Module 1: Introduction to Systematics

Learning Objectives:

At the end of the lesson, students should be able to:

a. define the basic concepts of biosystematics, taxonomy, and classification,
b. trace the history of systematic biology, and origin of life,
c. appreciate the importance of systematic biology to other fields, and
d. relate the challenges of systematic biology

Basic Concepts of Biosystematics, Taxonomy, and Classification

Biosystematics

The word systematics is derived from the Latinized Greek word ‘systema’, which means together,
applied to the system of classification developed by Carolus Linnaeus in the 4th edition of his
historical book Systema Naturae in 1735.

In biology, systematics is the study and classification of living things for the determination of
evolutionary relationships; in other words, grouping organisms based on a set of rules (or system),
the science of classifying organisms. Thus, biosystematics may be defined as 'taxonomy of living
populations'. The term biosystematics was coined by Camp and Gilly in 1943.

Blackwelder and Boyden (1952) gave a definition that “systematics is the entire field dealing
with the kinds of animals, their distinction, classification and evolution”.

C. G. Simpson (1961) considers that “systematics is the scientific study of the kinds and
diversity of organisms and of any and all relationships among them”.

The simpler definition by Ernst Mayr (1969), and Mayr and Ashlock (1991) is “systematics is the
science of the diversity of organisms”.

Christoffersen (1995) has defined systematics as “the theory, principles and practice of
identifying (discovering) systems, i.e., of ordering the diversity of organisms (parts) into
more general systems of taxa according to the most general causal processes”.

Systematics includes both taxonomy and evolution. Taxonomy includes classification and
nomenclature but inclines heavily on systematics for its concepts. So, study of systematics includes
a much broader aspect that includes not only morphology and anatomy but also genetics, molecular
biology, behavioural aspects and evolutionary biology.

Kinds of Systematics
Systematics can be divided into two closely related and overlapping levels of classification:
taxonomic and phylogenetic.

Taxonomic classifications (known as the Linnaean System) group living things together based on
shared traits - usually what they look like or what their bodies do.

For example, animals that lay eggs and have scales we call reptiles, and animals that give
births and have fur or hair we call mammals. More specifically, all humans share the same
characteristics and so belong to a group, or taxon, of the genus Homo, and species sapiens.

Phylogenetic classifications use evolutionary relationships of organisms; in other terms, it uses

the taxonomic names, but further group organisms by how evolutionarily related they are to one
another.

The German biologist Ernst Heinrich Haeckel was the first to use the word phylogeny, in
1866, and Darwin used it soon after. It combines the Greek phylos, "race," with geneia, "origin."

By looking at each organism's genes, we know that gorillas (taxonomic term) are more
closely related to humans than they are to cockroaches.

A phenetic classification system relies upon the phenotypes or physical appearances of

organisms.

A genotypic classification compares genes or genomes between organisms.

A polyphasic approach is the most popular which combines aspects of all the systems.
Module 2: Basic Principles of Systematics
Biology and Taxonomy
Learning Objectives:

At the end of the lesson, students should be able to:

a. define the different terms in systematics biology
b. discuss the scope and aims of systematics
c. discuss the component fields of systematics
d. discuss the history of systematics biology.

Taxonomy is the theory and practice of identifying organisms. In fact, taxonomy deals with the
principle involved in the study of classification of organisms.

It is the functional science which deals with identification, nomenclature and classification of
organisms.

The word ‘taxonomy’ is derived from the Greek words taxis (= arrangement) and nomos (= law)
and the term ‘taxonomy’ was coined by A. P. de Candolle in 1813.

There are several stages/levels/periods of taxonomy pertaining to important events in

classification.

I. Alpha () taxonomy

In this stage species are identified and characterized on the basis of gross morphological
features. It is the naming, identifying and characterization of species.

II. Beta () taxonomy

In this stage species are arranged from lower to higher categories, i.e., hierarchic system
of classification. It is arranging species in their natural systems of category.

III. Gamma (γ) taxonomy

In this stage intraspecific differences and evolutionary history are studied. It concerns
the evolutionary sequence, intraspecific variations and the interpretation of organic
diversity.

Diversity and Taxonomy

Alpha Diversity Alpha Taxonomy

It is the biodiversity in a specific area, In alpha taxonomy, analysis of species occurs

ecosystem or community. Alpha diversity is
typically expressed by species richness wherein they are identified, named and
(number of species) in that particular characterized. When species are discovered at
 ecosystem. This can be gauged by taking a this level, they are named as per the Linnaeus
count of the taxa in an ecosystem (species,
genera, families). system of binomial nomenclature. Here
priority is given to the one who publishes his
work first. All problems relating to species are
dealt here.

