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Introduction to
Invertebrates
RECONSTRUCTION OF
INVERTEBRATE PHYLOGENY
Cladistic Method
Downplaying the Linnean Categories
AN EXERCISE IN CLADISTICS
1
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T
Chapter 1
Introduction to Invertebrates
he study of invertebrates is a gateway to the vast diversity
of animal life. The astronomical numbers and myriad forms of
invertebrates delight the eye, challenge the mind, and provide
limitless opportunities for scientific discovery. Invertebrates
account for more than 99% of all species of animals and,
although fewer than 1 million living species have been named
so far, the total number of animal species on Earth may exceed
30 million. Clearly, most invertebrate species remain to be discovered and described, while among those already identified,
few have been studied in depth. Thus, any curious individual
equipped with the simplest observational or experimental tools
can make original and enduring contributions to science.
At first glance, the diversity of invertebrates may seem
incomprehensible, like contemplating infinity, but the countless individuals and species are variations on a relatively
few readily identifiable themes. One of these themes is the
ground plan (basic design, or Bauplan) of each taxonomic
group. For example, although there are millions of species of
arthropods, all have a segmented body, an exoskeleton that is
molted with growth, and jointed appendages. Knowing those
few traits enables anyone to identify an arthropod as such and
to appreciate the insect, crustacean, centipede, and spider
variations on the arthropod theme. The chapters in this book
have been written to highlight the ground plan of each major
group and thus to provide a foundation on which to understand the thematic variations.
Another way to simplify the diversity of invertebrates is to
establish the evolutionary relationships between the groups
of animals, to draw an evolutionary tree. This is accomplished
by identifying similarities in the themes of two or more groups
and uniting them on the basis of their unique similarities. So,
for example, crabs, lobsters, and shrimps (and a few minor
groups) are closely related to each other because only they,
among all crustaceans, share a design with five pairs of locomotory appendages. The evolutionary history of a group is
called its phylogeny and the actual depiction of evolutionary
relationships is known as a phylogenetic tree (or cladogram).
Although a phylogenetic tree, as a scientific hypothesis, is
subject to testing and change, it nevertheless provides a framework on which to organize and compare ground plans and
thus summarize much of the factual content of each chapter.
For those unfamiliar with the method for constructing or interpreting phylogenetic trees, an exercise in phylogenetic reconstruction is provided later in this chapter.
Yet another approach to invertebrate diversity is to view animal design in a framework of functional principles. Here we
seek to understand how the fundamental principles of physics
and chemistry impose limits — indeed, control — design. For
example, flight requires an airfoil (wing) to produce lift.
Although the composition of wings can differ (exoskeleton,
skin, feathers, aluminum) and the animals having them may be
unrelated, all necessarily have the shape of a wing. Thus, we
can predict that any animal that evolves flight will have something that looks like a wing. At a slightly more derived level are
the principles of physiology, development, and ecology. These
concepts together — physical, chemical, biological — allow one
to view living invertebrates as complex and fascinating expressions of a few principles. The subtlety and elegance of this
approach breathes life into invertebrate zoology.
This book integrates invertebrate structure and function in
an evolutionary context. The sequence of chapters, especially
in the first part of the book, is progressive, and several chapters (those whose titles include “Introduction to”)
describe major turning points in animal evolution, such as the
origin and significance of the multicellular body and, later,
bilateral symmetry. At each of these “steps,” the animal body
evolved a radically new design, which created novel ecological
opportunities and was inherited by all descendants. How those
new designs might have evolved, how they work in the context
of general functional principles, and how they enabled their
possessors to exploit new adaptive zones are the subjects of the
“Introduction to” chapters. This book, then, is a blend of factual descriptions and provisional syntheses that are subject to
scientific testing and revision. Throughout the book, we
emphasize that opportunities abound for discovery in this
dynamic field.
To provide comprehensive coverage of invertebrates is
a goal of this book, but a few taxa are described in less detail
than others. Because most schools have specialized courses in
protozoology, parasitology, and entomology, we provide only
abbreviated coverage of unicellular organisms (protozoa),
parasitic invertebrates, and insects (Hexapoda).
