What's in a name? A history of taxonomy

http://www.nhm.ac.uk/nature-online/science-of-natural-history/taxonomy-systematics/history-taxonomy/index.html

 

Taxonomy is arguably the world's oldest profession, and naming and classifying what's around us is part of the human condition. Scientific naming began with the Swedish botanist Carl Linnaeus in the eighteenth century. Scientists still use his system, but how much has the science changed from the days when Linnaeus, in a frock coat and with a powdered wig, classified the Earth's plants and animals?

Darwin's theory of evolution has allowed scientists to see diversity as the result of a dynamic process rather than a static picture. This makes the discovery, documentation and study of the diversity of life even more exciting now than in the past. As the conservation of biodiversity becomes ever more important politically, the work of taxonomists has impact not only within the scientific community, but also in society as a whole. Taxonomists today still go into natural habitats and discover new species; it is estimated that scientists have described only ten per cent of the Earth's species, so the task is enormous. Fieldwork involves plant presses, insect nets and hard work getting from place to place, just as it did in the past. Taxonomists house their collected specimens in museums, for their use and for that of future generations of scientists. Morphology is still important in the study of evolutionary patterns, so specimens continue to have a critical role in taxonomy. But today's researchers have at their disposal an armoury of ways of looking at the relationships between species--from electron microscopes for examining the tiniest organisms to DNA sequencers for looking at genes. In this seminar, Sandra Knapp, a botanist at The Natural History Museum, London, explores how taxonomy has changed while retaining its past, making it even more exciting and relevant now than ever before.

 

 

The ancient Greeks and Romans named and classified organisms, particularly those that were useful to them. They had names for medicinal plants and for the animals they hunted and those that they feared. This tradition was carried on in northern Europe, where the naming of plants was done by herbalists. In 1597 the Englishman John Gerard wrote a herbal, one of the first to be written entirely in the English language, which described the plants that were useful to people in his native land. Amongst these was the potato, which in those days was mistakenly thought to come from Virginia, in the United States--it does in fact come from Peru. The herbal also contained a reference to the tomato, which was known as the love apple, or poma amoris.

This is an example of one of the primary problems of the names of organisms. Names get changed with use--Poma amoris, the love apple, started life either as poma di Mori, the apple of the Moors or poma di ori, the golden apple. Over time, the vernacular name became corrupted to poma amoris, and the tomato was presumed to be a powerful aphrodisiac.

At about the same time, in the late sixteenth century, the science of scientific naming started. Plants and animals were given long, polynomial Latin names. At that time Latin was the language of scholarship--a logical choice for scholarly naming. For example, the humble tomato was given the long Latin polynomial Solanum caule inermi herbaceo, foliis pinnatis incisis which means the 'solanum with the smooth stem which is herbaceous and has incised pinnate leaves.'

In the eighteenth century, people working with plants and animals started trying to name things formally, and to use those names consistently. This became increasingly problematic as diversity increased. Initially, only local organisms were known but as other lands, such as the New World, were discovered and explored, the system became complicated. For example, some of the Jesuit priests who accompanied Pizarro when he conquered the Inca empire decided that there must have been two gardens of Eden because the vegetable productions of the Andes were so very different to those of Spain. Many similar theories abounded at the time because it was difficult to explain the diversity being encountered using the Biblical paradigm.

 

Linnaeus and a new way of naming

In the mid-1700s a young Swedish doctor, Carolus Linnaeus, went on a journey to Lapland. Linnaeus was an ingenious and complicated man, and one of the most arrogant people in the history of science. Later in his life he claimed that 'no man has ever transformed science in the way that I have.' His arrogance was, to a great extent, justified.

Linnaeus was a medical doctor, as were all botanists at that time. Until recently, all medicine had been based on herbalism and doctors had to study botany in order to learn their craft. So botanical taxonomy originated with the need of doctors to identify plants correctly. Linnaeus kept plant specimens in his own herbarium. He was fascinated by plants from all over the world and would often travel to examine other people's collections.