Beta Diversity Beta Taxonomy

It pertains to the organization of species in
It is the assessment of biodiversity involving
the natural system of hierarchical categories
the comparison of species diversity between
which is carried out based on the shared,
ecosystems. This includes equating the count
observable, structural factors and the
of taxa that are unique to each of the
evaluation of many attributes. Hence, this
ecosystems. It is the rate of change in the
taxonomy is linked with the search for a
composition of species across the habitats in a
natural system of classification. Each of the
community. It provides a quantitative measure
taxa hence would have diagnostic traits which
of the diversity of the communities which
are unique to that particular taxon.
undergo altering environments.

Gamma Diversity Gamma Taxonomy

It is used to refer to the net species richness
across a larger region. It measures the overall
diversity of various ecosystems in a
particular area. Between the component
ecosystems, it is the product of the alpha
diversity of component ecosystems and beta
diversity.
It revolves around the analysis of intraspecific
variations and evolutionary studies. Much
attention is paid to a causal interpretation of
organic diversity — study of speciation. But in
actual practice it is rather difficult to dissociate
them because these overlap and integrate. It is
only in some groups of animals wherein the
taxonomy reaches up to the gamma level few
groups of animals (some vertebrates,
especially the birds and a few insect orders like
Lepidoptera etc.). In most of the groups, it is
till the alpha and beta levels.

The nomenclatural and taxonomic system for botany introduced by Linnaeus and improved by
many generations of later botanists as its first goal to make possible information storage and
retrieval concerning plants and plant products.

While Linnaeus originally believed the systematic pattern he recognized in nature to divine
creation, post-Darwinian botanists have mostly interpreted it as resulting from evolution.

Many botanical taxonomists of today therefore regard systematics as the art of tracing similarities
between taxa, combining them into larger groups, and hypothesizing about their evolution.
While the number of taxonomic features available to Linnaeus was very limited, the gradual
refinement of technical equipment has both widened the scope of morphological studies and
facilitated the introduction of ancillary sciences like anatomy, embryology, serology,
cytotaxonomy, genetics, palynology, and chemotaxonomy.

Those have provided a number of new character sets, the incorporation of which has been
facilitated by methodologies such as numerical taxonomy, cladistics and DNA hybridization.

Biological Classification

Biological classification is a critical component of the taxonomic process. Biological classification

uses taxonomic ranks, in order from most inclusive to least inclusive: Domain, Kingdom, Phylum,
Class, Order, Family, Genus, and Species.

History

Systematics has its origins in two threads of biological science: classification and evolution. The
organization of natural variation into sets, groups, and hierarchies traces its roots to Aristotle and
evolution to Darwin. Put simply, systematization of nature can and has progressed in absence of
causative theories relying on ideas of “plan of nature,” divine or otherwise. Evolutionists (Darwin,
Wallace, and others) proposed a rationale for these patterns. This mixture is the foundation of
modern systematics.
Aristotle

Systematics as classification (or taxonomy) draws its Western origins from Aristotle. A student of
Plato at the Academy and reputed teacher of Alexander the Great, Aristotle founded the Lyceum
in Athens, writing on a broad variety of topics including what we now call biology. To Aristotle,
living things (species) came from nature as did other physical classes (e.g. gold or lead).
Today, we refer to his classification of living things (Aristotle, 350 BCE) that show similarities
with the sorts of classifications we create now. In short, there are three features of his
methodology that we recognize immediately: it was functional, binary, and empirical.

Aristotle’s classification divided animals (his work on plants is lost) using functional features as
opposed to those of habitat or anatomical differences: “Of land animals some are furnished with
wings, such as birds and bees.” Although he recognized these features as different in aspect, they
are identical in use.

Features were also described in binary terms: “Some are nocturnal, as the owl and the bat; others
live in the daylight.” These included egg- or live-bearing, blooded or non-blooded, and wet or dry
respiration.

An additional feature of Aristotle’s work was its empirical content. Aspects of creatures were
based on observation rather than ideal forms. In this, he recognized that some creatures did not fit
into his binary classification scheme: “The above-mentioned organs, then, are the most
indispensable parts of animals; and with some of them all animals without exception, and with
others animals for the most part, must needs be provided.” Sober (1980) argued that these
departures from Aristotle’s expectations (Natural State Model) were brought about (in Aristotle’s
mind) by errors due to some perturbations (hybridization, developmental trauma) resulting in
“terata” or monsters. These forms could be novel and helped to explain natural variation within
his scheme.

