Different Types of Variation Within a single population:

In case of Identification, we take morphological characters which are attributes of Genetics. Genes play
their role in these characters and each and every character is the outcome of gene. We identify the species
from these characters which vary from species to species, thus character variation is helpful in the
identification of specimen.
If these variations are permanent and native then these are called Genetic variation. But contain characters
are not permanent in nature and they may be due to one or other reasons and these variations may be
misleading in the identification of species. So, knowledge of these variations is important for identification.
Intrapopulation Variations:
These variations may be genetic or non-genetic.
Note: Keep in your mind that all these variations are found in a single population. Among different
populations variations already exist.

Types of Variations:
There are two types of variations
1. Genetic Variations
2. Non-Genetic Variations

Non-Genetic Variations:
Individual Variations:
It includes the following
 Age Variations
 Seasonal Variations in an individual
 Seasonal Variations in consecutive generations
 Social Variations
 Ecological Variations

Ecological Variations:
 Habitat Variations
 Variations induced by Temporary Climatic Conditions
 Host determined variations
 Density dependent variations
 Allometric variations (Heterogenic variations).
 Neurogenic/Neurohumoral variation.
 Traumatic Variations;
 Parasite induced variations.
 Accidental and Teratological Variations

Genetic Variations:
 Sexual Dimorphism
 Primary or secondary sex differences.
 Gynandromorphs (A gynandromorph is an organism that contains both male and female
characteristics.)
 Intersexes
 Reproductively Different Generations
 Sexual and uniparental Strains.
  Ordinary genetic variations (Discontinuous variations also called Genetic Polymorphism).
 Continuous Variations.

Details:
Non-Genetic variations, variations in which genes are not involved in determining the character.
Individual Variations
Age Variation: In case of insect eggs are not laid at a time and do not hatch at a time. Some individuals
come out at early stage as compared to others, so there is a difference of age, may be of size. These insects
pass through a series of Juvenile and Larval Stage, in that case they are totally different from the adults.
Younger age may be different from adult age, so while identifying a specimen we should consider the age
variations e.g. In Grasshoppers young ones are just like adults but in adults’ wings and genetalia are
developed which are absent in young ones. In Lepidoptera identification also takes place even at larval
stage. Keys are also available at larval stage.
Age variation is mostly important in case of parasites.
Seasonal Variations in an Individual: It is in case of those adults which live on the adults through several
breeding seasons. Some adults may have different colors during different seasons. In summer color of adult
may be faint while in spring it may have bright color.
 Butterflies in Spring Season have bright color while in summer they have faded color.
 Red spider Moth also have different. In normal it is reddish, but in winter they will become almost
blackish or dark green. It is mostly in birds during breading season color of feathers become bright
and in off season it is faded.
Seasonal Variations in Consecutive Generations: It is specially in case of insects because insects have
many generations in one year. Insects which hatch out in cold weather, they have different appearances
than those in Summer dry Season. It is case of that species that are short-lived and have many generations
in one year and each generation in a season may have different appearance. Butterflies of cold season have
different appearances than those of dry season.
Cyclo-morphosis: A seasonal Non-Genetic change in Phenotype.
Social Variation: These are variations which occur in social insects. Social insects like Hylomorphisms
have different casts. In Honeybee there are Queen and other castes. Castes are of different forms and shapes.
Individuals of the same species have different morphological appearances. There are definite groups of
Individuals in a colony but they are of different shapes and castes. These differences are called social
variations. In Hymenoptera different casts are present, they may be present in both sexes and these different
appearances occur due to the differences in larval food e.g., larvae which are fed on the fleshy food such as
royal jelly they develop into Queen and larvae which are fed on the pollen grains or nectar are developed
into worker bees. Such variations are also called as Intercolonial Variations.
Ecological Variations:
 Habitat Variations: Single species occur in different habitats, but in same region, they are often
visibly different.
Note: In this case there are certain difference between taxonomists. Some taxonomists say that spp at one
end of habitat should consider a separate that at the other end. While another taxonomist says that spp same
at both ends of habitat.
 Ecological variations are Non-Genetic Variations. Different authors have given them different
name, like micro sub species, Ecological Races, or non-genetic Eco phenotypes.
 Variations Induced by Temporary Climatic Conditions: Usually, small organisms are affected
mostly due to Temperature and Growth become stunted. These changes are for a given time period.
These changes may be Phenetics.
 Host Determined Variations: These variations are mostly found in the parasites of Plants and
Animals. Plants Parasites are all the sucking insects. These parasites are a source of trouble for
taxonomist because they permit confusion. Insects collected from one host may have different size
 and color from those insects which are collected from other hosts. These changes are related with
host and they provide a very potential source of taxonomy and often it is confused with
Microgeographic races (Sympatric Spp). These changes are usually expressed in Size Differences
but may also involve certain other Morphological or Physiological Characters. Now these host
determined Variations are not common. In Recent literature such species are referred as Sibling
Species’. There are Scale Insects such as Lecaniumcorni spp. They feed on Plant Prunus Spp (Spp
of Berries). These insects feed on both these plants. Those insects which feed Prunus spp have
larger body and shorter appendages while those which feed on Photinia spp have smaller body and
larger appendages, if host is changed then result will also be changed. Braconid wasp is a larval
Parasite belong to order Hymenoptera. When this spp was recorded on Blue Green Caterpillar of
Colias spp, one pupation parasite gives white cocoons. But when this parasite feed on Yellow Green
Caterpillar then resulting cocoons were Golden in color. Such Variations are not Permanent and
are Host Determined /Non-Genetic.
 Density Dependent Variations: (Density: No of Specimen /Individuals in a population.)
Whenever the population of species increases overcrowding takes place and such circumstances
reflected in and on Morphological Characters e.g., Locust, there are two phases in Locust Solitary
and Gregarious. When newly hatched nymphs are reared under isolated conditions, they give
pinkish Solitary Phase and when newly hatched nymphs are reared in crowded conditions, they
give Yellow Gregarious Phase. When these newly hatched nymphs are reared under less crowded
condition, they give rise to Transitional/Intermediate Phase. When Gregarious phase and pink color
nymphs are mixed the body, color is changed and is intermediate. These phases are different In
Anatomy, Color, and Behavior.
 Allometric Variations (Heterogenic Variations): These are the variations of measurements. It is
the growth of disapropriate size of a structure of a body or body part in relation to the rest of the
body. In insects it is very common e.g., in some Ants the Head becomes Enlarge as compared to
the rest of the body. In Stag Beetle mandible developed in disapropriate ratio i.e., one is large and
other is small. In Scrab beetle thorax o frontal horns developed in disapropriate manner. Some
segments of antenna grow more as the development of antenna in Thrips is also an example of
Allometric Variation.
 Neurogenic/Neurohormonal Variation: These variations refer to the color changes in animals
always in response to environment. In Insects they are color forming/pigmented bodies called
Chromatophores. They are affected by the environment. Dispersion and concentration of
Chromatophores may change the color. If Dispersal of chromatophore occur, the color will faint.
If concentration of Chromatophore occur color will be marked. In lower animals it is very less but
in higher it is very marked. In cold blooded vertebrates (Amphibian and Reptiles) this thing is well
developed.
Note: From Taxonomic point of view body color is not taken as character and it is not considered as a base
for identification. It has minor value.
Genetic Variation
Intrinsic/Inherited Variations: These variations are through the genes. These are genetically based
variations. These may be divided into the followings.
Sex Associated Variations: These are variations which are genetically determined within a population.
These may be sex limited or either sex associated Variations. These are expressed in one sex only. Primary
sex associated or Primary Sex differences involve primary sex organs which are utilized in reproduction
e.g., in case of Gonads, Ovary, Genetalia otherwise both these sexes are quite similar. Secondary Sex
difference or Secondary Sex Associated are more or less pronounced. In sexual dimorphism males are
totally different from Females. They should Not be misunderstood. They are very striking in certain groups
e.g., Hymenoptera the individual of one sex is very different from the other sex. In mango mealy bug and
African ants and Embiopteran Web-Spinner.
Alternation of Generation: It is also known as reproductively different generations. In many Insects
different generations exhibit different morphological Characters e.g. A genus of wasp is Cynics in this
genus one generation is different from bisexual generations. In aphids a gametic generation produce
wingless female but when this female reproduces through fertilization it will produce winged females.
Gynandromorph: They are such Individuals which show a male character in one part of the body and
female character in second part of the body. A single specimen exhibiting the characters of both sexes, one
half body part has one sex character and other half body part have other sex character. In some insect’s sex
characters may be scattered on the body in mosaic form. In such case Gynandromorphs are identified and
not declared as either male or female e.g., Papilio (Genus of Butterflies).
Intersexes: These are the Individuals which exhibit the blending of characters of both male and female in
scattered form. It is the outcome of imbalance in male female tendency gene or due to irregular fertilization
or due to upset in mitosis, due to certain external stimuli such as Physiological Change due to Parasitism.
Non-Sex Associated Variation: These are intrapopulation Variation and do not involve primary sex
characters. It is of two types Continuous Variations and Discontinuous Variations.
 Continuous Variations: These are most common type of individual variations which occur due to
slight genetic differences between individuals. No two individuals alike except twins
(Monozygotic). The study of these variations is one of the most primary tasks of a taxonomist
because no one individual is typical of the population. The variability of different characters of
some population is very different and different degrees of variability among related species. There
should be no lastly decision in the respect of variability of related species.
 Discontinuous Variation/Genetic Polymorphism: In certain species the number of a population
can be grouped into definite classes, determined by the presence of certain conspicuous characters.
Such discontinuous variation is termed as Polymorphism. And they are controlled by a single gene
transmitted by Mendalian Inheritance e.g. The spotting in lady bird beetle is an example of
Polymorphism. It has great Biological Importance since it provide selective difference between
apparently natural Characters. They are genetically different seasonal forms, the selection is very
strong e.g., Summer Generation different than Winter Generation e.g., Drosophila.
Sex Limited Polymorphism: Male is one form female is of different form in different season i.e.,
genetically different forms. Causes of this form found in Lepidoptera, Alfalfa, Butterflies has two striking
different female forms. One form resemble male in its brown color and second form is purely white.