RECONSTRUCTION OF
INVERTEBRATE PHYLOGENY
CLADISTIC METHOD
One of the noteworthy achievements of biological research
has been the establishment of a natural system, the recognition of species and the arrangement of those species in a hierarchic pattern of relationships. This pattern of evolutionary or
kinship relationships is called a phylogeny, or a “tree of life.”
The science that concerns itself with the discovery and depiction of phylogeny is known as phylogenetic systematics or
cladistics. The goal of cladistics is to discover and portray the
kinship relationships among species, ultimately in a phylogenetic tree, or cladogram.
In practice, one establishes a kinship relationship between
two named groups, or taxa (the singular is taxon), by detecting
a trait, or character, that is expressed in these taxa alone. Such
a uniquely shared character is called a synapomorphy
( shared derived character), and taxa united by one or more
synapomorphies are called sister taxa. Because a synapomorphy is shared by the sister taxa and no other taxon, it must
have evolved in the immediate ancestor of the sister taxa. Thus
the synapomorphy observed in the descendants actually originated in the ancestor as an evolutionary novelty, now called an
autapomorphy ( self-derived character). As the sole descendants of that one immediate ancestor, the sister taxa constitute
a monophyletic ( one origin) taxon. The ultimate goal of
cladistics is to reconstruct a comprehensive tree of life based
solely on monophyletic taxa.
An example of these concepts can be drawn from the arthropods (Fig. 1-1). For taxa, consider the crustaceans (such as
shrimps, crabs, and lobsters) and tracheates (insects, milli-
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Reconstruction of Invertebrate Phylogeny
Chelicerata
Tracheata
Crustacea
“Jointed appendages”
“Mandibles”
(Synapomorphy:
Monophyly)
(Symplesiomorphy:
Paraphyly)
Mandibulata
“Jointed appendages”
(Synapomorphy:
Monophyly)
“Chelicerae”
(Autapomorphy)
Arthropoda
“Mandibles”
(Autapomorphy)
“Jointed appendages”
(Autapomorphy)
FIGURE 1-1 Cladistics. Example of the use of cladistics in
phylogenetic reconstruction (see text). The observation of a uniquely
shared character (synapomorphy), “mandibles,” in Crustacea and
Tracheata unites them as sister taxa in a monophyletic taxon,
Mandibulata. Thus the stem species, or ancestor, of Mandibulata must
have first expressed “mandibles” as an evolutionary novelty
(autapomorphy). Similarly, the observation that “jointed appendages”
occur in only chelicerates and mandibulates and nowhere else indicates
that the character “jointed appendages” is a synapomorphy of the
two sister taxa, which form the monophyletic taxon Arthropoda. The
ancestral arthropod, therefore, first acquired “jointed appendages” as
an evolutionary innovation, or autapomorphy. If an observer, for lack
of complete knowledge, unites chelicerates and crustaceans into
what he believes is a monophyletic taxon because both share “jointed
appendages,” the union is false because “jointed appendages” are
not unique to chelicerates and crustaceans alone, but to all arthropods.
Thus the character “jointed appendages” did not evolve in the
immediate ancestor to chelicerates and crustaceans alone, but in
a deeper ancestry that also gave rise to the tracheates. The chelicerates
and crustaceans express this deeply ancestral trait as a shared
primitive character, or symplesiomorphy. The false union based on
a symplesiomorphy is called a paraphyletic taxon. The autapomorphy
“chelicerae” is a pair of pincerlike appendages on the second segment
of the head.
pedes, and centipedes, for example). Representatives of these
two taxa show that all share the character “mandibles” ( jaws
derived from appendages on the third head segment), which
occurs nowhere else and thus is a synapomorphy of crustaceans
and tracheates. The synapomorphy indicates that the sister
taxa (Crustacea, Tracheata) constitute a monophyletic taxon,
the ancestor of which first evolved mandibles as an evolutionary novelty, or autapomorphy. Once the monophyletic taxon
has been identified, it is given a formal name — in this case,
Mandibulata.
Now suppose that, instead of crustaceans and tracheates,
the study began with crustaceans and chelicerates (including
horseshoe crabs, scorpions, and spiders) and found that they
shared the character “jointed appendages,” which was then
hypothesized to be a synapomorphy of the two taxa (Fig. 1-1).
The character “jointed appendages,” however, is not shared by
3
crustaceans and chelicerates alone, but also by the tracheates.