Blooded Animals
Live-bearing Animals
Humans
Other Mammals
Egg-laying Animals
Birds
Fish
Non-blooded Animals
Hard-shelled sea animals: Testacea
Soft-shelled sea animals: Crustacea
Non-shelled sea animals: Cephalopods
Insects
Bees
Dualizing species (potential “terata” errors in nature)
Whales, seals, and porpoises – in water, but bear live young
Bats – have wings and can walk
Sponges – like plants and like animals
Aristotle clearly had notions of biological progression (scala naturae) from lower (plant) to higher
(animals through humans) forms that others later seized upon as being evolutionary and we reject
today. Aristotle’s classification of animals was neither comprehensive nor entirely consistent, but
was hierarchical, predictive (in some sense), and formed the beginning of modern classification.

Theophrastus

Theophrastus succeeded Aristotle and is best known in

biology for his Enquiry into Plants and On the Causes
of Plants. As a study of classification, his work on ivy
(κιττo ́ς) discussed extensively by Nelson and Platnick
(1981), has been held to be a foundational work in
taxonomy based (in part at least) on dichotomous
distinctions (e.g. growing on ground versus upright) of
a few essential features.

Theophrastus distinguished ivies based on growth form

and color of leaves and fruit. Although he never
presented a branching diagram, later workers (including Nelson and Platnick) have summarized
these observations in a variety of branching diagrams (Va ́csy, 1971) (Fig. 1.1).

Pierre Belon

Trained as a physician, Pierre Belon,

studied botany and traveled widely in
southern Europe and the Middle East. He
published a number of works based on
these travels and is best known for his
comparative anatomical
representation of the skeletons of
humans and birds (Belon, 1555) (Fig.
1.2).

Carolus Linnaeus

Carolus Linnaeus (Carl von Linn é ) built on Aristotle and created a classification system that has
been the basis for biological nomenclature and communication for over 250 years. Through its
descendants, the current codes of zoological, botanical, and other nomenclature, his influence is
still felt today. Linnaeus was interested in both classification and identification (animal, plant, and
mineral species), hence his system included descriptions and diagnoses for the creatures he
included. He formalized the custom of binomial nomenclature, genus and species we use today.

Linnaeus was known, somewhat scandalously in his day, for his sexual system of classification
(Fig. 1.3). This was most extensively applied to plants, but was also employed in the classification
of minerals and fossils. Flowers were described using such terms as visible (public marriage) or
clandestine, and single or multiple husbands or wives (stamens and pistils). Floral parts were even
analogized to the foreskin and labia.

Nomenclature for many fungal, plant, and other eukaryote groups is founded on the Species
Plantarum (Linnaeus, 1753), and that for animals the 10th Edition of Systema Naturae (Linnaeus,
1758). The system is hierarchical with seven levels reflecting order in nature (as opposed to the
views of Georges Louis Leclerc, 1778 [Buffon], who believed the construct arbitrary and natural
variation a result of the combinatorics of components).

• Imperium (Empire)—everything
• Regnum (Kingdom)—animal,
vegetable, or mineral
• Classis (Class)—in the animal
kingdom there were six
(mammals, birds, amphibians,
fish, insects, and worms)
• Ordo (Order)—subdivisions of
Class
• Genus—subdivisions of Order
• Species—subdivisions of Genus
• Varietas (Variety)—species
varieties or “sub-species.”

The contemporary standard hierarchy includes seven levels: Kingdom, Phylum, Class, Order,
Family, Genus, and Species, although other levels are often created as needed to describe diversity
conveniently (e.g., McKenna and Bell, 1997).
Georges Louis Leclerc, Comte de Buffon

Georges Louis Leclerc, Comte de Buffon, began his scientific career in mathematics and
probability theory. He was appointed director of the Jardin du Roi (later Jardin des Plantes),
making it into a research center.

Buffon is best known for the encyclopedic and massive Histoire naturelle, g ́en ́erale et
particuli`ere (1749–1788). He was an ardent anti-Linnean, believing taxa arbitrary, hence
there could be no preferred classification. He later thought, however, that species were real (due
to the moule int ́erieur—a concept at the foundation of comparative biology). Furthermore, Buffon
believed that species could “improve” or “degenerate” into others, (e.g. humans to apes) changing
in response to their environment. Some (e.g. Mayr, 1982) have argued that Buffon was among the
first evolutionary thinkers with mutable species. His observation that the mammalian species of
tropical old and new world, though living in similar environments, share not one taxon, went
completely against then-current thought and is seen as the foundation of biogeography as a
discipline (Nelson and Platnick, 1981).

