many species, many
groups are inadequately known, many others are difficult to identify, and taxonomic
resources especially specialists are in short supply. For additional discussion, see above
under Detailed planning. Collaboration with taxonomists is especially important to ensure
that they will be able to budget the necessary time.
Several simple steps facilitate identification. For example,
it is necessary to prepare and ship material so that it arrives undamaged (e.g.
Martin 1977). Hand delivery is often the best choice. Shipment details vary with the type
of specimen and the destination, but some organizations have developed formal internal
protocols for their own loans or other purposes.
Information provided with specimens can assist
identification (and in turn a specialist may then be able to supply useful ancillary
information about the taxa). If mass collections are supplemented by other
material, the task of identification can likewise be made easier. For this purpose, as
well as more generally, working with a taxonomist in the field can lead to optimal use and
placement of traps and a better understanding of basic biology of the target group, which
may help to focus on key hostplants, for example (e.g. Goulet 1996 for sawflies in
Strathacona Park, British Columbia).
Such arrangements exemplify collaborative work,
which has many benefits for both parties. For example, results from broader studies can
add to knowledge of species distributions at provincial or national levels, and unique or
duplicate specimens can be retained for collections and publications at no cost to the
specialist in return for identifications. Published results of such collaborations address
ecological and taxonomic themes (including descriptions of new species), for example
Fjellberg (1992), Campbell and Winchester (1994), Lindquist (1995), Behan-Pelletier and
Winchester (1998), Winchester et al. (1999), Marshall and Winchester (1999) and Coher
It is important to allow sufficient time for the
identifications. For example, the summary report about invertebrates of the montane
forest at Mount Cain, B.C. (Winchester 1999a) contains 4 published or in press papers and
an MSc. thesis, reflecting a long-term perspective and the sort of output, validated by
peer review, that is desirable (see Results and Completion below). Results from the major
study of arthropod biodiversity at Carmanah continue to be published 7 years after the
start of the project. Such a pattern will continue as the necessary taxonomic research is
completed, confirming the need in any study of arthropod biodiversity for long-term
attention to identification.
Proper payoffs from a study are not possible without explicit
attention to how data management, curation and publication will be planned for and
A data management system should allow all
information to be entered in a standard way from the beginning, so that data do not have
to be reformatted or re-entered at a later stage. Specific attention should be given
before the project begins to how data will be analyzed statistically (see under General
Planning above). Ordinary database or spreadsheet software (e.g. Access, Excel) can be
used for databases of smaller scale. Several systems allow much greater sophistication,
for example Biota, which helps to manage specimen-based biodiversity and
collections data by providing an easy-to-use graphical interface to a relational database
structure (Colwell 1999a). It is important that the taxonomic and geographic information
is reliable as well as accessible (Mickevich 1999).
Databases of larger scale require full-time database
managers. Extensive use of such data requires cooperative endeavours to develop standard
fields (cf. Noonan 1990) and to permit information with different origins to be accessed
electronically. Such larger concepts are being addressed by a number of working groups and
pilot projects (cf. Umminger and Young 1997; ITIS.ca 1999). Distributed databases
coordinated through a central infrastructure show particular promise.
Proper curation (including labelling, storage and
access) and retention of voucher specimens is essential for two reasons. First, specific
comparisons can be made during the project. Second, later work can be compared or
validated effectively. From a general viewpoint, of course, the value of museum
collections in these contexts has been stated many times (e.g. Danks 1988, 1991; Wiggins
et al. 1991). Such values include the basis for taxonomic work (including type specimens
and material for species descriptions) and for ecological work (including association
between species and habitats). For specific biodiversity projects, collections often have
to be revisited to answer taxonomic or ecological questions and to refine the database,
which can only be done if the material is properly organized. A well organized collection
saves resources (both for mounting, and the time of specialists), because a large
percentage of any trap catch is composed of relatively few species with many individuals,
and for many taxa a reference collection allows these species to be identified easily.
Such reference uses within the project itself mean that material should be properly
preserved and mounted, as indicated above.