Taxonomic characters:
The essence of original taxonomic characters is the analysis of material and synthesis of results
into a
classification. These steps although combined but they include two separate operations.
A. Finding and evaluating differences.
B. Discovering points of resemblance.
In both cases we deal with certain attributes of organism known as taxonomic characters.
The taxonomic characters may be defined as “Any attribute of an organism or group of organisms
by which it differs from an organism belonging to a different taxonomic category or resemble an
organism
belonging to same category”. The taxonomic characters are the attributes which permit placement
of an organism in the formal classification. Taxonomic characters have dual functions.
 A diagnostic aspect as indicator of differences (in lower category)
 Function as indicator of relationship (in higher categories)
Difference between the organisms of the same category i.e., Male versus female, immature versus
adult are not taxonomic characters.
Diagnostic Value of Taxonomic characters:
According to a recent estimate a higher animal may have 10,000 genes while the numbers of
characteristics are limited by the patience of investigator. Even two related species of same genus
may differ in 400-600 characters including all sort of physiological, ecological and psychological
reaction norms such as difference in instinctive behaviour. It would require a life time to prepare
an exhaustive species description with reference to all these characters. For the study of an
exhaustive species so many characters are not taken into consideration. These characters are
known as diagnostic characters. The most practical diagnostic character mostly relates to easily
visible structures. These characters many not be important for species but they serve as marker for
taxonomist. Very true diagnostic character or taxonomic character must be constant one for the
members of that category in case of certain variation. They must be constant for certain %age of a
population of a category.
KINDS OF TAXONOMIC CHARACTERS
Earlier taxonomists and even today many taxonomists use the morphological characters. No doubt
these characters are still in use but they are being supplemented by other kind of characters, which
have increased the reliability of classification. Each character has its own importance, again it
depends upon the taxonomist, to which character he gives the more importance. Classification
bases have increased by using all stages of life cycle of spp. i.e., characters of male and female
and immature stages. In certain cases, characters of immature stages are more important than adult
characters’ such as in species of anopheles. There is a complex of Anophelesmaculipennis. This
complex is identified on the basis of egg characters. In white fly characters of pupal stages are
important and species are identified on pupal bases. These characters may be grouped under five
categories.
1-MORPHOLOGICAL CHARACTERS
 General external morphology
 Special structure of body e.g., genitalia
 Internal morphology(anatomy)
 Embryology
 Karyology and other cytological differences.
2-PHYSIOLOGICAL CHARACTERS
 Metabolic factors
 Body secretions
 Genic sterility factors
 Serological, protein and other biochemical differences
3-MOLECULAR CHARACTERS
 Immunological differences
 Amino acids sequence of protein
 DNA hybridization
 DNA and RNA sequences
 Endonuclease analysis
 Other molecular differences
Molecular characters are latest technique for species identification.
4-ETHOLOGICAL CHARACTERS
 Courtship and other ethological isolating mechanisms
 Other behavior pattern.

5-ECOLOGICAL CHARACTERS
 Habitat of host
 Parasites
 Host reaction
 Seasonal variation
6-GEOGRAPHICAL CHARACTERS
 General bio geographical distribution/pattern
 Sympatric allopatric relationship of population

IN DETAIL STUDY
1. MORPHOLOGICAL CHARACTERS

1-EXTERNAL MORPHOLOGY:
Feature of external body pattern. These features vary according to kind of animals. It may be
feature of antennae, wing and thorax. These feature result of genetics, but we call them each
genotypic character as genotype. So, these characters may be very well studied and analyses by
using latest technique.
 Coloration of body, they provide diagnostic characters e.g., Butterflies and wasp spp. Have
different color for identification.
2. Special structure on body e.g., genitalia
Genitalia is very important character for separation of species, because while defining a species
we say reproductive isolation. This reproductive isolation will be difference of genitalia. It is
important for confirmatory test for species identification. Because copulatory organ has definite
structure. No other species will be able to copulate with other species due to different structure of
genetalia.
Why we supplement these morphological characters?
Morphology reflects only part of genotype and does not affect genetic relationship accurately.
Morphology in many taxa do not supply sufficient characters
Characters or any character may mislead because of special adoption.
Due to these reasons, we use other characters or supplemental characters.

3. Internal Morphology (Anatomy)

It provides numerous characters specially in case of higher groups, these internal structures of
head, thorax. Thy provide abundant characters.
4. EMBRYOLOGY:
It is the science of Egg. Especially comparative embryology in case of egg, Cleavage Pattern and
Gastrulation and other phenomenon of eggs are very important in case of higher categories.
 Cleavage Pattern is very important to differentiate between apterygote and
pterygote insects. Other characters of larval stages are also considered.
 In case of Anophelesmaculipenisis identified on the basis of egg characters, in this case
egg character is important to differentiate between sibling species or aleroididae.
  In case of Whitefly (Alerodidae) their taxonomy is based on pupal stage.
 In Lepidoptera larval stage is important for taxonomic delimitation.
5: Karyology and other Cytological Characters
Karyology: It is science of study of number of chromosomes.
 Characters involving chromosomal structure and number are called Karyological
characters.
 Number of Chromosomes varies in every species.
Cytology: It is the study of cell. If number of chromosomes do not compatible then there is
infertility.

2: PHYSIOLOGICAL CHARACTERS:
It is the study of function of different body organs. Physical characters initially have not been used
or unevenly used in taxonomy, otherwise these characters exceed morphological characters in their
Constancy and diversity.
 The disadvantage of these characters is that for the study of physiological characters we
need living species.
 Living species must have shot life cycle.
 The most suitable organism must be smaller in size.
 They should be easy to rear.
Note: It is fact that physiological character cannot be taken for taxonomic character.
Metabolic Factors:
These characters important for microbiologist as well as for bacteriologist as they do not have
enough morphological characters e.g., Enzyme activity is different in different bacteria.
Catabolism + Anabolism= Metabolism
Cell chemistry is also different in different in different species. So, these things are important for
microbiologist. Bacteria may be aerobic or anaerobic depend upon metabolism.
2: Serological and Protein and Other biochemical differences:
Serology: It is the study of physiological characters. It is a science which deals with interaction of
proteins, antigens and antibodies.
Antigen is the substance which when introduced in a body of living organism then they are capable
of producing antibodies of most introduced antigen. Vaccine is also made by using this principal
of protein reaction and biochemical differences getting ground base in taxonomic identification
3: Body secretions:
Body secretions are very consistent pattern for taxonomy. In insect’s waxy secretion. Every species
will have its own specific pattern of waxy secretion.
In coccids special waxy secretions, special glands secrete out certain waxy secretion.
Scales are also secretion of certain gland (mealy bug)
Different body secretion pattern is due to the fact that underlined area morphology pattern is
different.
4: Genic sterility factor:
Different species cannot interbreed this is sterility and these days it is considered as an isolating
mechanism in species. Taxonomically distant species are totally infertile while close species may
interbreed, but their offspring cannot interbreed e.g., Drosophila; it has large no. of species.

3-MOLECULAR CHARACTERS:
  Immunological differences
 Amino acid sequences of protein
 DNA hybridization
 DNA and RNA sequences
 Endonuclease analysis
 Other molecular differences
Serology was the first and earliest method of comparing protein, basic factor that protein of one
organism will show stronger antibody reaction to protein of closely related organism rather than
those which are distantly related. After that the recent work on enzyme or different body structures,
in that case chromatography is used, and that chromatography was known as Diffusion
chromatography. In this technique we take body fluids of different body parts which travel across
the filter paper when different organisms are treated on filter paper. Nowadays this
chromatography is replaced by electrophoresis in which we use Electric potential + hydrolyzed
charge to separate the fluids.
DNA & RNA Study
Large number of molecular techniques has been used. These techniques are proteins, nuclear DNA
or mitochondrial DNA, ribosomal nucleic acid techniques. In taxonomy the main purpose of
taxonomists is to determine overall similarities between species. Ribosomal RNA is very useful
for understanding the relationship between eukaryotes and prokaryotes. Studying DNA have
hybridization, the radio tanging (isotopes) techniques are used. When you mix the two DNA, one
treated and other is not treated and then hybridization techniques are employed, the separate stands
of both DNA are obtained. Similar one five hybridization reaction “it is called DNA sequences.
This level of taxonomy is known as molecular taxonomy .it is the latest technique.

4-BEHAVIOURAL CHARACTER:
These characters are also known as ethological characters. Ethology it is the science of the study
of behavior of the organism. Behavior varies from organism to organism and from genus to genus.
Courtship behavior and other isolating mechanism
Mating habits are different from group to group and organism to organism, and they provide very
important characters because they result in reproductive isolation. Their genital characters and
other sex characters and body secretions are different. They give isolating mechanism.
Other behavior patterns
They include other living patterns e.g.; each species of spider has different patterns of web making
and this pattern is used for isolating species. Similarly nest building behavior and pupation
behavior is different in different organism.

5-ECOLOGICAL CHARACTERS:
 Habitat of host
 Parasites
 Host reaction
 Seasonal variation
Habitat:
Surrounding or living area of organism. Each species of animal has its own range of tolerance
habitats. Each species has its own breeding season. There is a rule relating to ecological characters
and habitat.
GAUSES RULE: It states that no two species can coexist in same place because each species or
category has its own separate adaptive zone. That’s why on the basis of ecology we can define
them. Ecological characters are of great value for distinguishing sibling species. Sibling species
are mostly found on border areas.
E.g., genus of cricket Nemobius.
Genus of mosquito Anopheles.
They can be recognized on the basis their songs and ecological habitats. Similarly, anopheles are
difficult to identify morphologically. They are identified on the basis of habitat because some
species are found or live only in rice field. Some species are found or live-in standing water. Some
species are found in cool running water.
FOOD (Parasitic relation with host):
Different organisms have their own food preferences and on the basis of that they can be identified.
Bark beetle initially they have a few species but Similarly, when they are studied on the basis of
host plants it was found many species. Each species with definite constant characters boring pattern
and host preferences are different. Food also provides taxonomic characters.
Host parasite reaction
The knowledge of parasite is very useful for taxonomists because it can be used to differentiate
between groups, it is called host discrimination.
The method of using differences in parasites to distinguish between host is called host
discrimination e.g., two similar species of termites when their host were studied, they were found
distinct because trematopid beetle were found from the nest of the termites.
Parasitic discrimination
Discriminatory parasites are presumed to discriminate (i.e., re-enter the host search stage)
whenever they encounter the alternative host species for which they have lower fitness, therefore
searching until they either encounter the other superior host species or ultimately fail to do so e.g.,
bark beetle attacking Pinus because each species of Pinus is attacked by a definite species of beetle.
Seasonal variation:
Different organism appears only in definite season. On the basis of that we can identified definite
species.