This means that the “jointed appendages” of chelicerates and
crustaceans is not a shared derived character but rather
a shared ancestral trait, or symplesiomorphy, that evolved in the
stem species (ancestor) of all arthropods. The erroneous union
of crustaceans and chelicerates based on a symplesiomorphy is
called a paraphyletic taxon (Fig. 1-1). A paraphyletic taxon contains some, but not all, of the descendants of the stem species.
A major challenge of modern phylogenetic research is to identify and weed out paraphyletic taxa. In this example, “jointed
appendages” is actually an autapomorphy of the monophyletic
taxon Arthropoda, which includes the sister taxa Chelicerata
and Mandibulata (Crustacea Tracheata; Fig. 1-1).
A paraphyletic taxon fails to include all descendants of one
ancestor, but a polyphyletic taxon includes the descendants of
more than one ancestor. This mistake occurs when a shared
similarity results from evolutionary convergence rather than
common ancestry. Similarity attributable to common genetic
inheritance is called homology, whereas the superficial similarity that arises from convergence is known as homoplasy (or
analogy). Only homologous structures are useful in phylogenetic reconstruction based on monophyletic groups. An example of a polyphyletic taxon would be one that united birds,
bats, and insects together because all share “wings.” Bird
wings, bat wings, and insect wings, however, are unrelated
homoplasous structures, and the members of this polyphyletic
taxon descended from three separate ancestors, each of which
independently evolved its own unique wing.
The application of the cladistic method to species, groups
of species, and groups of groups of species leads to the
dichotomously branching hierarchic structure that typifies
a phylogenetic tree. Hierarchic structure means that species
are nested in larger, more inclusive taxa, which are contained
in still more inclusive groups. So a species of shrimp is a kind
of crustacean, which is a kind of mandibulate, which is a kind
of arthropod, which is a kind of animal. The hierarchy shown
in Figure 1-1 can also be summarized in tabular form in a phylogenetic hierarchy:
Arthropoda
Chelicerata
Mandibulata
Crustacea
Tracheata
A properly constructed phylogenetic tree is a deeply informative representation. The kinship relationships of species and
higher taxa are shown graphically and the synapomorphies are
indicated for each pair of sister taxa. The autapomorphies
associated with stem species, moreover, offer insight into the
form of the ancestors, which then may be compared with fossil
evidence, if such exists.
Although we try to choose simple, factually sound examples
to explain the method of phylogenetic reconstruction, in practice most reconstructions are more complex. It often happens,
for example, that apparent synapomorphies are contradictory:
Synapomorphy 1 unites taxa A and B and synapomorphy
2 unites taxa B and C. This results in two different and
competing trees. In this situation, systematists choose the tree
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Chapter 1
Introduction to Invertebrates
with the fewest branches, the tree that assumes the fewest
number of character changes — that is, the most parsimonious
tree. This principle of parsimony is one of the important
underlying assumptions of phylogenetic systematics (and of
science in general). It is an especially important tool in computer-aided cladistics. In such analyses, the software sifts
through large amounts of data in the form of numerically
coded morphological and molecular characters and, using different routines for joining or weighting them, generates more
than one tree. From among these alternatives, a most parsimonious tree is typically calculated and drawn.
In this book, all trees of phylogenetic relationship are
based on synapomorphies and, unless otherwise noted, are
plotted by hand, as in Figure 1-1. This explicit and direct
approach was adopted to allow you, the student, to evaluate
the choice and validity of characters used in the reconstructions. Trees are sensitive to new facts and to reevaluation of
assumptions about the nature of the characters themselves.
As you acquire new knowledge applicable to phylogenetics,
create your own trees and test them against those presented in
the text.
DOWNPLAYING THE LINNEAN
CATEGORIES
Modern cladistics is the method of choice for establishing
kinship relationships among taxa. Once monophyletic taxa
are identified and named on a phylogenetic tree or in a phylogenetic hierarchy, the hierarchic relationships among the taxa
are obvious and clear (see Fig. 1-1 and the phylogenetic
hierarchy set out in the last section). The question then is:
“What is gained by assigning a Linnean category — phylum,
class, order, and so on — to the taxon names already present?”
