Preview of Concepts Taxonomy is the science dealing with description, identification, nomenclature, and classification of living things.

A dichotomous key is a tool that allows users to identify items or organisms in a systematic and reproducible fashion. Dichotomous keys may be used in a variety of situations, such as for identifying rocks and minerals as well as for identifying unknown organisms to some taxonomic level (e.g., species, genus, family, etc.). What makes these keys distinctive is that they are ordered in such a way that a series of choices is made that leads the user to the correct identity of the item they are looking at. Dichotomous means, divided into two parts. !herefore, dichotomous keys always offer two choices for each step, each of which describes key characteristics of a particular organism or group of organisms. "magine that you are trying to construct a dichotomous key for the students in your biology laboratory class. !o do this you would need to make a list of some characteristics possessed by the members of the group, such as gender, hair color, height, type of clothing (#eans, dress pants, shorts, dress), whether or not they wear glasses, and so forth. $ou would then construct your key by setting up a series of bifurcating characteristics, starting with the most general characters and moving to the most specific such that, in the end, each member of the group can be clearly identified. %or example&
'. (ex female)))* (ex male)))+ *. ,air color red)))(ally ,air color not red)))-. ,air color blonde))).ulie ,air color black)))/ /. 0lasses worn)))Deanna 0lasses not worn)))1eslie +. (hoes high)top sneakers))).oseph (hoes not high)tops)))2 2. ,air color blonde))3ichael ,air color brown))David

!hus if we are looking at a male student with brown hair wearing cowboy boots we would start at step '. 4ecause he is male, we would be instructed to go to point +. At point + we would establish that his shoes are not high)tops, and so be directed to step 2, where the fact that our specimen has brown hair would allow us to determine that we are looking at David.

Exercise 1

"magine that you meet a blonde male member of this class wearing teva sandals. Who is it5 "magine that his girl friend has black hair but doesn6t wear glasses. Who is he dating5

Exercise 2 7se the dichotomous key to the principle orders insects provided to determine the identity of the following organisms&

8lick here for specimen '. 8lick here for specimen *. 8lick here for specimen -.

Systematic Approaches to Phylogeny hat is Systematics! (ystematics is a discipline within biology whose goal is the determination of the evolutionary history and relationships among organisms (termed phylogeny) and then the use of that phylogeny in classifying organisms. !o achieve that goal, a systematist will utili9e evidence from a wide variety of sources including paleontology, embryology, morphology, behavior, and molecular biology. :ver the last few centuries systematists have developed a number of different approaches for trying to show the relationships of organisms. As a result, different schemes have emerged for ordering these relationships into a classification scheme. !he main approaches in the last +; or so years may be classified as follows&

Phenetic (or numerical taxonomy) Cladistic (or phylogenetic) Evolutionary (or synthetic)

Phenetics "n the old days (before the twentieth century), classification was largely carried out by gentlemen scientists who based their identifications of organisms upon a few important characteristics and a gut feeling developed during many years experience. <henetic classification methods were invented by systematists who thought that this type of classification scheme was too sub#ective. !hese scientists argue that taxonomy should not be based on a few characteristics arbitrarily #udged to be important, but rather upon the degree of overall similarity between organisms. !hus they collect data on as many characteristics of the organisms being classified

as possible. =ach organism is then compared with every other for all characters measured, and the number of similarities (or differences) is calculated. !he organisms are then clustered in such a way that the most similar are grouped close together and the more different ones are linked more distantly. 4ecause of the enormous amount of data involved in this process, these calculations are usually made using specially programmed computers'. !he taxonomic clusters (phenograms) that result from such an analysis do not necessarily reflect genetic similarity or evolutionary relatedness. <heneticists would argue that this is :>, because nobody really knows the true evolutionary histories of these organisms and at least this system is ob#ective. ,owever, the lack of evolutionary significance in phenetics has meant that this system has had little impact on animal classification, and as a conse?uence interest in and use of phenetics has been declining in recent years.
1

!he dependence of <henetics on the use of computers made it particularly unpopular with some old)school biologists. <aul =hrlich, one of the early converts to <henetics, was reportedly once asked by an irate taxonomist at a conference, $ou mean to tell me that taxonomists can be replaced by computers5 =hrlich responded, @o, some of you can be replaced by an abacusA