6-GEOGRAPHICAL CHARACTERS:
They are very important for clearing the confusion in taxonomy because geography has its own
impact on species formation. These characters are of two types
General biogeographic patterns:
Biogeographer have divided the world into different geographic regions and they will be confined
to these geographic regions. Each region has its own fauna and flora. These regions are not rigidly
defined but they represent the distributional center. So, for clear cut identification of species a
taxonomist should have understanding of geographical history of these regions.
Sympatric allopatric relationship
It is very useful tool for solving the species or population patterns, whether two populations are
distinct species or not e.g., these days a chain or ring of forms which are known as allopatric
species is considered as indistinctive of polytypic species but on the other side it two or more
forms, they do not represent the allopatric formation. Then species are said to indicate independent
full species. Taxonomy is based on variation is different character.
Allopatric: a term applies to two or more populations which occupy mutually exclusive (but
usually adjacent) geographical areas.
Sympatric: a term applies to two or more populations which occupy identical or broadly
overlapping geographical areas.
Allopatric speciation: species formation during geographic isolation.
Sympatric speciation: species formation in the absence of geographic isolation.
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malaria crewsby nonscientists.
based their During
control
facility by
various choices following the numbers, which indicate the path
the that bperations on the results of field identifications of anopheline mosquito
purpose
divisionsof a key.follow.
Its mainThisdisudvantage
is the type which
is thatbestthefufills diagnof
relationship arvae (ie. 2. The fact that eritical characters were ilustrated
vell as deseribd mude theas well keys asusable by such personsPictorial as medical
as
previously isusednotisapparent
as followsto the eye. An eample bssed on the same ( rorpsmen
have been andemployed
engineers also in field by entomologists.
guides to vertebrates and key
flowering
plants.
usms.eonves
Failurei nenta
to impartpieturs
suechdtakey.w inuer-dlationsbip
picture is dtthe achief
rop ot oreanof
the traditional dichotomous Thisatisa gdance
not a serious defetdefect
for the
 L6G TAXONOMIC PROCEDURE PRESENTATION OF THE FINDINGS Lov

specialist who is accustomed to using and interpreting keys, but it is a4 in the groups listed in Table 13, smithi-ruficornis-flavicornis-californice
species
shortcoming from the viewpoint of the nonspecialist. To overcome this 4 may be a group that splits off very early from the other four
difficulty, three different types of key have been devised: the branching } (completa, emarginala, rufipes, and nigripes), the visible difference between
type (Fig. 30), the box type (Fig. 32), and the cireular type (Fig. 31). the groups may be ill defined (wings clear va. opaque}. To use such an
:
CALIFOASICA = FPLAVIGORNIS
EMARGINATA RUFIPES WIGRIPES SMITH RUF IGOR MIS
SOMPLETA
eyes legs legs omlennmnie ontencoe antennge antennae
eyes
red block black red ie aw
entire emorngingte
| 7

wn he STEAL
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Antennae Antennae Torsal
“Test Mert reste heirh
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) (No
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Fie. 30. Example of branching key based on analysis of characters in Table 13.
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"Eon dehamg = mbally cowie,
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| avies bolt WS) Geakih rearstee cf owmervis US) 0 Portier Ul gts Wit uel ee
ee a ee Pai tenet]
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ene be MALAMLA CONTROL IM WAR AREAS
© eee Mécatified fies Frestoes ond Garberg by Hl Haute US. PUBLIC HALTS apence

Fra, on). Pictorial key to lary ac of anopheline mosquitoes of the United States (08.

Public Health Service).

The data of Table 13 are used in each case, so that the types are
comparable, Example of circular key based on analysis of characters in Table 13.
Fie. 31,
Certain keys such as the indented (also the box-type and circular
type) are sometimes referred to as phylogenetic keys. Such an appella- unreliable character as the very first bracket might lead to many mis-
tion makes the silent assumption that the series of dichotomies chogen identifications, Furthermore, subsequent discovery of additional char-
parallels the phylogenctic history, The taxonomic record in well- acters may show that actually the form of the antennae is a more deep-
known groups has shown how casy it is to make mistakes in the interpre- seated character than the wings, and this would lead to a different
interpretation of the phylogeny. Finally, even the bracket-type key
tation of the phylogenetic value of characters. For instance, although
 16a PRESENTATION OF THE FINDINGS 169
PAXONOMIC PROCEDURE

can be constructed in such a way as to group together the most closely One of the first steps in a phylogenetic study is usually the tabulation
related forms, except that monotypic dichotomies sometimes have to be. of the characters shared by the groups concerned. Not only structural
placed out of order, features should be tabulated, but also biological, embryological, physio-
' logical, and geographical data to the extent to which they are available.
The Presentation of Phylogeny. Natural classifications are based on
phylogeny, although they can represent phylogeny only imperfectly; The second step consists in determining which of the tabulated char-
(Chap. 3). The great interest in phylogeny goes back to the early acters are primitive and which specialized. This often requires reference
Darwinian days. Darwin (1859) stated that all animals and plants wera to characters in related groups which fall outside of the study. Reduc-
derived from common ancestors, but he made no attempt to reconstru tion (e.g., loss of wings, fewer segments in appendages, etc.) is normally,
ct but not always, an indication of specialization. Narrowly adaptive
the genealogy of species and higher categories. It was Haeckel (1866)
who made a first attempt at presenting the relationships of all animals: characters which restrict or limit the habits of a species or group are
phylogenetically. A phylogeny is traditionally represented by a branch- usually specializations.
Since the more primitive species or groups are likely to retain the most
primitive characters, it is important to know where the most primitive
£ e
o = “t
| _ 2 s
forms are apt to be found. Here geographical distribution and habits
i E = a = “ £
=
aid greatly. New Zealand and Australia, and to a lesser degree South
a om = c = 3 o aa
= 5 3 = E _ = = America, are great reservoirs of primitive types. Outside of these areas
a E r 2 wa 3 a a
primitive groups may be widely but discontinuously distributed, fre-
‘ad a La] fo

Eyes
¥
Eyes
emargi- Legs | Legs | Antennoe| Antennae] Antennae Antennae
quently with highly localized, only distantly related, species. When the
entire | nate red | block black red Slack Fyellow
ptimitive groups have been located and the primitive characters recog-
nized, a rough approximation of the relative ages of the groups con-
Antennge Antennae Tarsal Tersal
sérrote
cerned is possible. Fossil evidence, when available, may aid greatly in
filiform segments segments
tin@ar bilobed confirming such conclusions. With many animal groups, however, only
Wings opaque Wings cleor limited help is normally available from this source.
Fig. 32. Example of box-type key based on analysia of characters in Table 13. Phylogenetic reasoning on the basis of degree of resemblance is con-
fused by several natural consequences of evolution. The first is con-
ing tree, somewhat as in human genealogies.
vergence due to adaptations to similar environmental conditions.
Ever since the invention
of the phylogenetic tree by Haeckel, it has been customary among taxon- Familiar examples are the distantly related but similar-appearing families
omists to express phylogenetic conclusions in the form of diagrams ‘ of water beetles, which possess a common streamlined form; the strikingly
(Jepsen, 1944). In spite of their numerous shortcomings, such diagrams similar structure of the forelegs in mantids (Mantodea) and mantispids
are useful summarizations (Neuroptera); and the superficially similar ectoparasites of vertebrates,
of taxonomic knowledge and provide a pic-
torial representation of the author’s concept of the evolutionary history which belong to at least six different orders of insects.
of a group. Often a simple diagram shows more than many ‘pages of | Second, phylogeny may be obscured by parallelism. The various
detailed discussion or description. species of Drosophila, for example, show similar mutations, such as orange
Some of the more useful kinds of —
diagram are discussed below, eye. Consequently, orange-eye variants in Drosophila are not mono-
: phyletic but cut across phylogenetic lines. They are part of the genetic
Phylogenetic Evidence. Before diagramming can be attempted, an
interpretation of the probable phylogeny must be reached on the basis pattern of the group asa whole, The same phenomenon is evident in the
white females of the various species of Colias, Mayr and Vaurie (1948)
of the taxonomic data. It is here that the systematist must muster
all his judgment and experience. have given examples of such characters in certain birds, and Michener
Because of the subjective nature of the
problem, it is difficult to lay down any hard and fast procedures for (1949) in eaturniid moths. Michener concluded that in the Saturniidae
attaining satisfactory results. a hind tibial spur has been independently lost at least 10 times in one
As has been remarked by Simpson subfamily, the epiphyses of the female at least 10 times in the family,
(1945), “Phylogeny cannot be observed. It is necessarily an inference
from observations that bear on it, sometimes rather distantly, and that and the articulation of the male genital harpes 7 times, He found that
reduction of the labial palpi has oceurred at least 4 times, and of the eves
can usually be interpreted in more than one way.”
See discussions, stats, and author profiles for this publication at: https://www.researchgate.net/publication/275891371

Insect Taxonomy—Basics to Barcoding

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 Insect Taxonomy—Basics to
Barcoding

K. Sreedevi, Naresh Meshram and P. R. Shashank

Abstract
The integration of new knowledge and methods of population biology,
phylogenetics, and other evolutionary disciplines into taxonomy is war-
ranted (Sites and Marshall, Trends Ecol Evol 18:462–470, 2004). The
analysis and interpretation of data used to delimit species have profound
implications in taxonomic research. Integrative taxonomy gives priority
to species delineation over the creation of new species names. The inte-
gration of all possible taxonomic approaches abridging the gaps of each
in arriving at correct species delimitation is the need of the hour in the
light of biodiversity inventory. Taxonomy needs to be pluralistic to im-
prove species discovery and description, and to develop novel protocols to
produce the much-needed inventory of life in a reasonable time. Insects,
being vast and diverse on earth , need much more integrative taxonomic
attention than other life systems. The unique characters of an organism
that unravels the diagnostic character differences that delimit the species
have to be assessed holistically.

Keywords 

Evolutionary disciplines · Holistic approach · Population biology ·

Phylogenetics

Introduction ages to the modern era. However, the taxonomic

studies originated in the eighteenth century with
Insects, ever since their appearance 350 million Carolous Linneaus work on Systema Naturae,
years ago, are the dominant species in the biot- first published in 1735. Aristotle was the first to
ic community widespread in all habitats of the recognize the hierarchical pattern in the diversity
earth. The various studies of insects prevail from of animals but the scientific method of classifica-
tion was put forth by Linneaus followed by work-
ers like Latreille, Fabricius, etc. This paved way
K. Sreedevi () · N. Meshram · P. R. Shashank for the emergence of taxonomy and with Charles
Division of Entomology, Indian Agricultural Research
Institute, New Delhi 110012, India Darwin’s theory of natural selection, systematics
e-mail: [email protected] gained the momentum.

A. K. Chakravarthy (ed.), New Horizons in Insect Science: Towards Sustainable Pest Management, 3
DOI 10.1007/978-81-322-2089-3_1, © Springer India 2015
4 K. Sreedevi et al.

