BIRD: linking the biodiversity community
Species is a taxonomic concept used in biology to refer to a population of organisms that are in some important ways similar. The idea of species has a long history. After thousands of years of use, the concept remains central to biology and a host of related fields, and yet also remains at times ill-defined and controversial. There are several main lines of thought in the definition of species:
- A morphological species is a group of organisims that have a distinctive form: for example, we can distinguish between a chicken and a duck because they have different shaped bills and the duck has webbed feet. Species have been defined in this way since well before the begining of recorded history. Although much criticised, the concept of morphological species remains the single most widely used species concept in everyday life, and still retains an important place within the biological sciences, particularly in the case of plants.
- The biological species or isolation species concept identifies a species as a set of actually or potentially interbreeding organisms. This is generally the most useful formulation for scientists working with living examples of the higher taxa like mammals, fish, and birds, but meaningless for organisms that do not reproduce sexually. It distinguishes between the theoretical possibility of interbreeding and the actual likelihood of gene flow between populations. For example, it is possible to cross a horse with a donkey and produce offspring, however they remain seperate species — in this case for two seperate reasons: first because horses and donkeys do not normally interbreed in the wild, and second because the fruit of the union is rarely fertile. The key to defining a biological species is that there is no significant cross-flow of genetic material betwen the two populations.
- A mate recognition species is defined as a group of organisims that are known to recognise one another as potential mates. Like the isolation species concept above, it is not applicable to organisims that do not reproduce sexually.
- A Darwinian or evolutionary or phylogenetic species is a group of organisms that shares a common ancestor; a lineage that maintains its integrity with respect to other lineages through both time and space. At some point in the progress of such a group, members may diverge from one another: when such a divergence becomes sufficiently clear, the two populations are regarded as separate species.
In practice, these definitions often coincide, and the differences between them are more a matter of emphasis than of outright contradiction. Nevertheless, no species concept yet proposed is entirely objective, or can be applied in all cases without resorting to judgement.
The naming of a particular species should be regarded as a hypothesis about the evolutionary relationships and distinguishability of that group of organisms. As further information comes to hand, the hypothesis may be confirmed or refuted.
As a result of the revolutionary (and still ongoing) advance in microbiological research techniques in the later years of the 20th Century, a great deal of extra knowledge about the differences and similarities between species has become available. Many populations which were formerly regarded as seperate species are now considered to be a single biological unit, and many formerly grouped populations have been split. At higher taxonomic levels, these changes have been still more profound.
The isolation species concept in more detail
In general, for large, complex, organisms that reproduce sexually (such as mammals and birds) one of several variations on the isolation or biological species concept is employed. Often, the distinction between different species, even quite closely related ones, is simple. Horses (Equus caballus) and donkys (Equus asinus) are easily told apart even without study or training, and yet are so closely related that they can interbreed after a fashion. Because the result is not usually fertile, they are clearly seperate species.
But many cases are more difficult to decide. This is where the isolation species concept diverges from the evolutionary species concept. Both agree that a species is a lineage that maintains its integrity over time, that is diagnosably different to other lineages (else we could not recognise it), is reproductively isolated (else the lineage would merge into others, given the chance to do so), and has a working intra-species recognition system (without which it could not continue). In practice, both also agree that a species must have its own independant evolutionary history (otherwise the characteristics just mentioned would not apply). The two species concepts differ in that the evolutionary species concept does not make predictions about the future of the population: it simply records that which is already known.
In contrast, the isolaton species concept refuses to assign the rank of species to populations that, in the best judgement of the researcher, would recombine with other populations if given the chance to do so. There are, essentially, two questions to resolve.
First, is the proposed species consistently and reliably distinguishable from other species?
Secondly, is it likely to remain so in the future? There are several broad geographic possibilities.