Cladistics "Phylogenetic Systematics# <henetics has been critici9ed because phenograms resulting from such analyses do not necessarily correspond to evolutionary histories (degree of relatedness) between organisms. !hus an alternative approach to diagramming relationships between taxa was developed, called cladistics. !he basic assumption behind cladistics is that members of a group share a common evolutionary history, and are thus more closely related to one another than they are to other groups of organisms. Belated groups of organisms are recogni9ed because they share a set of uni?ue features "apomorphies) which were not present in distant ancestors, but which are shared by most or all of the organisms within the group. !hese shared derived characteristics are called synapomorphies. "n contrast to phenetics, in cladistics groupings do not depend on whether organisms share physical traits, but on their evolutionary relationships. "ndeed, in cladistic analyses two organisms may share numerous characteristics but still be considered members of different groups. %or example, a #ellyfish, a sea star, and a human& #ellyfish and sea stars both live in the ocean, have radial symmetry and are invertebrates, so phenetic analysis might place them together in a group. ,owever, this would not reflect evolutionary relationships, as sea stars are actually more closely related to humans than they are to #ellyfish (both are deuterostomes). !hus in cladistics, the emphasis is not upon the presence of all shared traits, but upon the presence of shared derived (apomorphic) traits. "n the example above radial symmetry, a?uatic habitat and invertebrate structure are all traits that are believed to have been present in the common ancestor of all animals, and so such traits are not

considered to be very useful in determining relationships using cladistic analysis. 8ladistic analyses have some pretty strict rules. %or example, cladists always assume that new species arise by bifurcations of the original lineage (the lineage always splits in two). 3ost cladists assume that the original ancestral species no longer exists after this bifurcation, so each branching event results in two new species. "n addition, cladistic groupings must possess the following characteristics& '. All species in a grouping must share a common ancestor. *. All species derived from a common ancestor must be included in the taxon. !he application of these re?uirements results in the following terms being used to describe the different ways in which groupings can be made&

A monophyletic grouping is one in which all species share a common ancestor and all species derived from that common ancestor are included. !his is the only form of grouping accepted as valid by cladists. (%or example, turtles, li9ards, crocodilians and birds are all derived from a shared common ancestor. !hus a monophyletic grouping would place all of these together, rather than placing birds into a separate group.) A paraphyletic grouping is one in which all species share a common ancestor, but not all species derived from that common ancestor are included (for example, grouping turtles, li9ards and crocodiles as reptiles and separating that grouping from the birds). A polyphyletic grouping is one in which species that do not share an immediate common ancestor are lumped together, while excluding other members that would link them (for example, a hypothetical group the li9mams made by grouping together the li9ards and the mammals).

!hus, in cladistics, no matter how divergent in appearance 4 might be from 8, relative to A, if 4 and 8 share a common ancestor that is not shared by A, then 4 and 8 must be grouped together and separated from lineage A. "n the cladogram of the reptiles and birds above, you can see an example of such a situation. Steps for Constructing a Cladogram '. (elect a taxonomic group to be analy9edC for example, a group of vertebrates. *. %or each member of the group, determine some observable traits (characters), and note their states (a character state is one of two Dor moreE possible forms of that character). %or example, for a character fins, the possible states may be present and absent. %or the character number of forelimb digits, possible states may be ', *, -, /, or + -. %or each character, determine which state is ancestral (primitive or plesiomorphic) and which is derived (apomorphic). !his is usually done by comparison with a more distantly related organism termed the outgroup. "t is

hypothesi9ed that traits shared with the more distantly related organism(s) are likely to be ancient or plesiomorphic traits. (imilarly, traits that differ from the outgroup are postulated to have arisen since the group being considered branched from its shared common ancestor with the outgroup, and thus are likely to be derived or apomorphic. /. 0roup taxa by shared derived character states (synapomorphines). +. When in doubt, choose the most parsimonious tree. While similar structures may evolve independently in separate lineages facing similar selective pressures (convergent evolution), this is assumed to be a rare event. 3ost ma#or structures (eyes, horns, tails, fur, etc.) are assumed to have evolved or to be lost only rarely. !hus, when in doubt, choose a pathway that minimi9es the number of times a feature must be postulated to have arisen (or lost) separately. An Example of Cladogram Construction for $erte%rates
Trait &utgroup )rog "'o%e( finned fish# $es $es Turtle *angaroo +ouse ,uman