Systematics forms the basis for any life science ing up more than 58 % of the known global bio-
studies and advancement as it is important to all diversity. The insects are known to be the most
other fields of biology. It builds up the informa- successful and diverse animals on earth and are
tion on biodiversity of species. Taxonomy, a part closely associated with our lives and affect the
of systematics is the basic scientific discipline of welfare of humanity in diverse ways. At the same
biology that gives the identity and background of time, large number of insect species, including
the organism, on which all other related sciences those not known to science, continue to become
rely. Systematics, in addition to classification and extinct or extirpated from local habitats world-
naming, also deals with the relationships and en- wide. Our knowledge of insect biodiversity is
vironmental adaptations, thus drawing attention far from complete. Insects are the most exuber-
to the evolution and phylogeny. The morphologi- ant manifestation of earth’s many and varied
cal, physiological, ecological, behavioral, geo- life forms. The members of the class Insecta,
graphic, and molecular characters of the organ- arranged in 29 orders with more than 1 million
ism, in aggregate, are considered for the holistic described species (Grimaldi and Engel 2005; Ar-
approach in systematics. illo and Engel 2006), deserve serious attention
of the taxonomists. Four of these orders—the
Coleoptera, Diptera, Hymenoptera, and Lepi-
Insect Diversity doptera—account for 81 % of all the described
species of living insects. A growing number of
Now-a-days there is a good understanding of world checklists and catalogs are available on-
many things on the planet Earth and also the Uni- line for various families and orders, yet many
verse but we are still lagging behind on knowl- to find a place. Outfitted with search functions,
edge of how many species or life exist on the they provide another tool for handling the taxo-
earth. The term “biodiversity”, coined by Wilson nomic juggernaut of new species and nomencla-
(1988) as a contraction of “biological diversity”, tural changes. We can foresee a global registry
represents the diversity of life at all levels includ- of species in the near future that is updated with
ing genetic, species, and ecosystem diversity and each new species or synonym, allowing real-time
is the core of natural resources for sustainable counts for any taxon. It is imminent that insect
development and biotic capital for sustenance of biodiversity research must take cognizance of
life-support system (Kim and Byrne 2006). its material, taxonomy, insect pest management
The fundamental unit of biodiversity—spe- related requirements in light of recent develop-
cies—serves as focal point for studying the full ments viz molecular taxonomy, bioinformatics,
panoply of life, allowing workers to zoom in and information technology, and other advancements
out along a scale from molecule to ecosystem. The (Ramamurthy 2003). Though 2010 has been
species-centered view also provides a vital focus designated International Year of Biodiversity
for conserving life forms and understanding the by the Convention on Biological Diversity and
causes of declining biodiversity (Alder and Foottit the United Nations (Johns 2010), taxonomy, that
2009). The process of discovery and description of strengthens the knowledge of biodiversity, is on
all species is at very slow pace. This now appears constant decline (Wilson 1985; 2004).
unlikely to resolve the question in the near future,
if at all, for a variety of reasons such as the slow
rate of description of new species, the high level of Need for Insect Taxonomy
synonymy for most groups and the uneven distri-
bution of taxonomic effort which results in deficits Huge is the biodiversity of insects and little is
in the known number of species for many species- known from all the fronts. Species identity and
rich groups like insects (Stork 1993). information is the foremost step for advanced
Most eukaryote species awaiting description studies in any direction and that’s where system-
are insects (Raven and Yeates 2007), which are atics has a big role to play in. Despite the ongoing
the world’s most diverse group of animals, mak- biodiversity crisis, the number of new species de-
Insect Taxonomy—Basics to Barcoding 5

scribed per scientist has not increased in the past ography analysis are derived from specimens in
60–70 years, which has a huge impact on conser- collections. Global systematic work on insects
vation science (Terry Sunderland 2012). Many has a great deal to contribute to the understand-
species will become extinct before they are de- ing of evolutionary and geological events in the
scribed and one will remain continually unaware distant past.
of the total numbers of species that comprise Impacts of taxonomy on society are often
global biodiversity and this is acknowledged by beneficial, sometimes in unpredictable ways. A
the Convention of Biodiversity and its signatories case study related to the description of a new
as a “taxonomic impediment” (Terry Sunderland mealybug species in Africa on cassava reveals
2012). Taxonomy enables the facilitation of cer- the importance of taxonomy (Smith et al. 2011).
tain conservation issues like endangered species, Cassava (manioc or tapioca; Manihotesculenta),
species richness estimates, etc. for sustainable a drought resistant, staple food crop for over
management of the natural resources in a better 200 million people in sub−Saharan Africa was
way. An account on the role of taxonomy in spe- infested with new mealybug species, since de-
cies conservation was given by Mace (2004). scribed as Phenacoccusmanihoti in 1973. As a
Taxonomy not only produces fascinating result of misidentification and misdirected pest
knowledge on the characteristics of life but also eradication efforts, initial attempts to control
delivers basic and indispensible knowledge for this pest using natural enemies failed, severely
many fields of human interest and contributes in impacting the livelihoods of millions of people.
many ways to the sustainability of our planet. Re- Correct taxonomic identification came into res-
search in taxonomy and systematics involves the cue in locating its natural enemy, Anagyruslo-
study of virtually all available specimens of a tax- pezi, which established successfully by 1990 in
onomic group in order to ensure comprehensive 25 African countries implying a cost/benefit ratio
treatment, and is dependent on the availability of between 1:200 to over 1:600 (Smith et al. 2011).
well curated collections. In the course of these This is one of the examples where taxonomy has
studies, species previously unrecognized are fre- helped. In reality, many taxonomic works did
quently discovered. A single holotype specimen not get highlighted as it does not deal directly in
designated for each species is the standard of management of insect pests on crops, humans,
definition for that species. Much of research in and animals. Nevertheless, there will not be a
biology is ultimately dependent on the scientific proper applied research on an insect without tax-
name of the species. onomic details.
Taxonomy is the pivotal but hidden service The traditional taxonomy provides the most
behind sectors ranging from conservation and convenient and authentic classification based on
management of biodiversity to food security, the overall similarities, most visible characters
poverty reduction, health, biosecurity, new in- between species. It is pivotal in species recogni-
dustrial product development, and ecotourism. tion (with identification keys) and management
Wherever evolutionary history is relevant to a of biological collections. At the beginning, clas-
problem, systematics provides the resources. sification work was restricted to just taxonomic
Necessarily, then, systematics is at the leading details of the organism without considering the
edge of the study of evolutionary biology; and degree of relatedness between species. Later in
its central position is assured because new con- 1950s, the phylogenetic classification cropped
tributions of molecular and genetic research to up to take care of the evolutionary history of the
understanding the evolution of species have to organism. The different schools, (part of conven-
be related to the broader systematic concepts of tional taxonomy) that differ in their concepts of
the taxa concerned. Interpretation in biogeogra- phylogenetic classification but still converge on
phy depends substantially on an understanding of the basis of morphological similarities between
evolutionary history, and consequently is closely species, are presented hereunder.
allied to systematics; the basic data for bioge-
6 K. Sreedevi et al.

Conventional Taxonomy Phenetics or Numerical Taxonomy

1. Evolutionary or traditional taxonomy Phenetic systematics determines the relationships

2. Phenetics or numerical taxonomy of organisms through a measure of similarity,
3. Cladistics or phylogenetic systematics considering plesiomorphies (ancestral traits) and
4. Cladoendesis apomorphies (derived traits) to be equally infor-
mative. It aims at natural classification using nu-
meric algorithms like cluster analysis rather than
Evolutionary Taxonomy using subjective evaluation of properties. A priori
every character is given equal weight. From the
Evolutionary taxonomy, originated in early twen- twentieth century onward, it was superseded by
tieth century, attempts to classify the organisms cladistics, which considers plesiomorphies to
based on phylogenetic relationships coupled be uninformative for an attempt to resolve the
with degree of evolutionary changes. It takes phylogeny of earth’s various organisms through
taxon into consideration rather than a species. time. Today’s systematists generally make exten-
The characters differ in information content re- sive use of molecular biology and computer pro-
garding phylogeny and hence have different grams to study organisms. An alternative to these
weights. Both recency of phyletic splitting and matrix methods in phylogenetics and systematics
rate of divergence are given importance. Evolu- is cladoendesis.
tionary taxonomy evolved through the influence
of theory of evolution on Linnean classification
during post Darwinian period where the tree of Cladistics or Phylogenetic Systematics
life gained importance in scientific works with
publication of The Origin of Species. In 1930s, Cladistics got conceptualized in second half of
few biologists developed a Mendelian frame- the twentieth century and is termed as Phyloge-
work for Darwinian evolutionary theory, result netic systematics by Willi Hennig (also the title
of which was the evolutionary synthesis (Marc of his 1966 book). In cladistics, classification is
Ereshefsky 2007). Theodore Dobzhansky, Ernst mainly based on common ancestry and hence, be-
Mayr, and Gaylord Simpson were the few of lieves in cladogenesis, where two taxa originated
evolutionary taxonomists. The school consists of in the same branching event have a common an-
two tenets, firstly all taxa being a genealogical cestor that is not shared by any other taxon. Thus,
lineage and secondly constructing classification cladistics represents only monophyletic taxa in
that reflects both cladogenesis (branching) and their classifications. Those who follow cladistics
anagenesis (divergence). In cladogenesis, specia- perceive that the concepts of phenotypic differ-
tion occurs with the selection pressure (genetic ence and adaptive zone are ambiguous and are
revolution) on the isolated population from the applied inconsistently to different types of taxa
rest of the species where single lineage splits into (Hennig 1966; Eldredge and Cracraft 1980). Cla-
two branches whereas in anagenesis, speciation dists believe that the concepts of phenotypic di-
occurs in a single lineage. As a result, evolution- versity and adaptive zone are too malleable and
ary taxonomists see two types of taxa viz., mono- reject them as grounds for classifying taxa (Marc
phyletic and paraphyletic taxa arising from the Ereshefsky 2007). A group of cladists developed
processes of speciation through cladogenesis and the Phylocode—a phylogenetic code of biologi-
anagenesis, respectively. In brief, evolutionary cal nomenclature, which is considered alternative
taxonomists believe that classifications should to the Linnaean system (Cantino and de Queiroz
highlight only genealogical taxa, and those taxa 2004). The widely used and popular phylogenetic
can be either monophyletic or paraphyletic (Marc approach is cladistics.
Ereshefsky 2007).
Insect Taxonomy—Basics to Barcoding 7