Jean-Baptiste Lamarck

Jean-Baptiste Lamarck (who coined the word

“Biologie” in 1802) believed that
classifications were entirely artificial, but
still useful (especially if dichotomous). His
notion of classification is closer to our modern
keys (Nelson and Platnick, 1981). An example
of this comes from his Philosophie zoologique
(Lamarck, 1809), with the division of animal
life into vertebrates and invertebrates on the
presence or absence of “blood” (Fig. 1.4(a)).

Lamarck is best known for his theory of

Transmutation (Fig. 1.4(b))—where species
are immutable, but creatures may move through one species to another based on a motivating force
to perfection and complexity, as well as the familiar “use and dis-use.” Not only are new species
created in this manner, but species can “re-evolve” in different places or times as environment
and innate drive allow.

Georges Cuvier

The hugely influential L ́eopold Chr ́etien Fr ́ed ́eric Dagobert “Georges” Cuvier divided animal
life not into the Scala Naturae of Aristotle, or two-class Vertebrate/ Invertebrate divide of
Lamarck, but into four “embranchements”: Vertebrata, Articulata, Mollusca, and Radiata
(Cuvier, 1812). These branches were representative of basic body plans or “archetypes” derived
(in Cuvier’s view) from functional requirements as opposed to common genealogical origin of
structure. Based on his comparative anatomical work with living and fossil taxa, Cuvier believed
that species were immutable but could go extinct, (“catastrophism”) leaving an unfillable hole.
New species, then, only appeared to be new, and were really migrants not seen before. Cuvier
established the process of extinction as fact, a revolutionary idea in its day.

Étienne Geoffroy Saint-Hilaire

Although (like Lamarck), the comparative anatomist E tienne ́ Geoffroy Saint- Hilaire is
remembered for his later evolutionary views, Geoffroy believed that there were ideal types in
nature and that species might transform among these immutable forms. Unlike Lamarck, who
believed that the actions of creatures motivated transmutation, Geoffroy believed environmental
conditions motivated change. This environmental effect was mediated during the development
of the organism. He also believed in a fundamental unity of form for all animals (both living and
extinct), with homologous structures performing similar tasks. In this, he disagreed sharply with
Cuvier and his four archetypes (embranchements), not with the existence of archetypes, but with
their number.

Johann Wolfgang von Goethe

With Oken and Owen, Goethe was one of the foremost “ideal morphologists” of the 19th century
in that he saw universal patterns underlying the forms of organisms. He coined the term
“Morphology” to signify the entirety of an organism’s form through development to adult
as opposed to “gestalt” (or type— which was inadequate in his view). This is similar to Hennig’s
concept of the “semaphoront” to represent the totality of characters expressed by an organism over
its entire life cycle.

Goethe applied these ideas to the comparative morphology and development of plants (von
Goethe, 1790) as Geoffroy did to animals, creating morphological ideals to which all plants
ascribed. He claimed, based on observation, that archetypes contained the inherent nature of a
taxon, such as “bird-ness” or “mammal-ness.” This ideal was not thought to be ancestral or
primitive in any way, but embodied the morphological relationships of the members of the group.

Lorenz Oken

Oken was a leader in the “Naturphilosophie” (Oken, 1802) and an ideal morphologist. In this,
he sought general laws to describe the diversity in nature through the identification of ideal
forms. One of the central tenets of the Naturphilosophie was that there were aspects of natural law
and organization that would be perceived by all observers. He applied this to his classification of
animal life, and created five groups based on his perception of sense organs.

1. Dermatazoa—invertebrates
2. Glossozoa—fish (with tongue)
3. Rhinozoa—reptiles (with nose opening)
4. Otozoa—birds (with external ear)
5. Ophthalmozoa—mammals (nose, ears, and eyes).

Oken is also known for his attempts to serially homologize vertebral elements with the vertebrate
skull, suggesting fusion of separate elements as the main developmental mechanism. Although
falsified for vertebrates, the idea found ground in discussions of the development of the arthropod
head.