Many systematists today have abandoned the Linnean categories because they are unnecessary, because they were
established by Linnaeus for what he considered to be the
immutable levels of Creation, and because new cladistic ranks
of many taxa do not coincide well with the old Linnean categories. For some, liberation from the Linnean categories will
shift emphasis away from pigeonholing (“To what class does
this animal belong?”) to science (“What are the postulated
synapomorphies of Crustacea and Tracheata and how can they
be tested?”). For others, however, the Linnean hierarchy of
categories will continue to provide a familiar and constant
frame of reference.
This book embraces the new cladistic approach to systematics and largely dispenses with Linnean categories. The relative
ranks of taxa can be obtained from the phylogenetic trees,
the text sections that include “Phylogenetic Hierarchy of ” in
their title, and the chapter headings. To soften the transition
from the classical to modern approach, however, Linnean
categories are assigned to taxa as superscript abbreviations
when a taxon name is used as a text heading. The standard
abbreviations are: P for phylum, C for Class, O for Order, and
F
for Family. An uppercase S prefixed to a category abbreviation
identifies a supertaxon, a lowercase s identifies a subtaxon, and
a lowercase i indicates an infrataxon. Super-, sub-, and infrataxa
are ranked as follows:
Supertaxon
Taxon (for example, phylum, class, order)
Subtaxon
Infrataxon
In the phylogenetic hierarchy given earlier, the superscripts would be: ArthropodaP, CheliceratasP, MandibulatasP,
CrustaceaiP, and TracheataiP.
AN EXERCISE IN CLADISTICS
Figures 1-2, 1-3, and 1-4 as well as the following text provide an
exercise in cladistics for those who wish to practice building
a phylogenetic tree. The exercise is divided into three steps,
each of which is highlighted and explained here and in the
figure legends.
• Step 1: Study characters of a group of taxa in question and
search for apomorphies, first within species (Fig. 1-2)
Figure 1-2 shows one individual from each of six fictitious
species. The first task is to establish (or not) the uniqueness
(validity) of each species. To accomplish this, search for a character that is expressed uniquely in each species alone. Once
identified, that character is assumed to have evolved as an evolutionary novelty in the ancestor of that species and is thus
a species-level autapomorphy. An autapomorphy for each of
the six species is shown in Figure 1-2. In this example, these
species are then named appropriately according to their
autapomorphies. So, the species with the autapomorphy
“fan tail” is named Fan tail, the species with “long antennae” is
named Long antennae, and so on as shown in Figure 1-2. The
italicized species binomial always consists of a capitalized genus
name (for example, Fan) and a lowercase species name (tail).
In actual practice, both names would be latinized.
• Step 2: Find synapomorphies that link species and groups of
species pairwise into a most parsimonious tree (Fig. 1-3)
Figure 1-3 shows the most parsimonious phylogenetic tree
for the six species. The tree results from the recognition of
a character or characters (synapomorphies) that are shared
uniquely by two species or two groups of species. Notice that
at least one synapomorphy supports the pairwise union of
species (and higher groups) into sister taxa. For example,
species Shoulder leg and Fan tail are united in a monophyletic
taxon by the uniquely shared character (synapomorphy)
“stalked eyes.” This new monophyletic taxon has yet to be
named and thus is temporarily designated Nomen nominandum ( new name), abbreviated N. N.
Proceeding deeper into the tree, notice that the monophyletic taxon that includes Shoulder leg and Fan tail is united
with the species Scissors tail in another monophyletic taxon by
the synapomorphies “three segments” and “pliers claw.” This
new, higher-level taxon is also temporarily designated N. N.
Figure 1-3 shows the progressive pairing of taxa into monophyletic groups based on shared derived characters. But
a dilemma is encountered with Paddle foot at the base of the
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An Exercise in Cladistics
5
Head
Segments
Fan tail
Shoulder leg
Scissors tail
Five segments
Tail
Paddle foot
Long antennae
FIGURE 1-2 Exercise in cladistics, step 1 (see also text). Each of the six illustrated animals represents one
fictitious species. The first step in the analysis is to identify one or more characters that is unique to each species
(autapomorphy). The autapomorphies of each species in this example are shaded and reflected in the species’
name.
tree. The dilemma is whether the characters associated with
Paddle foot are plesiomorphies or apomorphies. Is the character “paddle foot” an autapomorphy of Paddle foot that is found
in that species and nowhere else, or is it a symplesiomorphy
occurring in Paddle foot and in other, as yet unstudied species?
