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Ovcerview The
importance of various aspects of taxonomy relative to biological control were
summarized by Gordh & Beardsley (1999). Taxonomy was defined as that
branch of biology which involves the naming, identifying and classifying of
organisms. Previous emphasis had been placed on the importance of taxonomy to
biological control by other researchers (Clausen 1942, Sabrosky 1955,
Schlinger & Doutt 1964, Delucchi 1966, Compere 1969, Gordh 1977, 1982).
For applied biological control workers, there is a need for names for the
natural enemies and hosts that are being deployed. Such names provide an
important mechanism for the dissemination of information. In theory taxonomy
is important to biological control researchers because classification are
developed which are intended to reflect evolutionary relationships. Such
classifications are helpful because they are intended to predict details of
biology and distribution. The
desire to arrange, organize, describe, name and classify is fundamental in
human activity. Such an urge operates at all levels of social organization.
In ancient civilizations names were applied to organisms, and the common
names of many organisms are in widespread usage today. There are however
several problems inherent in common names. Most serious is synonymy.
Frequently more than one common name is applied to a single organism
(synonyms), or the same common name is used for different organisms
(homonyms). Synonymy creates confusion and misunderstanding because the
biological characteristics and habits of similar organisms can differ
greatly. In its earliest form, the scientific name given to an organism was
often impractical. Scientific names during the lifetime of John Ray (Wray)
(1628-1705) consisted of a series of Latin adjectives catenated in such a way
as to describe the animal. The system was less ambiguous than the common name
system, but it was cumbersome because the name of an animal frequently was
several lines or a paragraph long. A
major contribution in the naming of organisms was made by the natural
historian and physician Carl Linnaeus (1707-1778), who is credited with
developing the current binomial system of naming organisms. The start of
zoological nomenclature is taken as the 10th Edition of Linnaeus' monumental
work, Systema Naturae. The notable exception
is the nomenclature of spiders which originates with the work of Karl
Alexander Clerck (1710-1765), Aranei
svecici. The accepted date
of publication of these contributions 1 Jan 1758, and this date is the
official starting point of zoological nomenclature. During
the following century taxonomic zoologists followed the lead of Linnaeus and
prepared descriptions of species for publication but named the animal with a
binomen. The binomen consists of two parts, the generic name and the specific
epithet. With the accumulation of taxonomic descriptions, problems developed
with synonymy, homonymy, the inconsistent application of bionomens, and
related nomenclatural difficulties. The first attempt to address these
problems was the "Strickland Code," prepared in 1846. This code was
developed by a panel of taxonomists, including Charles Darwin who was a noted
taxonomist of barnacles. Subsequently, an International Code of Zoological
Nomenclature was developed in 1906. This code has been altered slightly, but
continues to represent the basic guidelines for the formation and validation
of zoological names for taxa. The most recent revision was published in 1985.
Complicated problems of nomenclature, or matters requiring the fixation of
names in the interest of stability, are referred to the International
Commission of Zoological Nomenclature which serves as a kind of taxonomic
supreme court. Importance of Taxonomy to Biological
Control Danks
(1988) reviewed the importance of taxonomy to entomology. Its importance for
biological control was emphasized by Clausen (1942), and subsequently by
Sabrosky (1955), Schlinger & Doutt (1964), Gordh (1977) and Knutson
(1981). The Scientific Name.--The scientific name of an organism is of utmost importance
(Gordh 1977). It provides a key to the published literature regarding any
zoological taxon and without the correct name the researcher has no access to
knowledge published about an animal of interest. The scientific
name is a kind of shorthand method for conveying an enormous amount of
information about an organism which is available in published literature. All
the information which has been developed about any organism important in
biological control is stored under the scientific name for that organism.