Cladoendesis—New Approach to nucleotides or bases in either DNA or RNA seg-

Phylogenetic Construction ments extracted using different techniques.
The theoretical framework for molecular sys-
The term cladoendesis was introduced by N. J. tematics was laid in 1960s which plunged into
Kluge in early twenty-first century, meaning DNA–DNA hybridization during 1974–1986.
“branch coupling” that pays more attention on The advantage claimed for using hybridiza-
the connection between apomorphies of each tion rather than gene sequencing was that it was
taxon and characteristics of higher taxa, so that based on the entire genotype, rather than on par-
the characters of all the taxa are, from the very ticular sections of DNA. Another application in
beginning, considered to be interrelated within molecular phylogeny is DNA barcoding, wherein
a certain hierarchy (Kluge 2012). It is a method the species of an individual organism is identi-
of phylogenetic analysis opposed to various ma- fied using small sections of mitochondrial DNA
trix methods. The phylogenetic trees are not built (mtDNA) or chloroplast DNA that demarcates
each time as new ones but reconstructed based on species as lineages (Hebert et al. 2003).
the previous results where each character of each
taxon is compared with its ancestral condition in
the ground plan of the higher taxon (Kluge 2012). Insect Mitochondrial DNQA and DNA
Cladoendesis enables understanding of nature Barcoding
and evolution of metamorphosis in insects.
All the schools eventually yield phylogenetic Mitochondrial genes are often chosen for evolu-
trees, a visual representation of the fact that spe- tionary studies as they have a number of positive
cies are related by descent from a common ances- characteristics like: (i) maternal inheritance with
tor. Depending upon the type of school used in little or no recombination (ii) general conserva-
the construction, it may be called as phenograms tion of gene order and composition (iii) small
(arrived from phenetics) or cladograms (arrived size compared with the nuclear DNA and (iv) the
from cladistics). lack of introns (Gissi et al. 2008). mtDNA have
Molecular biology has taken the systematics proven to be informative in the study of species
towards a different turn. The convergence of un- diversity and evolutionary processes because it is
related in species under similar selection pressure easy to isolate and contains conserved sequences
and divergence of related in species under differ- that make it possible to use as universal primers
ent selection pressure yields to surge toward mo- (Otranto and Stewens 2002; Xie et al. 2006).
lecular taxonomy as an additional tool in support Extensive studies in the mtDNA of Dro-
of traditional taxonomy in diagnostics. Recent sophila species showed that the same genes are
advances in molecular techniques have greatly present in both mammals and invertebrates; how-
helped to resolve the controversial classification ever, their arrangements may differ (Clary and
schemes based either largely or entirely on mor- Wolstenholme 1985; Crozier and Crozier 1993).
phological attributes (Viraktamath 2003). The protein coding genes are the most frequently
sequenced mitochondrial genes for evolutionary
studies and phylogenetic analysis. Protein coding
Molecular Systematics genes commonly analyzed include; COI, COII,
16S, and 12S. In particular, the COI gene has
Early attempts at molecular systematics were also been widely sequenced. Yet, the specific region
termed as chemotaxonomy that made use of pro- chosen has varied from study to study (Lunt et al.
teins, enzymes, carbohydrates, and other mole- 1996; Caterino et al. 2000). The first subunit of
cules that were separated and characterized using mtDNA CO gene, corresponding to nucleotides
techniques such as chromatography. These have 1490–2198 of the D. yakuba sequence, has been
been replaced in recent times largely by DNA se- identified as an area of interest for “DNA bar-
quencing, which produces the exact sequences of coding” (Hebert et al. 2003). This region has a
8 K. Sreedevi et al.

rate of molecular evolution that is about three acters that should be less subject to convergent
times that of 12S or 16S rDNA, its third position evolution than other characters that might lead to
nucleotides showing a high incidence of base a confusion of grade and clade. A character can
substitutions. The success of universal primers be phylogenetically informative when nucleotide
for this gene enables the analysis of amino acid changes are shared by two or more taxa. A char-
substitutions to initially designate an unidentified acter can be phylogenetically uninformative when
organism to a higher taxonomic group before ex- all nucleotides are the same among taxa, or when
amining nucleotide substitutions to determine its only a single taxon has a different nucleotide.
species identity (Hebert et al. 2003). According to Hillis et al. (1996) three main
Hebert et al. (2003) named this technique applications of molecular systematics are,
“DNA barcoding.” Then, the Barcode of Life 1. Reconstruction of phylogenetic relationships
project was proposed to promote DNA barcod- of organisms.
ing as a global standard for sequence-based iden- 2. Studies of population structure, including geo-
tification of eukaryotes. Recently, the Barcode graphic variation, mating systems, heterozy-
of Life project entered a new phase with the gosity, and individual relatedness.
launch of the International Barcode of Life proj- 3. Identification of species boundaries including
ect (IBOL; International Barcode of Life 2012). hybridization.
The IBOL is a huge international collaboration of There are many methods to understand these mo-
26 countries that aims to establish an automated lecular variations; few molecular methods with
identification system based on a DNA barcode their applications were explained in Table 1
library of all eukaryotes. In the first 5 years, the
IBOL will focus mainly on developing a barcode
library, including 5 million specimens of 500,000 Molecular Markers Used in Systematics
species. The DNA sequences are used as genetic
“barcodes” that may potentially be used as a bi- A diverse range of novel molecular (DNA)
oidentification system for all animals and have markers are now available for taxonomic inves-
proven to be a useful identification tool for ver- tigations. Both DNA and protein markers have
tebrates such as birds (Hebert et al. 2004), fish revolutionized the biological sciences and have
(Ward et al. 2005), and hexapod orders such as enhanced many fields of study, especially sys-
Lepidoptera (Hajibabaei et al. 2006), Coleoptera tematics. This has been possible because of the
(Greenstone et al. 2005), Diptera (Smith et al. rapid advances in molecular biological methods
2007), Hymenoptera (Smith et al. 2008), Ephem- and bench-level protocols for wider application.
eroptera (Ball et al. 2005), and Hemiptera (Lee The utility of molecular markers as additional
et al. 2011). DNA barcoding does not substi- tools in systematic has led to “molecular system-
tute but complement conventional taxonomical atics”. Over a long time, significant contributions
studies. have been made in the field of insect systematics
The three main taxonomic applications that through morphometric traits, wherein a number
DNA barcoding has been previously used are: (1) of difficulties were encountered due to geno-
the identification of species previously defined type–environment interactions. The limitations
by other criteria, including rapid identification, in using morphological, physiological, and cy-
as well as linking specimens to established spe- tological markers for assessing genetic diversity
cies that are unidentifiable by other means; (2) and population dynamics have been largely cir-
the description of new species by interpreting cumvented by the developments in DNA-based
DNA diversity as an indicator of species diver- markers. Molecular markers, by nature, are neu-
sity; (3) the definition of operational units for tral to the stage of development, physiological
ecological studies (Rubinoff 2006). status, and environmental influences. Isozymes
DNA should be an excellent tool for inferring and other proteins as markers are often expressed
phylogenies: large number of homologous char- codominantly and discriminate homozygous and
Insect Taxonomy—Basics to Barcoding 9

Table 1   Some methods presently employed in identification of the species. (Adopted from: Singh (2012))
Methods Explanation
Hybridization Genetic materials from two different species are subjected to hybridize. Closely related
species show higher percentage of hybridization
DNA sequencing DNA segments of two species are sequenced from one end to the other and the
sequences of the two form the basis of establishing similarity or dissimilarity between
them
Restriction mapping Segments of DNA are isolated from different species and subjected to restriction map-
ping. Closely related species will have more similar restriction map
Chromosome banding The chromosomes of different species are examined through microscope. Banding of
chromosomes are also done for taxonomic purposes
Amino acid sequencing Like DNA sequencing, protein sequencing is also done. The amino acid sequence of a
given protein will be more similar between closely related species
Immunological methods Antibodies that recognize specific macromolecules, usually on the cell surface are
tested on different species. Antibodies that recognize macromolecules form one species
will often recognize closely related species, but not from distantly related species

heterozygous individuals. However, the limited Integrative taxonomy gives priority to morpho-
number of proteins and isozymes as markers and logical characters because of their greater com-
requirement of different protocols for each en- plexity and presumed multigenic origin, which
zyme/protein limit their utility. Unlike morpho- are believed to constitute a more secure basis for
logical and protein-based markers, several DNA separating species than small fragments of DNA
based markers are available to elicit the differ- sequence. The limitation of molecular system-
ences between individuals and populations, or atics—being an essentially cladistic approach,
they can be developed for each specific purpose. is that it assumes that classification must corre-
Although a large number of samples can be ana- spond to phylogenetic descent, and that all valid
lyzed quickly, a number of other factors such as taxa must be monophyletic also lead to integrative
cost, speed, and requirements of technical skills taxonomy. The recent discovery of extensive hori-
are the major concern. DNA-based markers can zontal gene transfer among organisms also pro-
generate large amount of high quality data com- vides a significant complication to molecular sys-
pared to several biochemical marker systems, but tematics (indicating that different genes within the
degree of polymorphism detected and the sta- same organism can have different phylogenies)
tistical dependability of the results vary among necessitating integration of both conventional and
marker systems. molecular taxonomy for holistic approach.
Molecular taxonomy, on other hand, can facil-
itate easy and rapid identification of the species
Integrative Taxonomy provided the gene sequence has been deposited
after authentic identification of the species mor-
Will et al. (2005) a slight critique of DNA barcod- phologically supported by all metadata and pho-
ing, used the term integrative taxonomy to mean tographs. Consistency index of molecular data is
a taxonomic process that was inclusive of all higher than that of morphological data. But ac-
available data sources and not just mtDNA COI curate identification always comes from the mor-
barcode data. Dayrat (2005) defined “Integrative phological characters and hence addition of the
taxonomy” as the science that aims to delimit the new species to the list of fauna is encouraged by
units of life’s diversity from multiple and com- conventional taxonomy, which can be supported
plementary perspectives. Thus, any study linking by molecular taxonomy. Both approaches have
different kinds of data by mapping morphologi- issues and limitations, which can supplement and
cal diversity on to a molecular phylogeny is inte- complement for holistic species identity.
grative (Yeates et al. 2011).
10 K. Sreedevi et al.

Schlick-Steiner et al. (2010) have offered the assisted programs based on character data (let it
most detailed treatment of the process of integra- be morphological in terms of numerical values or
tive taxonomy. They stressed that integrative tax- DNA sequences).
onomy does not replace traditional taxonomy, but Another dimension of ICTs in systematics is
that it uses complementarity among disciplines to in the field of automation of information with
improve rigor. Their integrative procedure relies the taxonomic expertise in their respective areas.
on an agreement among three “conclusive” dis- Lyal et al. (2008) stressed the need of accessi-
ciplines that is both proscriptive and restrictive, bility of user-friendly identification tools. Digi-
and disciplines and datasets are defined rather tization of the biological collections and descrip-
arbitrarily. Several studies cited as examples of tions helps in buildup of virtual information for
integrative taxonomy by Schlick-Steiner et al. the benefit of all researchers and scientists. The
(2010) used correlative approaches to compare transformation of the processes of science and its
molecular data and morphology (Malhotra and outcomes is possible only through ICTs which
Thorpe 2005; Yoder et al. 2005; Rissler and Apo- are adept at dissemination. The databases created
daca 2007; Roe and Sperling 2007). The determi- would largely help the present and future workers
nation of origin and their evolution trajectories of in respective line of work.
a species will trigger the species delimitation in a The taxonomists mandate should be built up
better manner (Padial et al. 2010). of databases, online publications of new taxa and
Yeates et al. (2011) surveyed the current taxo- monographs, development of interactive keys
nomic methods (inclusive of both morphological and virtual repositories, generation of distribu-
and molecular characters) employed in delimit- tion maps and predictive models utilizing both
ing the species and identified two challenges, the approaches, conventional and molecular for
which if met, will provide a more complete the benefit of speedy growth.
toolkit and a more robust research program in
integrative taxonomy and proposed alternatively
the term “iterative taxonomy” for the practice of Conclusion
treating species boundaries as hypotheses that are
to be tested with evidence. Delineating the spe- We attempted in this chapter to present an over-
cies boundaries accurately is crucial to discovery view of the genesis and the growth of taxonomy,
of life’s diversity (Dayrat 2005). both as a science and as a tool, of species iden-
tification and documentation. The integration of
new knowledge and methods of population biol-
Role of Information and ogy, phylogenetics, and other evolutionary dis-
Communication Technologies (ICTs) in ciplines into taxonomy is warranted (Sites and
Systematics Marshall 2004). The analysis and interpretation
of data used to delimit the species have profound
A biggest challenge of a systematist while deal- implications in taxonomic research (Yeates et al.
ing with the larger taxa with numerous species 2011). Dayrat (2005) stated that “integrative tax-
having possessing complex variations at species onomy” gives priority to species delineation over
and population levels is to look into the charac- the creation of new species names and also opined
ter evaluation (Ramamurthy 2003). Evaluation that a radical change in mentality is needed con-
of character itself is a complex task that involves cerning the creation of names in order to achieve
computational methods in screening the charac- this integration and to prevent the overabundance
ters (Ramamurthy 2003). Computer simulation of both synonyms and names of doubtful applica-
and modeling help in resolving certain phylo- tion from worsening.
genetic issues in building up the tree with in- The integration of all possible taxonomic ap-
tercession. Potential evolutionary relationships proaches abridging the gaps of each in arriving
are being evaluated with the help of computer at correct species delimitation is the need of hour
Insect Taxonomy—Basics to Barcoding 11