Richard Owen

Richard Owen was a vertebrate comparative

anatomist known for his role in founding the
British Museum (Natural History), the
definitions of homology and analogy, and his
opposition (after initial favor) to Darwinian
evolution. Owen (1847) defined a homologue as
“The same organ in different animals under every
variety of form and function.” Analogy was, in his
view, based on function, “A part or organ in one
animal which has the same function as another part
or organ in a different animal.”

Owen derived the general archetype for

vertebrates based (as in Oken) on the serial
homology of vertebral elements (Fig. 1.5).

Owen’s notion of homology and archetype was

tightly connected with the component parts that
made up the archetype—the homologues.

Charles Darwin

To Aristotle, biological “species” were a component of nature

in the same way that rocks, sky, and the moon were. Linnaeus
held that the order of natural variation was evidence of divine
plan. Darwin (1859b) brought the causative theory of
evolution to generate and explain the hierarchical distribution
of biological variation. This had a huge intellectual impact in
justifying classification as a reflection of genealogy for the
first time, and bringing intellectual order (however
reluctantly) to a variety of conflicting, if reasonable,
classificatory schemes.

The genealogical implications of Darwin’s work led him to

think in terms of evolutionary “trees,” (Fig. 1.6), the
ubiquitous metaphor we use today. The relationship between
classification and evolutionary genealogy, however, was not
particularly clarified (Hull, 1988). Although the similarities
between genealogy and classification were ineluctable,
Darwin was concerned (as were many who followed) with representing both degree of
genealogical relationship and degree of evolutionary modification in a single object. He felt quite
clearly that classifications were more than evolutionary trees, writing that “genealogy by itself
does not give classification” (Darwin, 1859a).

How to classify even a hypothetical case

of genealogy (Fig. 1.7)? Darwin’s Figure
presents many issues—ancestral species,
extinction, different “degrees of
modification,” different ages of taxa. As
discussed by Hull (1988), Darwin gave no
clear answer. He provided an intellectual
framework, but no guide to actually
determining phylogenetic relationships or
constructing classifications based on this
knowledge.

Darwin transformed Owen’s archetype

into an ancestor. Cladistics further
transformed the ancestor into a median.

Stammb ̈aume

Haeckel (1866) presented the situation in a

graphical form (Fig. 1.8), including both
genealogical relationships (as branches),
degrees of modification (distance from root),
and even Aristotle’s Scala Naturae beginning
with Monera at the root and progressing
through worms, mollusks, echinoderms,
tetrapods, mammals, and primates before
crowning with humans. In his 1863 lecture,
Haeckel di- vided the scientific community
into Darwinians (progressives) and
traditionalists (conservatives): “Development
and progress!” (“Entwicklung und
Fortschritt!”) versus “Creation and species!”
(“Scho ̈pfung und Species!”). He even coined
the word “Phylogeny” (Haeckel, 1866) to
describe the scheme of genealogical
relationships. Haeckel felt that paleontology
and development were the primary ways to
discover phylogeny (Haeckel, 1876).
Morphology was a third leg, but of lesser
importance. Bronn (1858, 1861) also had a tree
like representation and was the translator of Darwin into the German version that Haeckel read
(Richards, 2005). Bronn found Darwin’s ideas untested, while Haeckel did not.
August Schleicher constructed linguistic trees as Darwin had biological. A friend of Haeckel,
Schleicher “tested” Darwin with language (Schleicher, 1869). Interestingly, he thought there
were better linguistic fossils than biological, and hence they could form a strong test of Darwin’s
ideas.

Contribution of Systematics to Other Fields of Biology

There is immense contribution of systematics to other branches of biology and to humankind as a

whole. The study of organic diversity, prior to the rise of genetics, was entirely carried out by
systematists. Virtually all major evolutionary problems were brought to the forefront and solved
by systematists and even today new evolutionary problems are revealed by them. A similar
dependence on systematics exists in other areas of science.

Some of the noteworthy contributions are:

A. Patterned Diversity:
Organic diversity is not chaotic but patterned, revealing all sorts of regularities. The task of the
systematics is to discover the true nature and causation of these patterns. Rodents and lagomorphs
(rabbit etc.) have rootless gnawing incisors.

It is the task of the systematists to find out the reasons behind the causation of such similarity. Is
it due to common descent or due to adaptation to an equivalent adapting zone? All evolutionary
processes or phenomenon can only be studied reliably if a sound taxonomic foundation is present.

B. Applied Biology:
The contribution of systematics, both directly and indirectly, has been noticed in applied sciences
like medicine, public health, agriculture, conservation, management of natural sources etc.