Linnaeus was interested not only in plants but in all living organisms. He collected a huge variety of plants and animals and started looking for a system in nature. He was trying to describe all the things that had been put on Earth by the Creator, and therefore approached taxonomy with the tacit assumption that this task was finite.

In the course of his work, Linnaeus did two critically important things for the development of taxonomy. The first was to invent what is known as the sexual system. Instead of looking at the totality of a plant, as would a herbalist, he concentrated on one particular character--and organised all plants according to that character. The characters that he chose were the number of stamens, the male parts, and the number of pistils, the female parts. This was controversial at the time to the extent that he was accused of being a botanical pornographer! His rivals criticised his system as 'loathsome harlotry (scortationes quasi destabiles)'. It was thought that ladies shouldn't deliberately look at the sexual parts of plants. But despite this disapproval the sexual system soon caught on because it was so accessible and straightforward. It represented a democratisation of science in that anyone, not just a specialist, could look at a flower and count the number of male parts and female parts.

In this way Linnaeus made the first of the big advances that transformed the science of taxonomy. The second of his major achievements in this regard occurred by accident. In 1753 he published a book called Species Plantarum which was a list, organised according to the sexual system, of all the plants known to Linnaeus at that time. It was essentially a flora of the world, a field guide to the world's plants. In it he did something that was to completely change the way in which things were named.

 

The birth of the binomial

Until that time, plants were being named with polynomial, Latin names. As more and more species were being discovered, particularly in the New World, those polynomials were getting longer and increasingly unwieldy. So Linnaeus also assigned a 'trivial name' for each plant, a binomial name with only two parts. The idea was that this trivial name would be easy to remember and would trigger the memory of the plant's 'correct' polynomial name.

The binomial names were so much easier to remember that people soon started using them in place of the 'correct' names. Eventually they replaced the polynomial names completely, and became the correct names. The binomial system is the same one we use today--it's how the scientific names of all organisms are constructed. The first part of the name is called the genus and is always capitalised. The second part of the name is called the species epithet and is not capitalised. In the correct format of a scientific name a person's name (sometimes abbreviated) appears after the genus and species name, and this refers to the person who first coined the name. So the scientific name for the raspberry, Rubus idaeus L., can be broken down like this: Rubus (the genus name) idaeus (the species name) Linnaeus (the botanist who coined the name, often abbreviated). Taxonomists have formulated sets of rules for naming; all botanical naming begins with Linnaeus' Species Plantarum in 1753 and animal naming with his tenth edition of Systema Naturae published in 1759.

Animals often reflect their physical attributes, which is useful in aiding memory and providing a basic description. Sometimes those who attribute the names make mistakes; Solanum arboreum sounds as if it should be a tree, but is in fact a tiny shrub, which grows to only about 18 inches. Linnaeus also gave our own species their name, Homo sapiens, which means 'thinking man'--whether or not this name is appropriate is of course a matter of opinion.

Names are also used to honour people. This practice can sometimes result in cumbersome names; for example the plant Rahowardiana wardiana D'Arcy was named by a mentor of mine, after two of his mentors, R.A. Howard and D.B. Ward. Linnaeus was extremely fond of naming plants after people. He named the genus Magnolia after the French Huguenot botanist Pierre Magnol, whom he greatly respected, while the genus Sigesbeckia, a small creeping herb that grows in mud was named for Linnaeus' main critic, Johann Siegesbeck, with whom Linneaus quarrelled.

 

 

Cladistics and collections

Science involves a great deal of expertise. For a long time ideas about the relationships that were believed to exist between groups of plants and animals, that had been long-established by the scientific authorities of the day, were rarely challenged. Whoever was the expert had the last word. In the first decades of the twentieth century, a new generation of taxonomists started to question the accepted dogma, and suggest alternatives.