Much of the value of typical biodiversity samples of
arthropods comes from later use. About 94% of most Malaise-trap catches in ancient forests
are Diptera, but no taxonomic experts are currently available for many groups. Therefore,
most dipteran groups would not form the initial focus of a project. However, given the
ecological importance of the Diptera, there is great value in properly curating these
specimens. Material extracted from the Carmanah residues (preserved as explained above
under site selection) formed the basis for subsequent studies of mycetophilids, syrphids
and sphaerocerids (Coher 1995; Samoszynski 1998; Marshall and Winchester 1999).
Any project that will generate extensive material therefore
needs a plan for long-term curation at an institution equipped to do so. For example,
collections, especially from some of the projects referred to in this brief, have been
consolidated though the participation of the Pacific Forestry Centre in Victoria (Canadian
Forest Service), which now acts as a repository for all of the ancient forest arthropods
from projects in British Columbia. Conversely, the Royal British Columbia Museum was
unable to incorporate these collections.
Proper publication of results is essential to profit
from biodiversity studies in the long term, because general patterns are likely to become
visible only when a range of sites from different areas and habitats can be compared. On
the other hand, typical consultant and other reports that have not been validated through
the process of scientific publication are of very limited value. Recent findings, already
cited, that are visible from peer-reviewed publications include the facts that ancient
forests are source areas of high arthropod diversity; that this diversity applies to
several groups; and that certain habitats, and their associated species many of which are
undescribed, are found only in ancient forests (and not in second growth areas), including
unique microhabitats such as suspended soils. In turn, these findings suggest that
retaining diverse habitats is the key to meeting biodiversity commitments.
Such results can also be put to wider use beyond
dissemination to scientists. Objective information about the Carmanah Valley was made
available to a wide-range of user groups by means such as popular articles and a web site
(http://web.uvic.ca/~canopy, which lists other
relevant items). Such information for general audiences helped to foster the establishment
of Park status for the Carmanah site, for example.
Ongoing publications therefore lead to greater understanding
and appreciation of the systems studied. They also tend to lead to growing interest in the
sites, as manifested (e.g. for the ancient forest sites emphasized here) by increasing
collaboration for work on additional taxonomic groups, increasing interest in the sites
and the collections by potential graduate students, national collaborations with
interested agencies, and coverage by print and electronic media.
The costs of proper studies of biodiversity often have been
underestimated, because resources have to be made available over the whole period from
initial planning to completion and documentation, and because many tasks are very
labour-intensive. However, ongoing documentation of costs is very useful for steering
long-term projects. A synopsis of realistic costs for various stages of a study are given
for sampling and sorting
It is necessary to sample and sort using a reasonable number of
replicates, with a few selected techniques, for a diversity of taxa (as discussed above),
so that the total cost is significant. Tables 13, derived from careful documentation
of earlier efforts, shows sample elements of the cost of traps, and typical expenses for
sampling and sorting. The tables show that materials and technical assistance for a
general survey of terrestrial arthropods with initial sorting to order, and then to family
only for the flies, would be expected to cost more than $10,000 per site per season. This
cost would comprise $2016 (cost of traps, Table 1) + $1350 (installation and servicing,
Table 2) + [$1512 (family sort for Malaise Trap samples, Table 3) x 5 (assuming similar
catch and effort for other trap sets)]. It is not realistic to trim the trap and servicing
costs ($3366), because they reflect the intensity of effort that is necessary to secure
reliable data. The cost of sorting depends partly on the careful selection of taxa with
reference to the study objectives. Sampling and sorting aquatic insects is also very
time-consuming (e.g. Ciborowski 1991), taking at least as long as for the examples given
Table 1. Summary of minimum equipment costs ($$
Canadian) for a complete set of traps for a single site (from Winchester 1999b).
|No. of traps per site
|All traps at site
|Grand total per site
*Pitfall traps include cups and covers.
+(12) pans are the pans placed under the Malaise trap.