in the light of biodiversity inventory. Padial et al. Greenstone MH, Rowley DL, Heimbach U, Lundgren JG,
Pfannenstiel RA, Rehner SA (2005) Barcoding gener-
(2010) stated that taxonomy needs to be plural- alist predators by polymerase chain reaction: carabids
istic to improve species discovery and descrip- and spiders. Mol Ecol 14:3247–3266
tion, and to develop novel protocols to produce Grimaldi DA, Engel M (2005) The evolution of insects.
the much-needed inventory of life in a reasonable Cambridge University Press, Cambridge
Hajibabaei M, Janzen DH, Burns JM, Hallwachs W,
time. Insects, being vast and diverse on earth, Hebert PDN (2006) DNA barcodes distinguish spe-
need much more integrative taxonomic attention cies of tropical Lepidoptera. Proc Natl Acad Sci U S
than any other life system. The unique charac- A 103:968–971
ters of an organism that unravels the diagnostic Hebert PDN, Cywinska A, Ball SL, Dewaard JR (2003)
Biological identifications through DNA barcodes.
character differences that delimit the species Proc R Soc Lond B 270:313–321
(Sites and Marshall 2003) have to be assessed Hebert PDN, Stoeckle MY, Zemlak TS, Francis CM
holistically from all directions and taxonomic (2004) Identification of birds through DNA barcodes.
approaches. Perusal of various concepts and PLoS Biol 2(10):1657–1663
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 PRB 4.4710N 0 74.X0NIG P,APBRS
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The style ofTheychea
depends on the group. In ornithology
valid nanes nd synonvms is given, together with usually a complete
a detailedreferenc
desc
of the range.
Typical examples of each of these are litles ned not be as bad as these, however, to ctse dificulties fou
(nn Cali).
Duze, EUniw.P. 117. Catalogue of the Iemipteru of Anerica north o
":Ey',."
Pub. Bn., no. 2, XIV i 02 pp.
",",",",Za"," 'asataloguers,
tndercause forabstracters, ofreviewers,
complaintanhieuous
an incomnlste.
andbeingotheroverlooked
his orworkmisleadinr bibliographers. No author
title if it msquerade=
Mvr,lepidoptera.
141. Lit20South.
NewB. York,
Ca). A cd. Sci. M en., 1, 175: 3, I-1I.
pp.of New Guinen birds. American Museum of Natural
TORM OF TE TAXONOMIC AR IICLE hrrrrrrrür
Ttle. The itle is the first part of the paper encountered by the onfusion which resulted still persists in modern iterature. Women
ilthough it is often the last item to be aded in the preparatior
pper.
iare inItsits bibliographical
selection. The titleprominence should beandlongsignificane
enough to bewarran specit FrüEE
Ihe eontents of the paper but brief enough for easy indexing.
shoul be nenar the beinning of each series ofThewords.
vords ure preferable to polysyllabic terms. most Theimportan
title EE
'ise so that postal authorities can reconize it.
 l'AAUN UullL PUCLU t Pb 41 0 T130N1 P, IS
When more than one author is involved, the order of names is romplete
nations list inof previous
used all references to the species,
publications. In the with the names groups
better-known and combiof
mined by tbe nature of the contribution eachis solved
has made.by aWhen the unimals this is both unnecessary and uneconomical, and this fuction
has been rather
in which case theequally namesshared,
are usually the problem arranged coautho
in alphabetical reserved
unimals for bibliographical
(e.,areinsects) catalogues. Unfortunately,
have beenthatsobibliographical
incompletely catalogued, some groups
or existingofis
When the work has been disproportionately divided or there is a m satalogues
discrepancy
result. In suchin agecasesortheexperience,
name of the a senior author
and juniorappearsanauthorship ssential element of the full taxonomic treatment. This is especially truean
so out-of-date, synonymies are still
Introduction.
paragraph stating Every
the scopetaxonomie
of the paper papersenior
shouldwhereinclude
and, pertinent,
first.
introdv
the re
shen much of the literature prior to 1900 is more significant nomenclatur
lyroologically
than zologically, and the laterAnpublications are more significant
for the study, as well
uthebrief s the nature of theThese studiedfeatures
material. serveFrequ , 4ha rvlerthanwvvkiwe nomenclaturally.
tavooist. iunderstanding
sah orru of both is required
casualhistorical
the minds ofreader
review
the newinis theappropriate.
other andworkers student
field.of the group, as well as to r
tou
oi data: (1) scientific name (in its original form), (2)Hio
NM RgHig i,) Hl uiuJ Mw w Mi ul author," (3)HhdateWilof
publication, () reference,
example, (5) type locality, (6) present location of type
duction when they can be treated as part of the natural sequeni
Acknowledgments. Acknowledgments may be included in the 'optional). For
cxposition.
author's name. Some authors place them inusea footnote appended t
molopical SocietyThis system
AmcricaisSometimes
ofcontent. andin certain
regular other byjournals
the Annals
whichofpreced
arethep Oncideres
Mex.;
Oncideres
rhodosticta
Britishtrinodatus Bates, 185, Biol. Cent-Amer., Coleopl., 6:3867. (erdo
Mus. (Nat. ist.)l.
rily
SDIir. taxonomic in the acknowwledgments Vus. Ne synonygmy.FCasey, 1913, Mem. Coleopt., 4:352. IEl Paso, Tex.: U.S. Nat.
Methods Used and Materials Studied. In a revisional or n 'he above form is sufficient for a revision of a well-catalogued group.
graphie work it is desirable to include a statement on methods utilize In groups where remains the literature hasa full not been summarized adequately and
tollections, spcimens, or other materials studied. This enable the
ceader
vork. to evaluatemethods
Standard conclusionsmeasuring, and to judge the thoroughness
staining, sio ificnomenclature
names, incorrect andconfused, synonymy
correct, specifying the(i.e.,booksa list
and ofauthors
scien
preparations,
nethodls ned toetc., may beforreferred mounting,
in detail.to by name and reference. Only
mploying
shich have them) may be required. This should include all referehces
be described ically undernomenclatural
the actual or zoological
name (correct orsignificance,
incorrect) byarranged which thechronolog
author
ofBody of thedepend
course, Text.on Thethe material
scope andcomprising objectivestheof body of the textpapw
the particular vctually
tevice of areferred to them.
comma inserted Many theauthors
between specifieherenameuseandthetheconvenient
author IX
It is perhaps
includes () a sufficientoftothemention
definition highest thatcategory
a complete
includedsystematie paq
(family,(gener
tri vssihich
albus,hasSmith (not Brown)) to distinguish between a misidentification
tc.),
13) (2) a key and(or descriptions
synonymies keys) to all ofintermediate
the categoriescategories
intermediate treated (gener 'not
synonymyBron))is asnowhichfollows:
nomenclatural
has. A widelystatus,
used form and fora homonym (-s abus Jones
a full bibliographical
1)16)statement of the generic types, (5) comparisons with other gene LAncers roscf us 1ut8
auchkeysspecies,
to theandspecies of ech genus,
(8) statements as to (7)typesynonymies
Iocalitiesandanddescriptions
to location 'nucideres
Hex.: Britishrhodosticta
Mus. (Nat.Bates,
Hit.):I,Linsley,
Biol. Cent-.
1940, mer.,
Jour. Colopl.,
Bcon. Ent.,6 :86733 :562
Itype:(synon.
Lerdo
Iypes, generalwithdistribution,
omparisons other species, hosts
etc. and(forother significant
details biological
on preparation of descrda
lionsSynonymy.
aud keysInseemonographs,
Chap. 8). revisions, and catalogues wot
Birds it synoivrme that san be found i ie pvioa sundud o-kes: cudbpue
(18-90)g
ud advisable to giveof thethe development
complete synonymy of every species.custou
it is Dur More rcentof thechecklists
Britishof Mrueum
birds do(1873-182) and the orretly
not repeat svnonyma andtet cited o' Birdby Peter.
tIhe earlicrtostages of our taxonomie L,a7T7p,,ta,,at,t,,;
customry give not only the synonyms but alsoliterature
a more orit IV lot to be interpreted as a nomenclatural chnge.
 l'A.XUMIU' O H PbAATI0N O 1'AX0NIG APERS
distr.);
1945, Sci.Linsley, 1942, Proc.
Pub. Reading M us.,Calif.
no. 5:Acad.
v (key);Sci.,Dillon
() 44and:76Dillon,
(distr.);1946,DillonLc.,and6:313Di Genus Dirurus Vieillot
i 12:195Lkey,Ldistr.);,Schaeffer,
A 16,ue, se, A. Dicrurus
taire, p. 4.1841,Type,A listbyAprilsubsequent
Vieillot, 14, 1816,designation,
Analy'se d'uneCorvusnouvelle orithologieLinneusélémen
Can. Bnt., 88:19 (kevy). (G.Edolius
R. Gray, of the generaLe règne ed. 2,vol.p. 47.1,balicassius
of birds,animal,
So.,Oncideres cingulatuas, Hamilton (in part) (not Suy, 182), 1896, Trans. A mer.
28:11 distr.). subsequent Cuvier,
designation, De. 7, 1816,
Lanius for ficatus (G. R. Gray, 155, Cataloguebo
p. 350. Tvrpe,
Oncideres
Nat. Mus. Craighead, 1923, Can. Dep. Agr. Bul. 17 (n.), p. 12 (larva, ho7
U.S.Oncideres trinodatu Caseoy, 1918, Mem. Coleopt., :32, (typ: E Paso, of Drongo
the generaTickell,
nd subgenera Jour.of birds,
I3,ulescns Asiatie 58).Linnaeus
p. Soc. Bengal, vol. 2, Linnaeus.
p. 573. Tvpe, by
Ohcideresp. sp.,
America, pustulatus,
460, Fig. 368Essig
habits,(notditr.Le Conte, 18. 192, Insecta of Western N monotypy, Drong cuer Tickell e Lanius cerulescens
ChibiaEdoliua
nation, Iodgson,barbatus
86, IdiJ. Rev, E. vol.m 1,Corusp. hottentotus
Gray 324. Type, bylinneus subsequent
(G. R. desig
Grav,
The above synonymy might appenar in an abbreviated check lis I141,Bhringa
A listHodgson,
of the genera ofIndia
birds,Rev,ed. 2,vol.p. 47).
iuation and monotypy, 1886,
Bhringu tectirotris 1. p.
Hodigson. 325. Tvpe, by original desig
Bhuchangu Hodigson,
lesignation, Bhuchanga albirictus18, India Hodgson Rev, vol. , p. 326. 1877, Type, by sulbsequent
theC'haptia
British Museum,
Hodgpson,vol.Hodgson 245).Rev., vol.(Sharpe,
16,3, p.India
Catalogue of birds in
theWcheck
hen lista checkmay bestincreased
contansbyagiving terl page
bblorapy,referenceste which
uselulms
C'hapta muscipetoides =m Dicrurus aeneus1,Vieillot.
p. 326. Type, by monotypy.
betictus
located
Bates,by 1886
author, :367, date,
or, and page
more simply,in the
86:367. bibliography. Thus, Dissemurus Gloger, 1841, Gemeinnutzives Hand und Hilfsbuch der Natu
In a complete synonymy it is often desirabe to indicate the geschichte,
M usicusdetails,p. 37. Type, 180,
Reichenbach, by mootypy,
Avium Cuculusnaturale,
systea paradiseuspl. Linnaeus.
8, fig. 9. Figure
combinations under which each name has appeared. This may l fAad.generie no species
veniently
it through accomplished by taking thethenoldest
its various combinations, the noxtspecifie
oldest,numeetc.,
and sfolf lot i Sci.
(orvusParis, , p. 50. Typo, by tautonomy, Dicrurus musicusRendus
adsimilsvol.Bechstein.
included, ef. Bonaparte, 1854, Compt. Vil
M egcyene antennata ( White) Dicranotreptus
Figure of generie Reichenbach,
details, no 180, Avium
species included.systema
Type, naturale,
by subsequentpl. designation,
S, fig. 12.
Edoliusandmegarhynchuw
genera subgenera of birdis, Quoy andp. Gaimrd
5). (G. R. Gry, 155, Catalogue of the
Cyllene antenatus,
Craighead, Horn,A 18s, Trans.
Bul. 27,A p.ner.38 nt.(arvna,
Soc.,biol.):
8:135 Hoppinz,
(desr, syn.,
Ent.I Soc. A 123, Can.30: 441,
mer.,UiII, Dept. pr.,
pl.IIIl,
1 (revis. 1987 Summary. A summary is usually unnecessary in a strictly taxonomie
paper. When required, it should
WGyleIt
Arhopalus eurystethus L.LeConte,
Sonora, p.Mex.;17, pl.Mus.1,Comp.
Ml II.Proe.L Witpl,
1858, Acd. Nat.8Sci.103,Phila.
0I (T.,
1868 :s8 tyle.
pecifie,It notshouldin broad
be written a beseries
generalas terms.
briefofbutshortshouldparagraphs
not be inandtelegraphic
should be
Naturae, Fige. 9.Zool. Hrvard;: LaeConte, 1859, in Thomson, Refterences and Bibliography. References are generally treated either
the Inauthor's
the abovenameexample,
has againthebeencommaused,between
this time the specifie combintic
to distinguish betv L'"":"'E,;T";72E:;Ttgafar;;
new combination repeated reference
are costlyto topublishing
to thehaundle
sumeinbibliographical item is unnecessury.
bination (Clytus(Cgllene
antennatusantennatus,