Because of this, the correctness of the name needs to be emphasized. Accurate Identification.--The need for identification is great in biological control,
but the importance of accurate identification is greater. Two species which
are very similar morphologically are not always similar biologically. Subtle
differences in morphology or biology of closely related species can be
profound. Distinguishing between variation in taxonomic characters within a
species and difference in character states between species (individual versus
interspecific variation) is frequently difficult. Understanding the
functional significance of the observed anatomical features which serve to
distinguish between species is an area of research which has lagged behind
orthodox taxonomic studies. Apparent slight anatomical differences may reflect
significant differences in the biology of two organisms. So called minor
structural differences can mean the difference between pest and nonpest
status for species which are potential threats to agriculture, or between
establishment and failure to establish in the case of natural enemies. Some
examples follow: Pink Bollworm, Pectinophora gossypiella
(Saunders).--The gelechiid
genus Pectinophora contains
three described species: P. scutigera, P. endema
and P. gossypiella. Pectinophora
scutigera occurs in Australia,
Papua New Guinea, Micronesia and Hawaii; P.
endema is restricted to
eastern Australia (Common 1958), while P.
gossypiella occurs only in
Western Australia and other world sites. All species consume the flowers,
seeds and seed capsules of Malvaceae. Pectinophora
endema consumes only native Hibiscus in Australia, and is
not an agricultural pest. The remaining species consume other Malvaceae,
including Gossypium spp.
(cotton). Pectinophora gossypiella is one of the most
serious cotton pests, and larvae of this species can diapause within the
seeds of the host plant, which accounts for its widespread distribution. By
contrast, P. scutigera does not diapause
within seeds, is limited in distribution and is not considered a major pest
of cotton. Holdaway
(1926) gave the name of P. scutigera based on larval
differences. Later Holdaway (1929) described the structural characters of the
adult genitalia to separate the species. The validity of P. scutigera
as a species was originally challenged, but is now accepted (Zimmerman 1979). The importance of
correct identification of the bollworms focuses on the pest status of these
insects and quarantine enforcement. In Australia P. scutigera
is not a significant pest of cotton and its distribution is limited by
intrinsic biological characteristics. It does not play a significant role in
quarantine efforts. In contrast, P.
gossypiella is very
pestiferous in cotton. It occurs in the Northern Territory and Western
Australia but not in Queensland. Quarantine serves as an important barrier
restricting movement of this species. Quarantine is expensive to the state
and the commercial enterprise. Coffee Mealybug, Planococcus
kenyae (LePelly).--This insect of Kenya presents an interesting example of
early failure and delayed success in biological control caused by
misidentification of the pest species. The pest first appeared during the
1930's and caused serious losses to coffee in Kenya. First it was identified
as the common, widespread, citrus mealybug, Planococcus citri
(Risso). Later it was determined as a related Philippine species, P. lilacinus (Cockerell). Finally both of these
identifications were shown to be incorrect, but unfortunately, on the basis
of these names, a great amount of effort and expense was devoted to searching
for and shipping natural enemies of Planococcus
in the Asiatic tropics. Parasitoids which appeared promising when collected
could not be established in Kenya. The problem was resolved when the
taxonomist LePelley examined specimens of the pest. He found relatively
inconspicuous but consistent morphological differences which indicated the
that coffee mealybug was an undescribed species, which he then named
(LePelley 1935, 1943). It was then found that this mealybug also occurred in
Uganda and Tanzania where it was under natural biological control.
Parasitoids imported into Kenya from those areas produced complete biological
control. California Red Scale, Aonidiella
aurantii (Maskell).--The California
red scale gives an excellent example of the potential costs of incomplete
taxonomic and biogeographic knowledge of a pest species. This scale is a
member of a complex of species native to the tropics and subtropics of the
Old World (Africa through southeast Asia and the Orient) (McKenzie 1937). It
became a pest of citrus when introduced into the New World without its
associated natural enemies (Compere 1961). Many parasitoids associated with
closely related Aonidiella
species would not attack, or were not effective against A. aurantii.