 

Testing hypotheses

In the 1930s a German entomologist named Willi Hennig started developing an entirely new way of looking at relationships. His work was to change the way taxonomists work and still influences the way research is done today. Hennig proposed the practice of taxonomy as a series of tests of hypotheses. For example, three hypotheses could be applied to a group of three species, say the coelacanth, the lungfish, and a salamander:

 

  • Hypothesis One, the coelacanth and the lungfish are more closely related to each other than either is to the salamander.
  • Hypothesis Two, the salamander and the lungfish are more closely related to each other than either is to the coelacanth.
  • Hypothesis Three, the coelacanth and the salamander are more closely related to each other than either is to the lungfish.

 

A taxonomist would call this a three-taxon statement--the simplest set of hypotheses possible. To uphold or falsify the hypotheses, the taxonomist would amass evidence in support of or against them by looking at the organisms' characteristics--a term shortened to 'characters'.

The characters are each assigned a number. If say, character number one is present in all three species under study, A, B and C, while character two is present in species A and B and absent in species C; the taxonomist can define a group of A+B based on the distribution of character two. Character distribution can be mapped for all the species being studied, and this distribution can reveal groupings of species that are more closely related to each other than to anything else under study. Species and characters can be added to the study, and this method makes taxonomy--the relationship determining part of it--repeatable and shows others how the researcher arrived at their conclusions about taxonomy.

This methodology allows taxonomists to identify two types of groups of species. One type includes all the descendants of a common ancestor and is known as a monophyletic group. The other type, known as a paraphyletic group, includes most, but not all of the descendants of a common ancestor. A third type of group is the polyphyletic group, where the members do not share a common ancestor. With Hennig's developments, the task of taxonomy became to identify the monophyletic groups by defining species' characters unambiguously and plotting their distribution.

 

Cladistics

This new way of doing taxonomy was called cladistics. The word cladistics comes from clade, itself derived from the Greek word for branch. A clade is a group of organisms that share characters, and by extension, a common ancestor. Clades are monophyletic groups. Cladistics meant that scientists no longer relied on the gospel according to scientific authority. Now they were able to look at groups of organisms, study their characters and suggest alternative hypotheses.

In this way, an alternative hypothesis regarding the position of green algae in relation to other plants was suggested. Green algae were always considered to be a discrete and coherent group, all closely related to each other and relatively distantly related to land plants. But analysis of the distribution of certain characters such as type of cell division and microtubule organization revealed that the green algae are an artificial, paraphyletic, group. Despite superficial similarities, some members of the group are much more closely related to land plants than they are to other green algae.

Cladistics challenged the method of doing taxonomy that had been accepted without question for a very long time. In the 1960's when Hennig's book Phylogenetic Systematics (it was published in 1950 in German as Grundzüge einer Theorie der Phylogenetischen Systematik) was first translated into English and came into British and American academic circles, it was extremely controversial. But now, cladistics is widely accepted and through its application, much of the order of the natural world has been revealed. It was cladistic analysis that lead to the discovery that birds and dinosaurs are more closely related to each other than either is to reptiles. Cladistic ideas have become firmly established in all kinds thinking about plants and animals, and the way their relationships are depicted.

 

Collecting and collections

The most important elements of taxonomy are the specimens themselves, many held in museums and collections around the world. As well as relying on collections, taxonomists often go out into the field to find new organisms and collect specimens, and to see plants and animals in their native habitats. When plants are collected they are pressed in newspaper and dried--in the same way as traditional flower pressing. They are then mounted on paper so that they can be studied. Plant specimens from high up in trees are often collected with pole clippers, similar to those used by tree surgeons. They have several extension sections and are extremely efficient, but heavy to carry around. In some parts of the world, scientists hire local people to climb trees for them to access specimens. In other parts of the world, where the locals don't do this, the scientists sometimes learn to do it themselves.