-orsal .erve $es Cord 'egs .ature of egg @o Be?uires water

$es $es

$es $es Develops inside the mother 3arsupial $es $es $es

$es $es Develops inside the mother <lacental $es @o @o

$es $es Develops inside the mother <lacental Beduced @o $es

Be?uires ,ard shell water prevents drying "n egg @o @o @o "n egg @o @o @o

.ature of "n egg development ,air Presence of pouch /idpedal posture @o @o @o

"n this example, frogs share all ma#or traits with the outgroup (i.e., they show mostly ancestral or plesiomorphic traits), except that they have legs and slightly enlarged brains. !hese last two features are apomorphies that are widespread in the vertebrate lineage. %rogs are thus postulated to have branched from the main vertebrate lineage relatively early in the evolutionary process. !urtles show further modifications from the outgroup, most markedly the presence of a hard shelled egg, as well as an increased tendency toward larger brain si9eC therefore we would suggest that their lineage branched next from the ancestral

lineage. All three of the remaining groups are characteri9ed by an egg that develops inside the mother, suggesting that these three share a common ancestor not shared by frogs and turtles. 3ice and kangaroos share similar hair amounts, while humans and kangaroos share a generally bipedal posture. (o how do we know how to group these three organisms5 %irstly, we would suspect that the possession of hair, even in reduced amounts, might link humans to kangaroos and mice. (econdly, we would look to the other traits possessed by these groups. 4oth mice and humans show placental development and thus lack a pouch. !hus we would tend to link these two groups together more closely and the kangaroo more distantly. We would thus conclude that the cladogram for this group of organisms (minus the outgroup, which is not usually shown in these figures) should look something like the one below.

$ou will note that, in this solution, the tendency toward a bipedal posture was postulated to have evolved twiceC once in the marsupial lineage (kangaroos) and once in the placental lineage (humans). (uch features are said to be analogous and to have resulted from convergent evolution. ,owever, it is possible to draw the cladogram such that humans and kangaroos are postulated to have a common bipedal ancestor not shared by mice. ,owever, in this solution, the tendency towards placental development, along with all the re?uired anatomical changes (absence of a pouch etc.) must be postulated to have occurred twice (once in the mouse lineage and once in the human lineage). (uch a solution would re?uire more evolutionary steps than the se?uence that we have proposed here and, conse?uently, would not be as efficient or parsimonious as our initial solution. Exercises

8lick here for =xercise '. 8lick here for =xercise *. 8lick here for =xercise -. 8lick here for =xercise /.

Evolutionary "Synthetic# Systematics 1ike cladistics, evolutionary systematics is based strongly on evolutionary

relationships. ,owever, its supporters suggest that the degree of genetic differences between lineages should be used in addition to their genealogical (evolutionary) similarities when developing taxonomic classifications. Among other ways in which evolutionary systematics and a cladistic analyses differ& evolutionary taxonomists are more tolerant of multiple branches developing synchronously from an ancestral line (rather then #ust two as in cladistics), and they tend to treat organisms that are unchanged when a new branch arises from their lineage as the same species that existed before the branching event (i.e., the original species is not always believed to become extinct every time a branching event occurs). ,owever, the biggest difference between the two approaches is that, when creating groupings, evolutionary taxonomists seek to maximi9e the way in which the groupings they create communicate information about the group of organisms in that taxon (in the way the word mammal or reptile does for even an untrained person). !his type of division is also intended to increase the ease with which a field biologist, or even someone coming in from outside, can retrieve information about a group of organisms of interest to himFher using the taxonomic divisions formed. !axa generated by an evolutionary biologist will never be polyphyletic, but may be either monophyletic or paraphyletic in nature. %or example, as we saw earlier, birds and crocodilians diverged from the same ancestral reptilian line. A cladist would insist that these sister groups be placed in the same taxon, even though the amount of change from the common ancestor is much greater for the birds than it is for the crocodiles. An evolutionary taxonomist would suggest that the large number of similarities between crocodilians and reptiles would #ustify grouping them within the same general taxon, while placing birds in a separate taxon due to the large number of uni?ue characters possessed by members of this group.

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Introduction
"n this tutorial you will be learning about the 'innaean system of classification used in the biological sciences to describe and categori9e all living things. !he focus is on finding out how humans fit within this system. "n addition, you will discover part of the great diversity of life forms and come to understand why some animals are considered to be close to us in their evolutionary history.

,ow many species are there!