Table 2. Estimates of minimum costs for installation
and servicing of a complete set of arthropod sampling traps for one site, comprising 3
Malaise, 25 Pitfall, 10 Pan, 12 Lindgren, 6 Soil and 3 Litter traps or samples (from
Cost ($$ Canadian)
|Installation of all traps
|Single servicing of all traps
|Servicing of all traps for the
|Total seasonal cost (1 site)*
*Total cost is based on an experienced field crew of 2 people, working for $15
per hour and does not include travel time or benefits.
Table 3. Summary of minimum costs associated with
sorting for selected taxa in 30 Malaise trap samples collected from a single study area on
Vancouver Island (from Winchester 1999b). Total cost for one site is then calculated from
the mean sort time (see table), recommended number of traps per site (3) and recommended
number of samples per season (12).
|Ordinal sort All orders in samples
|No. of Orders
|No. of individuals
|Sort time (hours)
|Family sort after Order-level sort 17
Families of the order Diptera
|No. of Families
|No. of individuals
|Sort time (hours)
Estimated cost for 1 site @ $15 per hour:
Mean per trap sample
3 traps for the season
|Order-level sort (9.3 hours)
|Family-level sort after Order-level sort (2.8 hours)
The cost of obtaining independent identifications to species,
even if expertise exists and is available for hire (an assumption not often met), varies
with the group but normally is very high because in most groups developing the ability to
make accurate identifications requires specific long-term experience. For example, expert
identification to species is offered by the Entomology Department of the Natural History
Museum of London at a cost for each ordinary specimen equivalent to about $150, and even
more for difficult specimens or groups (see http://www.nhm.ac.uk/entomology/insident/).
On this basis, the identification of a modest range of arthropod material from one site
would cost many thousands of dollars. Specific identifications made more cheaply by
generalist contractors are likely to be unreliable.
Therefore, the direct costs of obtaining independent
identifications from qualified systematists normally are avoided by involving systematists
closely in the planning, execution and publication of biodiversity studies.
for curation and publication
Adequate follow-up requires curation at least of voucher
material, and the publication of results.
The cost of curation has been established for a number of
taxa. It is especially high for large vertebrates, but for most arthropods the cost of
bringing a specimen into a collection and documenting it ranges from about US $0.25 to
$4.00, or $5.00 per lot (West 1988; see also Anderson 1973). The subsequent annual storage
and maintenance cost for insect specimens, mainly for housing and adequate environmental
control, is much lower if the material is simply stored rather than worked. Nevertheless,
acquisition and long-term preservation of a substantial voucher collection costs thousands
of dollars (compare Lord et al. 1989b and preview in Lord et al. 1989a; Lord 1991) and two
thirds of the operating costs of typical museums are devoted to direct and indirect costs
associated with collections (Lord et al. 1989a). However, both the cost of obtaining
properly sorted and preserved material and its long-term value for research and reference
is many times the cost of curation (cf. Wiggins et al. 1993).
Publication in generally available, peer-reviewed, journals
requires significant time and expertise, even if page charges and similar costs can be
avoided by the choice of particular journals. Studies of biodiversity conducted by
consultants therefore suffer from the great weakness that not only may there be little
incentive to publish, but also the cost of employing qualified personnel to prepare
appropriate manuscripts (as opposed to much less rigorous reports) is not normally
included in the contract price.
|This review of some recent
results in the light of previous recommendations for biodiversity studies strongly
confirms the earlier advice. The necessary procedures can be summarized quite simply
do things properly:
- Think through the scientific objectives carefully, which will
help to define the target taxa and sites.
- Make sure that full advantage is taken of existing
information, specimens and publications.
- Plan the sample design and select traps and other components
after detailed consideration of the project needs.
- Plan explicitly for expert identification of material,
normally including a focus on species rather than higher taxa, and seek early
collaboration with systematists.
- Define the actions to be taken in detail, using clearly
defined protocols for sampling, sorting, specimen preservation, and data management.
- Ensure that resources and plans are in place for the long
term, including curation of specimens in a permanent repository, and publication in the
scientific literature and elsewhere.
Knowledge of arthropod biodiversity and resources for
reference can be built up for wide long-term benefit using this procedure, instead of
consuming funds on short-term projects that generate non-standard data of limited detail.