 

Entomologists sometimes collect insects in much the same way as their predecessors did, using butterfly nets with very long poles. They now also use a wide variety of mechanical methods of trapping insects, ranging from simple pitfall traps to more complex light traps and even more complicated methods. The Malaise trap consists of a netting frame that goes all the way to the ground. Insects fly into the netting, and in an attempt to escape they go upwards to find themselves in the peak of the tent. They are naturally attracted to the daylight from the pale top of the tent but on approaching it, drop into a bottle of alcohol. Workers often leave this sort of trap in a remote area for up to a week, and in this way collect a wide diversity of insects, including those which would never be trapped using a butterfly net. The jar of alcohol and insects--a sort of insect 'sludge'--is then sorted to major groups, and specimens are prepared in the usual way, by pinning. Some of the specimens collected in this way are so tiny they need to be glued to a tiny cardboard triangle through which the pin is inserted.

Specimens are prepared in much the same way that they always have been. Butterflies and moths are pinned through the body and put into special spreading boards, then their wings are spread flat out on boards, and then they are dried at extremely low heat in an oven. Using this method the wings stay flat and spread so the colour pattern can be clearly seen.

 

 

Taxonomy and technology

The technological advances of the twentieth century have made the process of examining organisms' characters much easier. For example, microscopes allow us to look at the shapes of chloroplasts, the bodies that convert light into energy, in particular plant cells. But one of the most significant innovations of all has been the scanning electron microscope. This microscope uses an electron beam to produce magnifying power that allows the viewer to count the bristles on an ant's nose, and to look inside the anthers of plants at the pollen, which is often fantastically sculptured and species specific in shape. Even plankton, microscopic marine organisms which were previously thought to have no clearly visible external characteristics, display a huge variety of shapes and characters under the revealing gaze of the scanning electron microscope.

Characteristics can also be studied at the cellular level. Extracts of plant or animal material containing enzymes can be placed within an electric field and different proteins will travel different distances. One can look at the banding patterns of these different proteins and analyze which populations share certain enzymes and how populations or species differ. This technique is called allozyme electrophoresis, and can be extremely useful for detecting hybridization in nature.

Banding patterns formed by chopped up DNA fragments from a fern.

An organism's interaction with its environment can also be helpful to the taxonomist. Behaviour has also become very important in determining taxonomic relationships and looking at how things are related to one another. Many parasitic insects are host-specific and only parasitize a single other species of plant or animal. Studying the behaviour of these interactions can give insect taxonomists clues about relationships and about the evolution of this peculiar way of life--parasitism.

 

The genetic code

One of the greatest technological revolutions of all has been the invention of methods enabling scientists to look directly at the genetic code. The discovery of the shape and function of DNA allows taxonomists to study the genes that code for the organism's characters.

 

Thinking point

Taxonomists of the past would be amazed to learn of the technologies that modern day researchers have at their disposal, and the information that those technologies reveal. Do you think that future developments will deepen our understanding of taxonomy to a level that we could not conceive of today?

All cells have DNA in the nucleus and also in their organelles, the small cellular components such as chloroplasts or mitochondria. All the DNA from plant cells for example, can be extracted, then separated into bands of different density which correspond to the different size genomes from different parts of the cell. One way of looking at DNA is to take a sequence of DNA, say the chloroplast DNA, which is much smaller than nuclear DNA, from the cells and chop it up with special enzymes. The resulting fragments can then be examined in the same way as allozymes are in an electric field. The banding patterns--the distribution of the bands--can be studied to assess the relationships between different plant species.

 

Output of a DNA sequencer

All DNA is made up of four nucleotides (deoxyribonucleotides--hence the abbreviation DNA); adenine (A) guanine (G), cytosine (C) and thymine (T). The DNA double helix consists of pairs of nucelotide bases--A always pairs with T, and G always pairs with C, which link the two strands together. Sequencers read single stranded DNA--like people read a line of text in a book. Each amino acid produced by living cells is coded for by a triplet of nucleotides; for example TTG codes for the essential amino acid leucine, while a triple with a single nucleotide difference, ATG, codes for methionine. Proteins that carry out cell functions are made up of a specific sequence of amino acids. So the sequence of the bases in the DNA has a direct link to the cellular machinery that allows all organisms to develop, grow and live in their environments.