Similarly, is the character “eight segments” an autapomorphy
of the original six species or is it a symplesiomorphy occurring
in those six plus additional species? The questions boil down
to this: How do we establish the apomorphy vs. plesiomorphy
polarity of the characters? The monophyly or paraphyly of the
group of six species hinges on the determination of character
polarity. The practical solution to this dilemma is to discover
whether or not “eight segments” and “paddle foot” occur
outside the group of six species, and this requires the examination of additional species. The extension of the cladistic
analysis beyond the taxa of immediate interest is called an
outgroup comparison. The goal is to determine character
polarity to resolve the plesiomorphy vs. apomorphy dilemma.
have eight segments. Some have evolved a body with fewer
segments, in this example, five and three segments. Or, alternatively, “eight segments” is an autapomorphy of a monophyletic
taxon that includes the original six species. The character “eight
segments,” as both synapomorphy and autapomorphy, is shown
in Figure 1-4.
In reference to the original analysis, the ingroup of six
species, apomorphies now support the monophyly of all
sister taxa. At this point, it is appropriate to replace the provisional N. N. designation with a formal Latin name. These
formal names, which are derived from taxon autapomorphies, are shown on Figure 1-4. For example, the monophyletic taxon supported by the apomorphy “stalked eyes” is
named Exophthalmia ( prominent eyes), the taxon supported by “three segments” is named Triannelida ( three
rings), and the monophyletic taxon that includes all six
species, based on the apomorphy “eight segments,” is designated Octoannelida ( eight rings). The phylogenetic hierarchy of Octoannelida is:
• Step 3: Perform an outgroup analysis and name the monophyletic taxa (Fig. 1- 4)
Figure 1-4 resolves the uncertain polarity of characters “paddle foot” and “eight segments” by performing an outgroup
analysis. The outgroup in this example is represented by a single
unnamed species and it is compared with the ingroup consisting
of the six original species. The outgroup species lacks the characters “paddle foot” (it has “normal” feet) and “eight segments”
(it has nine segments). Thus, the outgroup comparison enables
us to conclude that “paddle foot” is an autapomorphy of the
species Paddle foot and “eight segments” is a synapomorphy of
Paddle foot and the taxon (Cultrichela) that includes the remaining five ingroup species. Not all species of Cultrichela, however,
Octoannelida
Paddle foot
Cultrichela
Long antenna
Urocopa
Five segments
Triannelida
Scissors tail
Exophthalmia
Shoulder leg
Fan tail
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Chapter 1
Paddle foot
Introduction to Invertebrates
Long antennae
Five segments
Scissors tail
Fan tail
Shoulder leg
“stalked-eyes”
“three-segments”,“pliers-claw”
N.N.
“oar-tail”
N.N.
“jacknife-claw”
N.N.
“eight-segments”?, “paddle-foot”?
N.N.
Synapomorphy
?
N.N.?
Autapomorphy
Monophyletic taxon
FIGURE 1-3 Exercise in cladistics, step 2 (see also text). The six fictitious species are united pairwise into
monophyletic taxa on the basis of uniquely shared characters (synapomorphies). The observed synapomorphy
of each pair of sister taxa (together a monophyletic taxon) evolved as an evolutionary novelty in the ancestor of
the monophyletic taxon and is indicated on the tree as an autapomorphy. Each of the monophyletic taxa is given
a general temporary designation, abbreviated N. N. Notice the uncertainty associated with the characters “eight
segments” and “paddle foot.” Both characters could be either apomorphies or plesiomorphies. If they are
apomorphies, then the N. N. taxon that includes all six species is monophyletic; if they are plesiomorphies, then
the taxon is paraphyletic. The plesiomorphic vs. apomorphic interpretation of these characters can be resolved
only by comparison with a taxon or taxa outside this group of six species. Such an outgroup comparison is shown
in Figure 1-4 and is described in the text.