(a) Epidemiology:
The famous case of epidemiology of malaria, caused by Anopheles maculipennis, was
reported throughout Europe, yet malaria was restricted to local districts. Large amounts of
money were wasted and yet no one understood the connection between the distribution of
the mosquito and that of malaria. Finally, the key to the problem was provided by
systematics.

A. maculipennis was found to have a number of sibling species with different habitat
preferences and breeding habits. Only some of the maculipennis complex was responsible
for the transmission of malaria and that too in a given area. This information allowed
control measures to be directed to those spots where they were effective.

(b) Biological Control:

Another example is the biological control of insect pests. For a brief period it appeared that
biological control had become obsolete owing to the success of chemicals (pesticides etc.),
Because of the adverse effect of these chemicals on human health applied entomologists
had to revert increasingly to biological control. The work of the taxonomist or systematics
is to find out the parasites that attack the various pests (and at which stage of the pest) and
thereby the application of these parasites can bring about a successful control of these pests.

(c) Wildlife Management:

We are aware that deforestation and indiscriminate killing of animals have resulted in dis-
balance of nature. Many species of plants and animals have become extinct and many are
on the road to extinction. Taxonomists have contributed to environmental protection by
identifying all such endangered plants and animals that are endangered due to anthro-
pogenic causes.

(d) Determination of sequential events:

Dating of rocks is generally done through radioactive decay, but sedentary rocks can be
dated through their enclosed flora and fauna. The taxonomist plays a vital role by identi-
fying such flora and fauna and formulating a clear picture of the geological events. Such
work has often been of great value, particularly in the success of oil industries in America.

(e) Environmental problems:

Various environmental problems have been successfully tackled by systematists.
Environmental problems such as pollution, leads to the persistence of certain non-
biodegradable pollutants in the environment. Tracing the movement of these pollutants in
the food chain requires the identification of the various species that constitute the biotic
community. Present advances in systematics have revealed that various species of plants
and animals act as indicators of pollution. The identification of such species in a particular
location helps in the rapid and inexpensive monitoring of the pollutants that degrade the
concerned environment.

(f) Soil fertility:

 Many animals and microbes play important role in increasing soil fertility. It is essential to
locate such species and detect the role they play in soil fertility, so that such species can be
utilised in agricultural management practices.

(g) Introduction of commercially important species:

Based on sound systematics various commercially important species have been established
in India and other countries. Correct identification of such species and the role they play in
the ecosystem provides useful information for their introduction.

Apis mellifera (the Italian honey bee), Cyprinus carpio (common carp) etc. are some exotic
species that have been successfully introduced in India. This has been made possible only
through correct identification by the systematists.

C. Theoretical Biology:
Population thinking has come into biology due to taxonomy and systematics. One of the two roots
of population genetics is systematics. The problem of multiplication of species was solved by
them.

It was systematics who continued to uphold the importance of natural selection when early
Mendelian’s thought that mutation had eliminated the role of natural selection as an evolutionary
understanding of mimicry, provided the first clear proof of the importance of natural selection in
evolution.

The development of ethology and the study of the phylogeny of behaviour were developed by the
taxonomist and naturalists. Thus, systematics has contributed to a healthy balance in biological
science.

Challenges: Biosystematics, Can It Deliver?

It is often argued that we cannot hope to conserve and manage our biological diversity effectively
if we do not know the makeup of the Earth's biological constituents (e.g. May 1988, 1994;
Wilson 2004; Wheeler et al. 2012). However, knowing the constituent taxa is of limited use,
unless there is basic understanding of where those taxa occur in relation to landscape features and
tenure (i.e., their spatial distribution). In other words, it is equally important to map the
distributions of species as it is to recognize them formally (Wheeler et al. 2012). Systematic
monographs and taxonomic revisions (and the museum collections on which they are based)
provide these two basic elements: the circumscription of species, and the geographical distribution
of those species.