 

DNA sequencing

When single stranded DNA is put through an automated sequencer, the computer produces output that looks like many overlapping lines. Each base shows up as a different colour: thymine is red, guanine is black, adenine is green and cytosine is blue. The peaks correspond to the presence of particular nucleotides and from this a sequence--ATCGTTA for example--can be read. These sequences need to be aligned to make sure they are all in the correct 'reading frame'.

Sequencing, however, is not the end of the story. To determine relationships between organisms is much more complicated than that. Patterns and sequences are sometimes extremely complex and can mislead the researcher. For example, some plants from the tobacco genus Nicotiana show two phylogenetic trees, one constructed using a sequence of DNA from the nucleus and the other a sequence from the chloroplast genome. Chloroplasts in plants are inherited maternally--so the paternal genome never shows up in a tree based on chloroplast sequences. Nicotiana contains many species that are of hybrid origin and are allopolyploid--they have twice the normal number of chromosomes, a set from one of each parent. The trees based on chloroplast and nuclear sequences conflict due to hybridity--if we did not know these species were of hybrid origin, we would be seriously confused!

 

Many patterns seen at the DNA level are still not understood. For example sometimes genes convert one to another or only one parental type is inherited. Nicotiana tabacum, which is cultivated tobacco, is a tetraploid hybrid species, with two sets of chromosome pairs--its two parents are different diploid species, each with one set of chromosome pairs. It inherits the chloroplast genes only from the maternal parent, Nicotiana sylvestris, and so it groups with that species on the tree based on chloroplast sequences. Tobacco groups with its paternal parent, Nicotiana tomentosiformis, on the tree based on nuclear sequences. But nuclear sequence /paternal, chloroplast sequences /maternal is not always the pattern seen. Sometimes, both sets of sequences group a hybrid species with its mother and complicated patterns can result. So sequencing does not provide all the answers as was originally hoped.

 

The genomes of some species have been sequenced completely. The only plant to have had its entire genome sequenced so far is Arabidopsis thaliana, a humble little weed in the mustard family with very simple flowers. The taxonomy of Arabidopsis is not yet well understood, despite us knowing all about this one species. To really determine relationships and study evolutionary patterns in nature, we must know about more than one thing. Ideally one would need three characters, to construct a three taxon hypothesis as described earlier.

 

Botanical taxonomists have long been interested in the development of flowers because many sexual characters, which are found in the flowers, are used to classify plants. Researchers are now looking at the genes that control flower development, at the ways they are turned on and off and how they relate to each other. What is being sought is essentially the genetic basis for what Linnaeus called the sexual system over 300 years ago.

 

 

The future of taxonomy

These are exciting times for taxonomy. New techniques are being developed all the time, and these often throw up as many questions as provide answers.

 

Thinking point

How important is the science of taxonomy? Would the conservation of biodiversity be possible without the information provided by taxonomists?

Taxonomy's challenges for tomorrow relate to the two threads that construct its fabric--to name and identify species and to determine the relationships between them--with the aim of ultimately understanding how nature's wild, wonderful diversity is generated. During the science's history, while new ways of looking at characters and methods of depicting relationships have been developed, many new species have continued to be discovered. Linneaus knew of 23 species of Solanum, the plant genus to which the potato and tomato belong, in 1753. By 1852 the next Solanum monograph, compiled by a French botanist working in Montpellier, Michel-Felix Dunal, described 900 species. Now there are thought to be more than 2000 species of Solanum. And this is just one example from one genus in one part of the world, the New World tropics.

While ideas about relationships are changing, and new species being described, the world around us is constantly changing. It sometimes changes for the better, but more often for the worse. Deforestation and habitat destruction are immense problems on a global scale. For example, Eastern Paraguay is about twice the size of Great Britain and was once largely covered with semitropical rainforest. In 1945, about 50 percent of eastern Paraguay was covered by forest. By 1991 that forested area had shrunk to just 15 percent. Today patches of green are few and far between. Paraguayan biologists and conservation workers have a very hard task ahead to save those few remaining patches--but they are trying and succeeding.