In other words (contrary to some hopes or expectations), proper studies of arthropod
biodiversity cannot be done over very short time frames. Cheap and limited efforts in
isolation will produce worthless results. In particular, it is worth noting that even when
the specimens have been collected only a very small percentage of the work has been done.
The major data come from sorting and identification, and their major value comes from
analysis and dissemination.
Securing funding for biodiversity studies for the long term
rather than the short term therefore is the key need, both for overall scientific payoffs
and for logistic reasons such as maintaining secure sites, keeping trained field and
laboratory crews in place, and allowing appropriate taxonomic expertise to be developed.
For many years, lip service has been paid to long-term biodiversity issues, such as
environmental sustainability. It now has to be matched by long-term support for research,
so that a sound factual foundation can be established from studies of biodiversity that
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Recommended procedures for biodiversity studies (based on Danks 1996), and
examples of the benefits of following them. For further details and examples see text.
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Establish long-term baselines
||Comparison with other systems
||Identification of spider fauna from ancient forest
sites shows the general importance of the predator guild there
||Winchester and Ring 1999
||A reference baseline has been made available for
important ancient forest habitats. Also, detailed work has led to tested and effective
protocols for study (see below)
||Winchester and Ring 1999; Winchester and Fagan 2000
|Obtain specific answers, e.g.:
|Impact of change
||Experimental design allows valid comparisons
||Differences in the fauna associated with forest age
were established, including the presence of rare species that persist only in ancient
||Spence et al. 1996; Winchester 1998
|Uniqueness of habitat
||Focus on key groups
||Characteristic species of oribatid mites were
identified from ancient forests
||Behan-Pelletier and Winchester 1998
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Gather previous and background knowledge
||Unnecessary repetition avoided
||Existing information on design, sampling techniques,
analysis and costs were used to plan a major monitoring programme efficiently
|Scientific focus for feasibility
||Lasting scientific benefit from proper foundation
||Detailed information for selected taxa (e.g. oribatid
mites) shows the uniqueness and richness of the fauna and hence its importance for
||Winchester and Ring 1999
|Early collaboration with systematists and ecologists
||Material and data collected can be interpreted, not
||Collaboration with systematists and collections
enhanced identification and preservation of data and specimens
||(See Identification and Specimen curation below)
|Resources adequate for all phases
||Samples exploited efficiently
||Appropriate trade-offs were analyzed to help design a
major sampling program
||Curated, labelled material set aside for future
||Handling of data appropriate for objectives
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Identification to species
||Information referenced properly in the biological
literature for future use
||Species names are the universal biological currency
||e.g. Danks 1988
||Detailed information of value for the project
||Site specificity of oribatid mites and staphylinid
beetles (and hence the importance of sites for various purposes) was shown by
species-level but not by genus- or family-level identifications
||Behan-Pelletier and Winchester 1998; Winchester 1997b
|Accessible, discrete, stable, well characterized and
representative; and appropriate for selected taxa
||Sampling less costly, sites easily recognized and
sampled, protected from damage, background information available and results more widely
||Including these criteria improves efficiency, although
there may be other trade-offs
|Balance between scientific utility and feasibility
||Best compromise between the features of organisms that
may address project objectives (such as diversity, habitat specificity, dispersal ability,
feeding habits, and others), and the practicalities of sampling, sorting and
||Many benefits from a proper focus on the objectives
coupled with planning from sampling, identification and other standpoints
||For sample considerations see Samways 1992, Kremen et
al. 1993, Danks 1996, Winchester 1999b
||Select several taxa (not one), and also endeavour to
||Focus is feasible though not too narrow, and
additional material is available for future use
||(For discussion see text)
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Long enough to provide information about
long-term natural events of interest
||Answers address project objectives
||Long-term studies (long
duration mimicked by matched sites of different ages) can show the long-term effects of
fire and logging
||See EMEND website
|Long enough to compensate for annual and
seasonal variations in life cycles and in populations accessible for sampling
||Samples accurately represent diversity
||Only season-long sampling of staphylinid
beetles provided a valid annual data set, because the species collected shifted through
|Multiple methods, and that are appropriate
for the taxa selected
||Provides proper coverage for the selected
||A full range of species is obtained
through combined catches from Malaise, flight-intercept and pan traps, and behavioural
||Marshall et al. 1994; Blades and Maier
||Allows comparison with results elsewhere,
in same or different project
||Direct comparisons of faunas were
favoured by identical protocols among different sites by the same as well as different
||cf. Winchester 1999b
||Labour cost per specimen collected is
||Appropriate design of work in the canopy
(including the use of passive trapping) led to cost-effective sampling
||Winchester, pers. obs.