White, 185).Horn, 1880) and an original Since
uy addthey materially costs,tvpeseting and printing.
parenthetical references howvever,
are toandbe
Generie
ynonymy, synonymy is handled in much the sme way as as
treatment,except that tpe
the generie in the(andcase of new synonymy
its designator, if any)orisfullcitedbibliog
in pl E
herus,typeas cited
locality and type location. The synonymy of the genus l
as an example.in Varie's (1949) revision of the Dicruridae, may be hould
nay bebegreatly
selected.increasd
Frequently the valueparenthetical
by including of the terminalcomments
bibliographv
on the
 TAXONOMIC PROCEDURE
1'111/P,tlt.11'/0N OJ/ 'l',1.'<0NOM tc l'tlPt-:1:S 189
188
ln tho tYJ)(J6cript, Ioomctes are entered beneath a marginal line in the
nature of the subject matter covered. Unverified references may
included when necessary for completeness, but they should be mark text {see oxam?le~hove) rather than at the bottom of tho page, becauso in
with an asterisk or some other device to indicate the fact that the author the. fin~I pubhcat,on, pagination is entirely different from the original
pag1rul.tlon.
has not seen them. If the work cited is by SC\'eral authors, only the first. need be reversed
Bibliographical it.ems should receive full citation, including author
for alphabetical pul'poses. Thus,
title, publication, volume, pages, date, etc. Text referencesto the ter ·
nal bibliography may be made by enclosing an author's name and da · :\fcAtce, \V. 1...., and J, R. M1dJoch. 1922. Chl\ngcs ln OSmC'.S ol' American Rhyu-
{sometimesalso page) in parentheses. Two or more references to public rhot.a., chiefty Eme.,:inae. Proc. B'iol. Soc. Wash., 36:95-06. ·
tions by a single author in t,he same year may be designated by append
The original style of capitalization and it,11ics may be followed. How-
letters (Smith, 1940a; Smith, 19406). The nuthor-datesystem of bibli '
e-ver, many titles are set entil'ely in capitals; others are set entirely in
graphical reference is far more satisfactory than tho straight numberin
low~r <:850, except tJmt the first word and scientific and place 1lll.mes are
system which is sometimes used. The number system tells nothi ·
cap,tahzed. The full title should be given in all but the briefest of foot-
about the reference; moreover, the author-date system permits t
notes, because i·eadei·s obtain "aluable leads in this way.
addition of references during the preparation of the manuscript withou
Abbreviations of journals should follow such standard works as the
the necessity of renumbering all references beyond the point of insertio ·
IVorld Liu of Sci,mtific Periodi.cal$ or the list of Abbreviation$ 11.sed in the
A formal "Bibliography" implies completeness of coverage of the su ·
Department of Agricult-ure for Titles of Pul,lir.atwn• {Whitlock Carolvn
ject. "Literature Cited" indicates restriction of referencea and is ·
U.S. De!'t. Agr. Muc. A,b. 337, pp. 1-278, 1939. Price 30 ce~ts). • '
explanatory. The c1t,at.1on: {'\Vhecler, L889a) is specific enough because it refers to a
Examples of footnote entries and terminal bibliographies are giv
two-_page paper. On the other hand, it may be necessary to refer to a
below. Numerous exceptionswill be encountered, especially various go
particular page of a larger work, thus: (Wheeler, 1910, p. 263). ln this
ernment documents, but a majority of literature citations will fit into ·
case the page is indicated in the citation. and the complete work is listed
or another of the simple styles illustrated. It is becoming standard p
in the bibliography.
tice to list the year of the publicntion immediatelyafter the author's na
since this sequence sgreee with thnt of the author-datesystemof referen PREPARATION OF THE MANUSCRIPT FOR PUBLICATION
Bibliography Aside from matters in\'olvcd in the actual organization and construc-
Wheeler, William Morton tion of a taxonomicpaP.Cr, there are some points which should be kept in
1889a. Homologues in embryo Hemiptera oi the appendages to the 6n1t abdo mmd 10 order to facilitate editorial handling after the paper has been
nal segment of other insect embryos. Anurican Naturalil.t, Vol \
X)..'lTI, pp. 644-645.
submitted for publication. Editors arc much more apt to accept readily
1889&. Uber drusensrtige Geblldc iro ersten Abdomioalsegment der Hemipt and publish quickly papers which are in good form and require a minimum
enembryonen. ZoolO{li-flcktr Anzeiga\ Band Xll, pp. 500-504, 2 figs. amount of editing. Most publications h,we special form requirements,
UUO. Ant1J, t.lmir structure, development and behavior. New York. Columbia :,nd much editorial time can be saved by careful advance reference to t-he
'Univcr&ity Prese, pp. xxvi + 664, front. 286 figs. 8vo (Columbia iom·nnl in which the paper is lo be published.
University BioJogit'Al Series, IX).
. . Typi~~· AU manuscript submitted for publication should be typed.
Literature Cited l he origmal draft$ may be on yellow paper, but the final copy should be
Wheeler, W. M. 18890. Hcrnologues in embryo Hemipters of the appendages to'
on standard (8.K by 11 ln. or 8 by 10 Yz in.) white paper, entirely double-
the futst abdominal l!legmcnt of other insect embryos. Amer. Nat.., 23:0H-646. •J>aeed (some publications require triple spacing), and "1th a wide margin
1889b. Uber drneenartjge Gebilde im cr&tcn Abdominelsegment der Hemip- lor adding proof marks and for editing. If approximately the same
terenembryonen. Zool~ Anz., 12:500-504.. number of lines is typed on each page, the editor can conveniently e.sti•
1910. Ania, their structure, development and behavior, mate the size of the final printed paper. However, some editors require
Presa, New 'York. xxvi + 664 pp.
that pages end with completed paragraphs. Pages should be numbered
(Footnote] consecutively in the upper right-band corner. Inserted pages are num-
• Wheeler, \\'. l\t. IAA9. .4mer. Nat., U :644-645.
bered alphabetically (e.g., 65a). Whole sheets should be used for i~er-
100 TAXONOMIC PROCEDURE l'R.Ei'1tR.AT/ON OF TAXONOMIC PAP/!118 191
tions, regardlessof length of inserted matter. When it becomesnecessary. always advisable to have other persons read a. manuscript, before it is
to cut and rearrange, sheets should be assembled by pasting, not. by submitted for publication. A fully corrected carbon copy of the manu-
pinning. All tubular material should be typed on separate shoots, smce: script should be kept by the author for use in case the original is lost.
it is usually set in a different type from the text. . . Proofreading. Most scientific journals permit the author to read
Underlining. Underlining indicates that the material so marked 1~ t ~ proof on bis papers before publication. Some few journals placethe entire
be printed in italics. Jn a, taxonomic,manuscript submittedfor publica-, burden of the proofreading on the author and bold him responsible for
uon, underlining should be limited to scientific names of genera_and, typographical or other errors which may pass undetected. In any event,
species which appear in the text. JS'ew names should not _be underlined, where the author secs the proof, proofreading becomes a very important
because the editor will usually mark these with a wavy line to indicate part of his scientific responsibility. The scientific value of bis paper can
boldface. Indications of style or sizes of type for titles, headings, sub- be greatly lessened by unfortunate typographical errors. Such errors
headings, sideheadings, and the like should b_e left to the editor: I~ are sometimes obvious to the reader, but t.bey may be insidious and
general, marks which the author makes mer~ly interfere with_ the editor s wholly misleading.
work though marginal notes as to the relative rank of headings may be In general, the submission of proof to the author is to permit the
belpf:11 • elimination of errors for which the printer is responsible. Author's
Legends and Text Citations to Illustrations. Titles and legends errors are his own responsibility, and some publications charge authors
should be self-explanatory. The manuscript of these titles should for corrections other than printers' errors. Changes in proof are costly
typewritt(ln,double-spaced on separate sheets (several titles oil a sing and therefore should not be made unless necessary or unless t.he author
shoot), and assembled in numerical order at the end of the manuscnp is willing to assume the cost of the change.
following the bibliography. A sbo1~ identifying title ~ay be placed o Proofreading cannot always be done satisfactorily by one person. It
each plate for purposes of identification, hut this title will not be pnn ; is advisable to supplement the personal rending by having someone else •
Usuallyin the process of handling, titles and legends go to the typesette, road slowly from the original manuscript, while the proofreader (pref-
with the rest of the manuscript, whereas illustrations are sent to th erably the author) carefully reads the proof. Special attention should
engraver. The printer may never see the original drawings. . · be given to punctuation,spelling of scientific names, numbers, and dates
The place of insertion of the illustrations should be marked rn th, of all kinds. When corrections are necessary, they should be made '
manuscript and also in the galley proof. Illuetrafions are usually num according to the standard system of proofreaders' marks (Fig. 4 l).
bered starting with each article, but some journals number. plates.con, )'lost authors see only galley proofs of their papers. These arc long
secutively throughout a volume. In any event, a new ".°rics of figu , sheets with the text continuous and not broken into pages. For most
numbers or letters should be used on each plate. Many journals des, journals a galley is the equivalent of about three printed pages. Some
nate figures with Arabic numbers, plates with Roman numerals. publicationsalso submit page proofsto the authors. In such cases proof-
figures should be referred to in the text by number. . readingcannot be restricted to individual words which were corrected in
Revision of the Manuscript. Some few authorshave sufficient maste . the galley proof but must include the whole lino in which the word
of the English language so that they can write directly in final form fo appeared. Modern liootype machines cannot change a single letter in a
publication. Other equally competent scientists find it necessary to , word but must reset the whole line. JJ a word was inserted, it may have
revise page after page not once but many times. T. D. A. Cockerell wus: been necessary to reset several lines or perhaps the remainder of the
an example of the former type of writer, whereas, by his own testimony, paragraph. The author should carefully check everything which has
Charles Darwin ,vas an inveterate reviser. · been reset. Corrected proof should be returned at once to t.!Je editor or
'I'release (1951) recommendscareful readingof the manuscript 10 times,- printer in order to avoid delay in publication. The printing of an entire
each time for one of the following:(1) consistency, (2) sentences, (3) clear·; issueof a periodical may be held up by a single tardy author.
ness, (4) repetition, (5) connectives, (6) euphony, (7) punctuation, (~)', l11ustrations. The object of illust.rat.ions in taxonomic papers is to
style, (9) accuracy, and (10) length. Authors of taxonomic papers se • · present precise, comparative information which c,mnot be so well
dom follow the details of this recommendation,but most papers would expw.;ed in words or which is needed to elucidate the written text.
benefit by more revisionsthan are usually given. It often helps to put _a Thus accuracy, simplicity, and intelligibility are prime considerations.
manuscript aside for a while before the final revision is made. It is ln the prepnrat.ion of illustrntions, adv·ancc consjderntion should Uc given
 LECTURE # 3