The failure of early attempts at biological control were due, at least in
part, to the inability to differentiate this species from such closely
related species as A. citrina. Some parasitoids in
the Orient appeared promising to entomologists, but these species failed when
introduced into California because their preferred hosts were other species
of Aonidiella. This was
apparent after Howard McKenzie made a careful revision of the genus Aonidiella and showed that the species
could be separated on the basis of microscopic differences. (also see <bckeys>) Of
equal importance to accurate determination of pest species in biological
control is the correct identification of the entomophagous organisms which
are found in association with target pests and which are being considered for
utilization in biological control. Sometimes such natural enemies belong to
groups of small to minute insects, the species of which often resemble one
another. Taxonomic knowledge needed to differentiate species level taxa in
such groups has accumulated slowly and with great effort. In many groups
knowledge remains incomplete. Some examples of the problems involving natural
enemy taxa important to biological control are as follows: Among
the Aphelinidae, an important family of entomophagous Chalcidoidea, the
genera Aphytis and Marietta appear closely related
on the basis of morphology. Superficially it is difficult to place some species
in the correct genus. Biologically the differences between the genera are
profound. Aphytis species
are primary parasitoids of armored scale insects while Marietta species are hyperparasitoids, usually associated
with armored scale insects or other Coccoidea. Since hyperparasitoids are
viewed as deleterious to biological control, importation or deliberate
movement of Marietta could
adversely affect biological control. The
family Encyrtidae, another large group within the Chalcidoidea, contains a
vast array of genera whose species are primary parasitoids of phytophagous
insects. However the same family also contains genera whose species are
mostly secondary parasitoids (e.g., Cheiloneurus,
Quaylea). Recognition of
these hyperparasitoids and their elimination requires a taxonomic knowledge
of the Encyrtidae. Failure to do so could result in the introduction and
establishment of undesirable species, which is thought to have occurred in a
few cases. A few genera of encyrtids (e.g., Psyllaephagus) contain both primary and secondary
parasitoid species, which demands careful biological and taxonomic study to
separate the beneficial primary and undesirable hyperparasitoids prior to
releases. In
the case of the California red scale, not only did difficulty in distinguishing
the pest from related species retard biological control, but this such was
also encumbered by a lack of knowledge about a very important group of
armored scale parasitoids, the genus Aphytis.
DeBach et al. (1971) showed that this lack of knowledge delayed achievement
of biological control of California red scale by 50 years. Early explorations
for natural enemies revealed the presence of Aphytis parasitoids at several localities in the Orient.
Specimens from these collections were determined as Aphytis chrysomphali
Mercet, a species already present in California which was not especially
effective. Therefore, no effort was made to propagate and release new
oriental Aphytis until after
World War II (Compere 1961). The two most effective natural enemies of red
scale, Aphytis lingnanensis Compere and A. melinus DeBach, were not recognized as distinct species
until 1948 and 1956, respectively. These species might have been introduced
into California many years earlier had a proper understanding of the taxonomy
of Aphytis existed..
Similarly, Aphytis holoxanthus DeBach, the most
effective parasitoid of Florida red scale, Chrysomphalus aonidum
(L.), apparently was first collected around 1900, but was ignored because it
was confused with another species. Aphytis
holoxanthus was made
available for biological control in 1960 when DeBach recognized it as a
distinct species (DeBach et al. 1971). Trichogramma is a cosmopolitan genus of tiny parasitoids which occur
as more than 120 species. All species for which the biology is known develop
as primary internal parasitoids of eggs. Trichogramma
has been used extensively against lepidopterous pests in classical biological
control or inundative release programs. Some programs have produced
contradictory results, with some workers
claiming success and others admitting failure. Poor taxonomic knowledge has
contributed to conflicting assessments. Early researchers rarely deposited
voucher specimens for their research and without material to compare it was difficult
or in some instances impossible to determine what species of Trichogramma was used in a
release program. In one example, most references to Trichogramma minutum
Riley, T. evanescens Westwood and T. semifumatum (Perkins) made prior to 1980 probably are in
error. It is now known that Trichogramma
contains many anatomically similar species which can be distinguished only by
microscopic differences on antennae and genitalia. Traditional reliance on
body coloration is if limited utility and has been shown to depend on
environmentally induced variation. Many species display dark coloration at
the base of the forewings, and the name T.