Two major challenges for biosystematics are therefore to: (1) systematically catalogue and map
the Earth's known species, and (2) discover, describe and document new species (the as-yet-
unknown or missing species). The former task includes resolving synonymy of the numerous
cases where species have been described more than once, redescribing and diagnosing taxa
currently placed in synonymy that prove to be specifically distinct, and resolving other problems
of nomenclature (e.g. homonyms). Both tasks are needed in order to optimise conservation effort
of the world's biodiversity (Scheffers et al. 2012; Costello et al. 2013; Stork & Habel 2014). Other
challenges for biosystematics are to: (3) reconstruct the evolutionary history or tree of life and
establish the phylogenetic relationships among species and (4) incorporate phylogenetic
diversity (taxonomic distinctiveness) as a component of biodiversity into conservation
planning. The rationale behind the latter task is that phylogenetic diversity captures the
evolutionary legacy or distinctiveness of a taxon or taxonomic group, but so far this has apparently
contributed little towards practical nature conservation (Winter et al. 2013). Isaac et al. (2007),
however, have developed a method for incorporating both phylogenetic diversity and extinction
risk for threatened species that are both ‘evolutionarily distinct and globally endangered’ (i.e.
EDGE species), which can be used to generate global priority lists for conservation. Moreover,
there is a sense of urgency to complete these challenges because of increasing levels of extinction
against a background of declining systematic expertise (Wilson 1987, 1992, 2004; Dirzo &
Raven 2003; Mallet & Willmott 2003; Yeates & Raven 2004; Oliver & Lee 2010; Probert 2010;
Maddison et al. 2012; Wheeler et al. 2012; Yen & New 2013). Not only are professional
taxonomic specialists across all biota declining, but this expertise is mismatched against the
species richness of taxa and the geographical location of biodiversity (e.g. Gaston & May 1992;
New 1996b; Godfray 2002; Tautz et al. 2003; Wheeler et al. 2004; Yeates 2009; Probert 2010;
Boero 2010; Löbel 2014).

The general view is that the task of describing and naming all of the world's species is probably
unachievable (New 1993, 1996b; Stork et al. 1996; Wheeler et al. 2012; Costello et al. 2013) and
that complete inventories of invertebrates even at relatively small spatial scales are logistically
impossible (Harvey et al. 2011). Kitching (1993a) estimated that between 50–80% of species
among some Australian rainforest canopy taxa are unnamed, and Yeates et al. (2003) and
Austin et al. (2004) estimated that at least 75% of the Australian Insecta (c. 205 000 known
species) have not been formally described or are yet to be discovered. Kristensen et al. (2007)
estimated that only about one third of the global Lepidoptera (c. 160 000 species described so far)
has been systematically catalogued, with around 1000 new species being described annually in
recent years. Insect conservation in Australia is thus a formidable task, being carried out by a small
number of informed specialists using incomplete taxonomic and biological knowledge
(Samways 2007; New & Yen 2013; New & Samways 2014). Not surprisingly, insects receive low
priority on the conservation agenda in all state agencies, and setting priorities for insect
conservation is difficult and sometimes controversial.

Several solutions have been proposed to overcome this enormous taxonomic impediment for
insect conservation biology. One proposal is that taxonomic attention should be diverted
towards small geographical regions known to support high levels of endemism where
detailed inventories may be possible (Samways 2007), or focus on a limited set of ‘priority’
taxa (Ehrlich 1992, 2003; Kitching 1993a,b; New 1993; Stork et al. 1996; Samways 2007;
Kitching & Ashton 2013). These proposals may be preferable than attempting to catalogue the
entire biological diversity of the planet because most species will almost certainly go extinct before
they are even discovered, let alone named and classified. Ehrlich (1992, 2003) argued that this
prioritised taxon set be used for inventory and monitoring – to design conservation area networks,
to evaluate ecosystem health and to gauge the effectiveness of management and persistence of
ecological and evolutionary processes. Given finite resources for taxonomic research, which
groups should be prioritized for conservation biology? The answer to this question will require
careful consideration – we suggest at least two of the essential criteria listed in Table 1 be
considered for selection of taxonomic groups: (1) the taxon is reasonably well known
taxonomically (i.e. total inventory is estimated to be 90% complete, and/or morphospecies have
been circumscribed through the availability of parataxonomists and well-curated reference
collections); and, (2) the taxon is known to be informative as bioindicators. Terrestrial insect
and other invertebrate groups in Australia that fit both criteria include landsnails (Stanisic 1999;
Slatyer et al. 2007; Braby et al. 2011), some spiders (Churchill 1997; New 1999), dragonflies
(Brooks 1996; Clausnitzer et al. 2009), dung beetles (Hill 1996; Spector 2006; Nichols &
Gardner 2011; Monteith 2015), ground beetles (Grimbacher et al. 2007; Michaels 2007; but see
New 1998), butterflies, some diurnal moths, some nocturnal macromoths (New 1996b, 2011;
Kitching et al. 2000; Lomov et al. 2006; Ashton et al. 2011) and ants (Andersen & Majer 2004;
Andersen et al. 2004).