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An Exercise in Cladistics
“Outgroup”
Paddle foot
Long antennae
“Ingroup”
Five segments
Scissors tail
Fan tail
10
11
Scissors tail
Exophthalmia
8
9
Five segments
Triannelida
6
7
Long antennae
Urocopa
4
5
Paddle foot
Shoulder leg
Cultrichela
“Eight-segments”
3
2
Octoannelida
1
Synapomorphy
Autapomorphy
Monophyletic taxon
Sister taxa
FIGURE 1-4 Exercise in cladistics, step 3 (see also text). An outgroup analysis shows that the character
“paddle foot” is an autapomorphy of the species Paddle foot. It also indicates that the character “eight
segments” is a synapomorphy of the taxon (species) Paddle foot and its sister taxon, Cultrichela, which
includes all other species. Thus the synapomorphy “eight segments” establishes the monophyletic
taxon Octoannelida for the six species in this study. The autapomorphies of the monophyletic taxa are:
1, Octoannelida: “eight segments”; 2, Paddle foot: “paddle foot”; 3, Cultrichela: “jacknife claw”; 4, Long
antenna: “long antenna”; 5, Urocopa: “oar tail”; 6, Five segments: “five segments” (reduced from eight),
“pliers claw” (modification of “jacknife claw”); 7, Triannelida: “three segments” (reduced from eight);
8, Scissors tail: “scissors tail”; 9, Exophthalmia: “stalked eyes”; 10, Shoulder leg: “shoulder leg”;
11, Fan tail: “fan tail.”
7
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Chapter 1
Introduction to Invertebrates
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Ecology
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9
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INTERNET SITES
General
http://life.bio.sunysb.edu/marinebio/mbweb.html (State University of
New York. Invertebrate images, references, career opportunities, and news regarding marine biology.)
www-marine.stanford.edu/HMSweb/careers.html (Marine biology links,
careers, and more.)
www.ucmp.berkeley.edu (University of California at Berkeley Museum of Paleontology. Explore their invertebrate collections and follow links to impressive images of living invertebrates.)
www.ucmp.berkeley.edu/exhibit/phylogeny.html (Take the Web Lift
to Taxa to discover the relationships that connect all organisms.)
www.nmnh.si.edu/departments/invert.html (Smithsonian National
Museum of Natural History. Good coverage of certain
groups of invertebrates, such as squids and their relatives.)
http://mbayaq.org (Monterey Bay Aquarium. The next best thing
to being there. Explore their beach, rocky intertidal pool,
open water, and kelp forest exhibits, several of which are
three-dimensional and several that are interactive.)
www.bioimages.org.uk/index.htm (BioImages. A U.K. virtual field
guide.)
www.biosis.org/zrdocs/zoolinfo/gp_index.htm (BIOSIS Internet Resource Guide for Zoology.)
Lab Manual
www.lander.edu/rsfox/310labindex.html (Lander University OnLine
Invertebrate Lab. Illustrated anatomical descriptions of
approximately 100 species in support of invertebrate teaching and research.)
Professional Societies
www.invertebrates.org (American Microscopical Society. Publishes
the international journal Invertebrate Biology. The journal’s
Web site links to other sites on invertebrates and opportunities in the field for research, jobs, and education.)
www.museum.unl.edu/asp (American Society of Parasitologists.
Much useful information and many images at their Web
site.)
Cladistics
tolweb.org/tree/phylogeny.html (Tree of Life. Provides a phylogenetic classification of all taxa of living organisms based on
molecular and morphological traits. Coverage is uneven
and often outdated, but new information is being added
constantly. Good source of contemporary references to
invertebrate systematics.)
www.nhm.ukans.edu/downloads/CompleatCladist.pdf (An online version of Wiley, E. O., Siegel-Causey, D., Brooks, D. R., and
Funk, V. A. 1991. The Compleat Cladist: A Primer of Phylogenetic
Procedures. Special Publication No. 19. University of Kansas
Museum of Natural History, Lawrence, KS.)
www.gwu.edu/~clade/faculty/lipscomb/Cladistics.pdf (Diana Lipscomb’s 1998 workbook, Basics of Cladistic Analysis
[George Washington University, Washington, DC.])
erms.biol.soton.ac.uk (European Register of Marine Species.)
www.nhm.ac.uk/hosted_sites/uksf (U.K. Systematics Forum. A database of taxonomists and the Web of Life.)
animaldiversity.ummz.umich.edu/index.html (University of Michigan Museum of Zoology Animal Diversity Web.)