THE INTERNATIONAL CODE OF ZOOLOGICAL NOMENCLATURE

The International Code of Zoological Nomenclature (ICZN or ICZN Code) is a widely accepted

convention in zoology that rules the formal scientific naming of organisms treated as animals. The rules

principally regulate:

How names are correctly established in the frame of binominal nomenclature,

Which name has to be used in case of conflicts among various names,

How names are to be cited in the scientific literature.

The Code was first published in 1905, although it has precedents such as Merton's Rules and Strickland's

codes going back to 1843. The 1st edition was first proposed in 1895 in Leiden (3rd International

Congress for Zoology) and published in three languages in 1905 (French, English, German; only French

was official). The 2nd edition was from 1961, the 3rd edition from 1985, the present edition is the 4th

edition (effective since 2000). The Code is elaborated by the Editorial Committee and issued by the

International Commission on Zoological Nomenclature. The Editorial Committee for the 4th edition was

composed of seven persons appointed by the Commission. Such new editions of the ICZN Code are not

democratically approved by those taxonomists who are forced to follow the Code's provisions, neither

do taxonomists have the right to vote for the members of the Commission or the Editorial Committee.

The Code deals with zoological nomenclature, which is defined in the Glossary as

"The system of scientific names for animal taxa and the provisions for the formation, treatment, and use

of those names."
 A Pictorial Key to the Order of Adult Insects

winged wingless
(go to page 64)

front wings hardened, leathery or front wings membranous

parchmentlike at least at the base (go to page 61)

chewing sucking
mouthparts mouthparts

without with front wings leathery front wings of

pincer-like pincer-like at base and uniform texture
cerci cerci membranous at end

DERMAPTERA HOMOPTERA
HEMIPTERA (leafhoppers, planthoppers,
(earwigs) (true bugs) cicadas, spittlebugs)

front wings with A. front wings hard,

branched veins without veins

jumping insects walking insects

A. hind femur enlarged A. hind femur not enlarged
B. tarsi with four or B. tarsi with five segments
fewer segments COLEOPTERA
A (beetles)

B
A

ORTHOPTERA DICTYOPTERA
(crickets, katydids, (roaches, mantids,
grasshoppers) walkingsticks)
60
 continued from key page 60

two wings four wings

wings with few or A. wings usually covered

no scales; without with scales
coiled proboscis B. mouthparts consist of
coiled proboscis
A. pronotum pronotum not
extended over extended over
abdomen abdomen A
B

A LEPIDOPTERA
(butterflies and moths)

very slender wing with no fringe of hairs, or if

fringe of hairs as long as present, not as long as
ORTHOPTERA wing is wide wing is wide
(pigmy grasshoppers)

THYSANOPTERA (go to page 62)

(thrips)

end of abdomen without end of abdomen with

noticeable appendages style or thread-like tail

A. with haltere-like B. with halteres A. style-like tail B. two or three

organs in front of wings behind wings thread-like tails

A
B
B

COLEOPTERA DIPTERA HOMOPTERA EPHEMEROPTERA

(male stylopids) (flies, mosquitoes, (male scales) (mayflies)
gnats, midges)
61
 continued from key page 61

hind wings equal to or hind wings smaller

larger than front wings than front wings
(go to page 63)

no long abdominal appendages abdomen with two or

three thread-like tails

tarsi two or three segmented tarsi with more than three

segments (usually five)

EPHEMEROPTERA
(mayflies)

A. piercing-sucking B. chewing mouthparts

mouthparts

B
B. antennae as long as
body; wings and body
often with scales
A A. antennae shorter than
body; no noticeable scales

HOMOPTERA A
(cicadas, leafhoppers,
planthoppers, spittlebugs) B

HYMENOPTERA
(bees, wasps, ichneumons) TRICHOPTERA
(caddisflies)

PSOCOPTERA
(barklice, booklice)

62
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mouthparts mouthparts at end of beak-like

close to eye structure some distance from eye

MECOPTERA
wings never held (scorpionflies)
flat over abdomen

B. antennae apparently wings held

A. bristle-like flat over abdomen
inconspicuous antennae with several segments

B
A

ODONATA
(dragonflies, damselflies)

hind wings with enlarged hind wings without enlarged

anal area folded fan-like; anal area folded fan-like;
wings tend to curl around wings do not tend to curl
the body lengthwise around the body lengthwise

NEUROPTERA
MEGALOPTERA (lacewings, mantispids,
(dobsonflies, fishflies, owlflies, antlions)
alderflies)

all legs of hind legs modified

walking type for jumping

ORTHOPTERA
(tree crickets)

A. cerci usually long; B. cerci short; with

more than eight segments two to eight segments

B
A

ISOPTERA
PLECOPTERA (termites)
(stoneflies) 63
 continued from key page 60

antennae present antennae absent

legs present legs absent

head and thorax head and thorax

separate fused

DIPTERA
(louse flies, bat flies)

HOMOPTERA COLEOPTERA
(scales) (female stylopoids)

A. collophore present; both collophore and

B. spring-like organ usually present spring-like organ absent

B
long tail-like three tail-like
A appendages absent appendages present

COLLEMBOLA
(springtails)

body not flattened A. body flattened laterally

B. or dorsoventrally

A THYSANURA
(silverfish)
B

(go to page 66)

(go to page 65)

64
 continued from key page 64

body flattened dorsoventrally body flattened laterally

sucking mouthparts no sucking mouthparts

externally visible externally visible

SIPHONAPTERA
A. antennae longer B. antennae shorter (fleas)
than head than head
A
B

antennae longer antennae shorter

than head than head

HEMIPTERA DIPTERA
(true bugs) (louse flies and bat flies)

A. tiny insects; tarsi with B. large insects; tarsi

two or three segments with five segments
A

PSOCOPTERA DICTYOPTERA
(booklice, barklice) (roaches, mantids, walkingsticks)

A. head wider than thorax at B. head narrower than thorax at

point of attachment to thorax point of attachment to thorax
A
B

MALLOPHAGA ANOPLURA
(biting lice) (sucking lice)

65
 continued from key page 64

abdomen and thorax not abdomen and thorax

narrowly joined together narrowly joined together
A

body covered body not covered

with scales with scales

HYMENOPTERA
(ants)

tarsal claws tarsal claws

absent present

LEPIDOPTERA
(female cankerworm) piercing-sucking chewing
mouthparts mouthparts
THYSANOPTERA
(thrips)

cornicles absent A. cornicles usually

present
A

with distinct without distinct

head and eyes head and eyes

HOMOPTERA
(aphids)

HOMOPTERA
(female scales)

HEMIPTERA
(bed bugs)

abdominal forceps present; abdominal forceps absent

entire body rather hard (go to page 67)
and brown colored

DERMAPTERA
(earwigs)

66
 continued from key page 66

A. mouthparts at end of beak-like mouthparts not elongated,

structure some distance from eye close to eyes

cerci present cerci absent

A
MECOPTERA
(scorpionflies) A. antennae longer B. antennae shorter
than one-third of than one-fourth of
body length body length
B
A

body leathery and body soft and

usually grey or pale colored
dark colored

PSOCOPTERA COLEOPTERA
ORTHOPTERA (barklice and booklice) (female stylopids)
(crickets)

three to five tarsal segments

A. basal segment of front tarsi about
same size as ones immediately following

ISOPTERA
(termites)

67