||Significance of any differences between
sites, treatments, years, etc., can be properly evaluated
||Adequate statistics on variance of samples
of staphylinid beetles, obtained through multiple trap replicates at each site, allowed
real differences between forest habitats to be estimated
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Execution of sampling
||Variation caused by careless procedures is
||Standard written protocols were developed,
and training of technical staff carried out, prior to fieldwork, sorting and preservation
of arthropods from Alberta forest sites
||Alberta Forest Biodiversity project: see http:// www.fmf.ab.ca
|Sorting and preparation
|Sufficient time for proper sorting
||Explicit analysis of time and labour are
made, so that short cuts are not required that will compromise the objectives
||Analysis ensures that sufficient personnel
||Winchester 1999b; and see Table 2
|Sorted and prepared according to
||Identification of as many specimens as
possible is favoured
||Small flies preserved by critical-point
drying can be identified much more rapidly
||Marshall et al. 1994
||Ensures that comparison among samples, and
with projects elsewhere, are valid, and material can be identified
||(see Methods above)
|Specific protocols established early
||Cross-contamination and mislabelling of
samples avoided, quality of specimens maintained
||(see Quality control above, and also see
|Component of study
||Benefit of proper procedure
||Specific example of benefit
|Material prepared and shipped with
appropriate care, and adequate data and context provided
||Identification is assisted
||Proper preparation and shipment and
information about host plants assists identification
|Mass collections supplemented with other
specimens if possible
||Sampling includes site visits by
||(cf. ALAS project: see ALAS website )
||Assistance, reliability and delivery of
||Allowing retention of specimens of
interest by taxonomic specialists and fostering joint subprojects led to relatively rapid
analysis and publication on several groups of arthropods from important ancient forest
||Campbell and Winchester 1993; Coher 1995,
1999; Behan-Pelletier and Winchester 1998; etc.
|Sufficient time allowed
||Project report as complete as possible
||Sampling, analysis and residence at the
same institution for 7 years allowed for continuing publications to profit more fully from
the original sampling
|Tracking of information
||An explicit plan allows all data to be
tracked from the beginning of the project
||Biota database design has provided a means
to assemble and make available a wide range of results
||Communication and comparison are favoured
|Component of study
||Benefit of proper procedure
||Specific example of benefit
||Reference material allows specific
problems to be solved during the project, and also allows findings to be followed up and
used more effectively
||Faunal differences in ancient forests were
verified through material housed at the Pacific Forestry Centre
||References cited above; and see text
|Proper peer review and disemination of
||Information is validated and accessible
||Journal papers and chapters in thematic
publications have shown key aspects of arthropod diversity and habitat associations
||Credible information can be put to wider
||Demonstration and information for the
public (popular magazines, general articles, documentaries, talks, web page) has helped to
foster wider support for the core research and for the conservation needs based on it
ca/~canopy/ , and items listed there
|Interim results clearly identified
||Data are made available quickly, but
results that are not yet definitive are not misapplied, because users are cautioned
||Information on sampling effort helps to
define futher actions; information on the proportion of undescribed species helps to
support further work
||(Chiefly documents of limited circulation,
but see website cited above)
|Ongoing publications as more information
and identifications become available
||Full value drawn from project, not just
||(see Identification - Sufficient time,
Prepared on behalf of the Biological Survey
by H.V. Danks and N.N. Winchester
Published by the Biological Survey of Canada
(Terrestrial Arthropods) 2000. ISBN 0-9692727-9-0