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TAXONOMY
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Overview The
importance of various aspects of taxonomy relative to biological control was
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. A panel of taxonomists,
including Charles Darwin who was a noted taxonomist of barnacles, developed
this code. 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 that 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.
Taxonomists to refine their taxonomic analyses f groups can use this
information. 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. The taxonomist who must rely on preserved specimens, yet they must
be made aware of such differences, cannot easily obtain behavioral
differences between populations. 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 that 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,
scientists whose taxonomic interests originated with their involvement in
applied biological control have conducted a considerable amount of basic
research, particularly with entomophagous forms. 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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REFERENCES: [Additional references may
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