- The proposed species are sympatric—they occupy the same habitat. Observation of many species over the years has failed to establish even a single instance of two diagnosably different populations that exist in sympatry and have then merged to form one united population. Without reproductive isolation, population differences cannot develop, and given reproductive isolation, gene flow between the populations cannot merge the differences. This is not to say that cross breeding does not take place at all, simply that it has become negligable. Generally, the hybrid individuals are less capable of successful breeding than pure-bred individuals of either species.
- The proposed species are allopatric—they occupy different geographical areas. Obviously, it is not possible to observe reproductive isolation in allopatric groups directly. Often it is not possible to achieve certainty by experimental means either: even if the two proposed species interbreed in captivity, this does not demonstrate that they would freely interbreed in the wild, nor does it always provide much information about the evolutionary fitness of hybrid individuals. A certain amount can be inferred from other experimental methods: for example, do the members of population A respond appropriately to playback of the recorded mating calls of population B? Sometimes, experiments can provide firm answers. For example, there are seven pairs of apparently almost identical marine snapping shrimp (Altheus) populations on either side of the Isthmus of Panama (which did not exist until about 3 million years ago). Until then, it is assumed, they were members of the same 7 species. But when males and females from opposite sides of the isthmus are placed together, they fight instead of mating. Even if the isthmus were to sink under the waves again, the populations would remain genetically isolated: therefore they are now different species. In many cases, however, neither observation nor experiment can produce certain answers, and the determination of species rank must be made on a 'best guess' basis from a general knowledge of other related organisims.
- The proposed species are parapatric—they have breeding ranges that abut but do not overlap. This is fairly rare, particularly in temperate regions. The dividing line is often a sudden change in habitat (an ecotone) like the edge of a forest or the snow line on a mountain, but can sometimes be remarkably trivial. The parapatry itself indicates that the two populations occupy such similar ecological roles that they cannot coexist in the same area. Because they do not crossbreed, it is safe to assume that there is a mechanism, often behavioral, that is preventing gene flow between the populations, and therefore that they should be classified as seperate species.
- There is a hybrid zone where the two populations mix. Typically, the hybrid zone will include representatives of one or both of the 'pure' populations, plus first-generation and back-crossing hybrids. The strength of the barrier to genetic transmission between the two pure groups can be assessed by the width of the hybrid zone relative to the typical dispersal distance of the organisms in question: the dispersal distance of oaks, for example, is the distance that a bird or squirel can be expected to carry an acorn; the dispersal distance of Numbats is about 15 kilometres, as this is as far as young Numbats will normally travel in search of vacant territory to occupy after leaving the nest. The narrower the hybrid zone relative to the dispersal distance, the less gene flow there is between the population groups, and the more likely it is that they will continue on seperate evolutionary paths. Nevertheless, it can be very difficult to predict the future course of a hybrid zone; the decision to define the two hybridizing populations as either the same species or as seperate species is difficult and potentially controversial.
- The variation in the population is clinal — at either extreme of the population's geographic distribution, typical individuals are clearly different, but the transition between them is seamless and gradual. For example, the Koalas of northern Australia are clearly smaller and lighter in colour than those of the south, but there is no particular dividing line: the further south an individual Koala is found, the larger and darker it is likely to be; Koalas in intermediate regions are intermediate in weight and colour. In contrast, over the same geographic range, black-backed (northern) and white-backed (southern) Australian Magpies do not blend from one type to another: northern populations have black backs, southern populations white backs, and there is an extensive hybrid zone where both 'pure' types are common, as are crossbreeds. The variation in Koalas is clinal (a smooth transition from north to south, with populations in any given small area having a uniform appearance), but the variation in magpies is not clinal. In both cases, there is some uncertainty regarding correct classification, but the consensus view is that species rank is not justified in either. The gene flow between northern and southern magpie populations is judged to be sufficiently restricted to justify terming them subspecies (not full species); but the seamless way that local Koala populations blend one into another shows that there is substantial gene flow between north and south. As a result, experts tend to reject even subspecies rank in this case.