semifumatum was often
applied to such forms. The latter species is now recognized as endemic to the
Hawaiian Islands based on one collection (Pinto et al. 1978). Biological Control Contributions to
Taxonomy There
exists an element of reciprocity between the biological control worker and
taxonomist which must be fully developed to maximize the usefulness of
taxonomy as an adjunct to biological control. Biological control workers can
offer taxonomists important data necessary to complete taxonomic
identifications. The kinds of important information include zoogeographical,
biological, behavioral ecological and hybridizational data. Zoogeographical Data.--Biological control researchers frequently engage in time
consuming and expensive foreign exploration. Often the results of this work
are not published and the imported material is not studied. Such material can
provide potentially important data for taxonomic studies in terms of
understanding geographical variation and expanding known limits of
distribution. Biological Data.--Because it is believed
that there are trends toward habitat specialization and host specificity in
many groups of parasitic Hymenoptera, data on host range and host preference
can be obtained in the field and in the insectary. This information can be
used by taxonomists to refine their taxonomic analyses f groups. Also,
information on pest species, such as host plant preferences, can be shared
with specialists. Behavioral Data.--Subtle differences in behavior between populations of what
appears to be one species may point to taxonomic differences between two or
more closely related species. Behavioral differences between populations
cannot be easily obtained by the taxonomist who must rely on preserved
specimens, yet they must be made aware of such differences. Once behavioral
differences are known, the taxonomist may find encouragement to search more
for minor anatomical differences which can be used to distinguish between
closely related taxa. The
kinds of important behavioral differences are many. For example, courtship
behavior in Aphytis appears
to be controlled primarily by species specific sex pheromones released by
virgin females. Males are attracted to the pheromone released by conspecific
females. Also, males produce a pheromone which appears to calm the virgin female
and render her sexually receptive. Males and females do not normally respond
to members of the opposite sex belonging to other, even closely related
species (Rosen & DeBach 1979). Additionally, other kinds of behavior,
such as host finding, may also be indicative of taxonomic difference between
populations which show no readily apparent anatomical differences. Ecological Data.--Closely related
species often differ substantially in their ecological requirements.
Important data must be kept on the ecological associations of entomophagous
arthropods collected for biological control purposes. Factors such as
elevations and season are important, but less apparent ecological data, such
as the type of plant community in which the species occurs, can also provide
valuable clues to the taxonomist who is attempting to differentiate similar
forms. Host specificity among related species of parasitic Hymenoptera is
often reflected in their association with specific plants which harbor their
insect hosts. Thus, information on the plant hosts on which parasitoids are
collected may prove useful to taxonomists. Hybridization Studies.--Most
classical taxonomists do not have access to insect rearing facilities, and as
a consequence these taxonomists are restricted in their ability to test
reproductive compatibility and to make judgements involving the biological
species concepts. While most museum taxonomists would acknowledge
reproductive compatibility as a viable approach to the study of species
limits, in reality they are limited to conceptual acknowledgment only.
Biological control researchers with access to laboratory and insectary
facilities are able to provide detailed information regarding reproductive
compatibility and reproductive isolation. This kind of information is
important as is illustrated in such groups as Trichogramma (Pinto et al. 1986). Sources of Taxonomic Expertise It
is often difficult to find specialists sufficiently expert in the taxonomy of
pests and natural enemies who are willing to provide biological control
workers with the unequivocal identifications required. This has been
especially true for groups of minute parasitoids that are of major
importance. Dwindling public support for natural history museums and for
taxonomic research in general has intensified this problem since the 1960's.
Many biological control specialists have been required to undertake
systematic research in an effort to solve taxonomic problems associated with
their own research. Thus, a considerable amount of basic research, particularly
with entomophagous forms, has been conducted by scientists whose taxonomic
interests originated with their involvement in applied biological control. An
example is the detailed study of the aphelinid genus Aphytis by Rosen & DeBach (1979). As a result, Aphytis now is recognized as
among the best understood genera of Hymenoptera used in biological control.
Similarly biosystematic studies by Dr. E. R. Oatman and colleagues have
elucidated Trichogramma in
the 1980's, and work with Muscidifurax
by E. F. Legner has shown great diversity in a group that was previously
regarded as monotypic. [ Please refer to Research ] Directories of
taxonomic specialists are published periodically (e.g., Blackwelder & Blackwelder
1961), and although helpful, they are quickly outdated. An effective method
of locating taxonomic expertise is by consulting the most recent -volumes of
the Zoological Record. Word of mouth approach is very effective also. Principal Groups of Natural Enemies Please see <keys.htm> and Groups for keys to
families of entomophagous arthropods, and details of principal families. |
|
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