 

In 1992, world leaders attended the Earth Summit in Rio de Janeiro to discuss the problem. The result of that meeting and years of hard work by those interested in the conservation of biodiversity was the United Nations Convention on Biological Diversity, to which about 170 countries, including Great Britain, are signatories. Under the convention (called the CBD) countries are obliged to monitor biodiversity and to promote sustainable development. At first glance, this does not appear to have a huge amount to do with taxonomy, but one of the articles of the Convention requires every signatory country to identify and monitor the biodiversity of the organisms within its borders. How can biodiversity be identified and monitored?

 

Biodiversity and the role of taxonomy

As the world becomes increasingly urban and globalised, people become disconnected from the natural world around them. To conserve diversity we need to also be able to develop some sort of feeling for it. In Great Britain or the United States it would be relatively easy to grab a field guide and to identify the animals and plants we see around us. But tropical ecosystems are much more complicated; such great numbers of species live in the rainforest that it's very hard to identify and catalogue them. But knowing about organisms is part of what makes people care about them. And biodiversity isn't just about the big, easily recognised species, it's also about countless small species that might, individually, seem totally insignificant. Biodiversity is about the totality of things--just like in the first days of taxonomy.

Biodiversity is the term given to the variety of life on Earth and the natural patterns it forms. Present biodiversity is the fruit of billions of years of evolution, shaped by natural processes and, increasingly, by the influence of humans. It forms the web of life of which we are an integral part and upon which we so fully depend. This diversity is often understood in terms of the wide variety of speices on Earth. So far, about 1.75 million species have been identified, mostly small creatures such as insects. Scientists reckon that there are actually about 13 million species, though estimates range from 3 to 100 million. Biodiversity also includes genetic differences within each species, for example, between varieties of crops and breeds of livestock. Chromosomes, genes, and DNA--the building blocks of life--determine the uniqueness of each individual and each species.

Yet another aspect of biodiversity is the variety of ecosystems such as those that occur in deserts, forests, wetlands, mountains, lakes, and agricultural landscapes. In each ecosystem, living creatures, including humans, form a community, interacting with one another and with the air, water and soil around them. It is the combination of life forms and their interactions with each other and with their environment that has made Earth a uniquely habitable place for humans.

Taxonomy's role is vital, because without taxonomists documenting the natural world, we no longer have the fabric that underpins not only biology, but also the science of conservation. Taxonomy is a science that keeps pace with the present but also draws upon the wealth of knowledge accumulated throughout its history. At the same time it is constantly making advances, and looking at organisms and their characters in new and ever more creative ways. Who knows what the next generation of characters will be?

 

Taxonomy in the twenty first century will seem completely different than that of the past--but like good fabric, it still retains both of its threads. Taxonomists do not discard their past, but build upon it in ever more creative ways. While retaining our past, we look forward to increased relevance, both to society at large through our participation in global efforts to conserve the world in which we live, and scientifically, as we refine ever further our hypotheses about life itself. But being a taxonomist today is more difficult than it has ever been--you need to be able to drive both a jeep and an automatic sequencer--but sadly, we are a shrinking community, perhaps as endangered as the organisms we study.

But the future is not bleak--linkages between science and society mean that the role of taxonomy is becoming more appreciated now than ever before. We still have a lot to do, even more so now than when Alfred Russel Wallace penned these somewhat prophetic words:

 

the most perfect collections possible in every branch of natural history should be made and deposited in national museums, where they may be available for study and interpretation.

 

If this is not done, future ages will certainly look back upon us as a people so immersed in the pursuit of wealth as to be blind to higher considerations. They will charge us with culpably allowing the destruction of [that] which we had it in our power to preserve; and while professing to regard every living thing as the direct handiwork and best evidence of a Creator, yet, with a strange inconsistency, seeing many of them perish irrecoverably from the face of the earth, uncared for and unknown.