!his is not an easy ?uestion to answer. About ' -F/ million have been given scientific names. @early *F- of these are insects. =stimates of the total number of living species generally range from '; to ';; million. "t is likely the actual number is on the order of 'to '/ million, with most being insects and microscopic life forms in tropical regions. ,owever, we may never know how many there are because many of them will become extinct before being counted and described. !he tremendous diversity in life today is not new to our planet. !he noted paleontologist (tephen .ay 0ould estimated that GGH of all plant and animal species that have existed have already become extinct with most leaving no fossils. "t is also humbling to reali9e that humans and other large animals are freakishly rare life forms, since GGH of all known animal species are smaller than bumble bees.

hy should we %e interested in learning a%out the diversity of life!

"n order to fully understand our own biological evolution, we need to be aware that humans are animals and that we have close relatives in the animal kingdom. 0rasping the comparative evolutionary distances between different species is important to this

understanding. "n addition, it is fun to learn about other kinds of creatures.

hen did scientists %egin classifying living things!

4efore the advent of modern, genetically based evolutionary studies, =uropean and American biology consisted primarily of taxonomy , or classification of organisms into different categories based on their physical characteristics. !he leading naturalists of the 'Ith and 'Gth centuries spent their lives identifying and naming newly discovered plants and animals. ,owever, few of them asked what accounted for the patterns of similarities and differences between the organisms. !his basically nonspeculative approach is not surprising since most naturalists two centuries ago held the view that plants and animals (including humans) had been created in their present form and that they have remained unchanged. As a result, it made no sense to ask how organisms have evolved through time. (imilarly, it was inconceivable that two animals or plants may have had a common

8arolus 1innaeus 'K;K)'KKI

ancestor or that extinct species may have been ancestors of modern ones. :ne of the most important 'Ith century naturalists was a (wedish botanist and medical doctor named >arl von 1innJ. ,e wrote 'I; books mainly describing plant species in extreme detail. (ince his published writings were mostly in 1atin, he is known to the scientific world today as Carolus 'innaeus , which is the 1atini9ed form he chose for his name. "n 'K-+, 1innaeus published an influential book entitled Systema Naturae in which he outlined his scheme for classifying all known and yet to be discovered organisms according to the greater or

lesser extent of their similarities. !his 1innaean system of classification was widely accepted by the early 'Gth century and is still the basic framework for all taxonomy in the biological sciences today. !he 1innaean system uses two 1atin name categories, genus and species , to designate each type of organism. A genus is a higher level category that includes one or more species under it. (uch a dual level designation is referred to as a binomial nomenclature or %inomen (literally two names in 1atin). %or example, 1innaeus described humans in his system with the binomen Homo sapiens , or man who is wise ))Homo is our genus and sapiens is our species.
genus species species genus species species

1innaeus also created higher, more inclusive classification categories. %or instance, he placed all monkeys and apes along with humans into the order Primates . ,is use of the word <rimates (from the 1atin primus meaning first ) reflects the human centered world view of Western science during the 'Ith century. "t implied that humans were created first. ,owever, "t also indicated that people are animals.
order family genus species species genus species species genus species species family genus species species

While the form of the 1innaean classification system remains substantially the same, the reasoning behind it has undergone considerable change. %or 1innaeus and his contemporaries, taxonomy served to demonstrate the unchanging order inherent in 4iblical creation and was an end in itself. %rom this perspective, spending a life dedicated to precisely describing and naming organisms was a religious
8harles Darwin 'I;G)'II*

act because it was revealing the great complexity of life created by 0od. !his static view of nature was overturned in science by the middle of the 'Gth century by a small number of radical naturalists, most notably Charles -arwin. ,e provided conclusive evidence that evolution of life forms has occurred. "n addition, he proposed natural selection as the mechanism responsible for these changes. 1ate in his life, 1innaeus also began to have some doubts about species being unchanging. 8rossbreeding resulting in new varieties of plants suggested to him that life forms could change somewhat. ,owever, he stopped short of accepting the evolution of one species into another.

hy do we classify living things today!

(ince Darwin6s time, biological classification has come to be understood as reflecting evolutionary distances and relationships between organisms. !he creatures of our time have had common ancestors in the past. "n a very real sense, they are members of the same family tree. !he great diversity of life is largely a result of branching evolution or adaptive radiation. !his is the diversification of a species into different lines as they adapt to new ecological niches and ultimately evolves into distinct species. @atural selection is the principal mechanism driving adaptive radiation.

Beturn to 3enu

<ractice Lui9

@ext !opic

!his page was last updated on . 8opyright M 'GGI)*;;+ by Dennis :6@eil. All rights reserved. "llustration credits