The second proposal is to accelerate the rate at which new species are discovered and
described, at least by an order of magnitude (Wiens 2007; Bacher 2012; Wheeler et al. 2012;
Costello et al. 2013; Novotny & Miller 2014). This proposal includes several initiatives, such as
increasing the efficiency of the publication process (e.g. moving towards early view publication,
online registration through ZooBank (Yeates 2009)); open-access to comprehensive online
taxonomic databases and literature, and development of other information technology platforms
or bioinformatics (e.g. Global Biodiversity Information Facility, Integrated Taxonomic
Information System, Tree of Life, All Taxa Biodiversity Initiative); increased resources for
specimen collection and curation, integrated with digital imaging and DNA barcoding; and,
perhaps most importantly, training of more taxonomists and better coordination between amateur
and professional scientists. Boero (2010) recommended that for the science of taxonomy to survive
it must become a multidisciplinary science, integrating molecular biology and computational
science with traditional morphological approaches. We suggest that a fourth dimension be added
to raise the profile/relevance of biosystematics – integration and collaboration with conservation
biologists, conservation agencies and relevant community groups (e.g., citizen science
programmes).

The cost in terms of the amount of both time and effort required to complete the taxonomic
inventory (should that be possible) is debatable, but clearly considerably more investment is
needed than at present (Wheeler et al. 2012; Costello et al. 2013). Although there has been a global
increase in the number of people describing species (Joppa et al. 2011b; Bacher 2012;
Costello et al. 2013), the professional workforce of traditional taxonomists has been in a steady
state of decay for several decades, at least in Australian entomology (particularly those associated
with premier natural history museum collections such as the Australian National Insect
Collection). In our opinion, many of the people who describe new species or who have their name
attached to publications describing new species, are not necessarily experts nor ‘good’
taxonomists. Many new generation taxonomists have little or no formal training in the principles
of the International Code of Zoological Nomenclature (ICZN) (International Commission on
Zoological Nomenclature 1999), have limited understanding of species concepts and the
importance of types, do not prepare formal diagnoses, have limited knowledge of morphology,
and typically publish works of single species rather than the comprehensive revisions of higher
taxa (e.g. genera, families) that require high levels of taxonomic expertise.

Some proposals to accelerate taxonomic description, such as web-based taxonomy and/or side-
stepping peer review (Scoble 2004; Maddison et al. 2012), are not without problems and
inevitably will produce poor quality science, resulting in long-term taxonomic chaos that will
eventually need resolution. Moreover, taxonomists who reveal the existence of unrecognized taxa
on the internet run the risk of having those candidate taxa described by unscrupulous individuals
using non-refereed, and often in privately published, works (Oliver & Lee 2010). The rules of the
ICZN currently do not regulate the quality of taxonomic research, just the use of taxonomic names.
We contend that predictions that the global inventory (recent estimates vary from c. 10 million
species (Chapman 2009) to 3.7–6.1 million species (Hamilton et al. 2010, 2013) and 6.8 million
species (range 5.9–7.8 million) for terrestrial arthropods (Stork et al. 2010,2015)) can be
completed within five decades by boosting taxonomic capacity (Wheeler et al. 2012;
Costello et al. 2013) are overly optimistic. Extrapolations from species discovery curves fail to
appreciate that the relationship between the cumulative number of species over time is non-linear.
In well-known groups, a disproportionate amount of time/effort is required to document the as-
yet-unknown species (Joppa et al. 2011b), largely due to the diminishing pool of missing species
that are harder to ‘catch’, refinement of species boundaries (hypothesis testing) as new evidence
becomes available, and inevitable synonymy. The inventory of Australian butterflies, for example,
is not expected to be completed until about the year 2090 (i.e. 75 years from now), despite the fact
that the pool of missing species is estimated to be only around 9% of the total fauna (Fig. 3).
Historically, the taxonomy of Australian butterflies as a science has been driven by private
researchers comprising largely amateur experts using traditional methods; this is still very much
the case today (Moulds 1999; Edwards et al. 2001). Clearly, if the inventory of even such well-
known groups is to be completed within a few decades the rate of taxonomic resolution needs to
change drastically.

Review Questions:

1. Differentiate among biosystematics, taxonomy, and classification; cite the basic

underlying concepts.
2. Tracing the history, who are the notable people, and what are their significant
contributions in the development of systematic biology?
3. Discuss the importance of systematic biology to other fields of biology.
4. What are the challenges of systematic biology that post problems in the
scientific and vigorous studies of biological diversity?