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TAXONOMY OF NATURAL
ENEMIES ----CLICK on underlined categories to
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Matter, depress Ctrl/F ]: Importance of Taxonomy to Biological
Control Biological Control Contributions to
Taxonomy Sources of Taxonomic Expertise Principal Groups of Natural Enemies 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. 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. 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
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