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Chapter 10 - Classification And Phylogeny Of Animals
10-1
Chapter 10 - Classification And Phylogeny Of Animals
4. A species has a geographic range and evolutionary duration, which defines its distribution through
space and time. Geographic ranges may be large or worldwide distributions, cosmopolitan, or
restricted ranges, endemic. A species geographic range can change over its evolutionary duration.
B. Typological Species Concept (Figure 10.3)
1. Before Darwin, species were considered fixed and with essential and immutable features.
2. Typological species concept centered on an ideal form for the species; variations were “imperfect.”
3. A type specimen was labeled and deposited in a museum to represent this “standard” specimen;
today the practice continues, but as a name-bearer to compare with potentially new species.
4. Variation in a population is now recognized as normal.
C. Biological Species Concept
1. Theodosius Dobzhansky and Ernst Mayr formulated this during the evolutionary synthesis.
2. A species is a reproductive community of populations (reproductively isolated from others) that
occupies a specific niche in nature.
3. The ability to successfully interbreed is central to the concept.
4. The criteria of “niche” tie in ecological properties.
5. However, controlled breeding experiments can be difficult to conduct.
6. Molecular and other studies may detect sibling species with similar morphology.
7. The biological species concept lacks an explicit temporal dimension and gives little guidance
regarding the species status of ancestral populations relative to their evolutionary descendants.
8. Proponents of the biological species concept often disagree on the degree of reproductive isolation
necessary for the determination of distinct species.
9. The biological species concept has received strong criticism for a number of reasons:
a. A species has limits in space and time; boundaries between species may be difficult to locate.
b. Species are both a unit of evolution and a rank in taxonomic hierarchy.
c. Interbreeding is not an operational definition in asexual organisms.
D. Evolutionary and Cohesion Species Concepts
1. Where do we set the species boundary in a lineage leading back to a fossil form?
2. Simpson provided the evolutionary species concept in the 1940s; it persists with modification.
(Figure 10.4)
3. An evolutionary species is a single lineage of ancestor-descendant populations that maintains its
identity from other such lineages and that has its own evolutionary tendencies and historical fate.
4. This definition accommodates both sexual and asexual forms as well as fossils.
5. The cohesion species concept is as follows: the most inclusive population of individual shaving the
potential for phenotypic cohesion through intrinsic cohesion mechanisms.
E. Phylogenetic Species Concept
1. The phylogenetic species is an irreducible (basal) grouping of organisms diagnosably distinct from
other such groupings and within which there is a parental pattern of ancestry and descent.
2. This is a monophyletic unit that recognizes the smallest groupings that undergo evolutionary change.
3. It discerns the greatest number of species but may be impractical.
4. This system disregards details of evolutionary process.
F. Dynamism of Species Concepts
1. The various competing concepts of species have a common underlying principle, that a species
constitutes a segment of a population-level lineage—the general lineage concept of species.
2. Disagreement is a sign of dynamic research and argument.
3. We cannot predict which species concepts will remain useful in the future.
4. It is possible that disagreements regarding species boundaries may identify interesting cases of
evolution in action.
5. No one concept is comprehensive or final; all need to be understood to understand future concepts.
G. DNA Barcoding of Species
1. DNA barcoding is a technique for identifying organisms to species using sequence information from
a standard gene present in all animals.
2. This method can be used for specimens in nature as well as natural-hisotry museums, zoos, aquaria,
and frozen-tissue collections.
3. DNA barcoding often permits the origin of a specimen to be identified to a particular local
population.
10.3. Taxonomic Characters and Phylogenetic Inference
10-2
Chapter 10 - Classification And Phylogeny Of Animals
A. Homology
1. A phylogeny or evolutionary tree is based on the study of characters that vary among species.
2. Character similarity that results from common ancestry is called homology.
3. Different lineages may develop similar features independently; this is convergent evolution.
4. Characters that are similar but misrepresent common descent are nonhomologous or homoplastic.
B. Using Character Variation to Reconstruct Phylogeny (Figure 10.5)
1. Reconstructing phylogeny requires determining ancestors and descendants.
2. The form that was present in the common ancestor is ancestral.
3. Characters that arose later are derived character states.
4. An outgroup shows if a character occurred both within and outside the common ancestor.
5. A series of species that share derived characters form a subset called a clade.
6. The derived character shared by members of a clade is called a synapomorphy of that clade.
7. By identifying the nested hierarchy of clades or branches, we can form patterns of common descent.
8. Character states ancestral for a taxon are plesiomorphic for that taxon; sharing of ancestral states is
termed symplesiomorphy.
9. Identifying the level at which a character state is a synapomorphy may identify a clade.
10. A cladogram is a nested hierarchy of clades.
11. To obtain a phylogenetic tree, we must add important information on ancestors, duration of lineages,
and amount of change.
C. Sources of Phylogenetic Information (Figure 10.6)
1. Comparative morphology examines shapes, sizes and development of organisms.
a. Skull bones, limb bones, scales, hair and feathers are important morphological characteristics.
b. Both living specimens and fossils are used as phylogenetic information.
2. Comparative biochemistry analyzes amino acid sequences in proteins and nucleic acid sequences in
nucleotides.
a. Direct sequencing of DNA and indirect comparisons of proteins sequences are comparative
methods.
b. Some recent studies use biochemical techniques to analyze fossils.
3. Comparative cytology (karyology) examines variation in number, shape and size of chromosomes in
living organisms.
4. Fossils can provide information on the relative time of evolution; radioactive dating can confirm age.
5. Some protein and DNA sequences undergo linear rates of divergence; this allows us to calculate the
time of the most recent common ancestors.
10-3
Chapter 10 - Classification And Phylogeny Of Animals
6.Phenetic taxonomy was another school of classification that abandoned producing phylogenetic
trees in favor of measuring overall similarity; but it has little support today.
C. Phylogenetic Systematics/Cladistics (Figure 10.10)
1. Willi Hennig first proposed cladistics or phylogenetic systematics in 1950.
2. It emphasizes common descent and cladograms.
3. For example, chimpanzees, gorillas and orangutans are included in Hominidae with humans.
4. Cladists do not assert that amphibians evolved from fish or birds from reptiles; they contend that a
monophyletic group descending from a paraphyletic group contains no useful information.
5. Likewise, extinct ancestral groups are always paraphyletic since they exclude a descendant that
shares their most recent common ancestor.
6. Some cladists indicate that primitive and advanced are relic ideas derived from a pre-Darwinian
“scale of nature” leading to humans near the top; groups were defined based on “higher” features
they lacked.
7. Cladists avoid paraphyletic groups by defining a list of sister groups to each more inclusive taxon.
D. Current State of Animal Taxonomy
1. Modern animal taxonomy was established using evolutionary systematics and cladistic revisions.
2. Total use of cladistic principles would require abandonment of Linnaean ranks.
3. A new taxonomic system called Phylocode is being developed as alternative to Linnean taxonomy.
4. This system replaces Linnean ranks with codes that denote the nested hierarchy conveyed by
cladograms.
5. Sister groups relationships are clear descriptions that will replace “mammals evolved from reptiles”
paraphyletic statements that are problematic.
6. The terms “primitive,” “advanced,” “specialized” and “generalized” are used for specific
characteristics and not for groups as a whole.
7. Likewise, avoid calling a living species of group of living species “basal”; rather, the term is best
reserved for describing branch ponts on phylogenetic tries.
8. Cladistics causes confusion by using “bony fishes” to also include amphibians and amniotes, reptiles
to include birds, but not some fossil forms, etc.
10.5. Major Divisions of Life
1. Aristotle’s two kingdom system included plants and animals; one-celled organisms became a
problem.
2. Haeckel proposed Protista for single-celled organisms in 1866.
3. In 1969, R.H. Whittaker proposed a five-kingdom system to distinguish prokaryotes and fungi.
4. Woese, Kandler and Wheelis proposed three monophyletic domains above kingdom level—Eucarya,
Bacteria and Archaea—based on ribosomal RNA sequences.
5. Retaining Whittaker’s five kingdoms, the Eucarya are paraphyletic unless Protists are broken up into
Ciliata, Flagellata and Microsporidia, etc. (Figures 10.11)
6. More revisions are necessary to clarify taxonomic kingdoms based on monophyly.
7. “Protozoa” are neither animals nor a valid monophyletic taxon.
8. “Protista” is not a monophyletic kingdom but is most likely composed of seven or more phyla.
10.6. Major Subdivisions of the Animal Kingdom
1. Phylogenetic relationships among animal phyla have been difficult to resolve by
morphological and molecular characters.
2. Animal phyla have been informally grouped based on embryological and anatomical traits.
3. Protozoa constitute many phyla and none of them belong within the animal kingdom.
4. Eumetazoa include all animal phyla except Porifera and Placozoa and is divided into Radiata and
Bilateria.
5. Bilateral animals are divided into deuterostomes and protostomes based on their embryological
development, with further division of Protostomia into taxa Lophotrochozoa and Ecdysozoa.
Lecture Enrichment
1. Describing the problems with Aristotle’s land-air-sea classification provides students with an idea of the
limitations of some classification schemes logical for their time.
2. One effective but time-consuming method to understand traditional, phenistic and cladistic classification is to
work through one simple set of animals or plants by developing a table of traits and a similarity matrix. For
10-4
Chapter 10 - Classification And Phylogeny Of Animals
instance, ask for six common animals, form a chart of features they share then build a phylogeny on number of
shared traits.
3. Ask students why drug research proceeds first to rats, then to monkeys, and then moves to human trials. Ask for
applications of the predictive power of a phylogeny that more closely matches evolutionary history.
4. Several decades ago, some scientists predicted numerical taxonomy would make classification a technician’s job.
A decade ago, molecular systematists proclaimed they would provide the ultimate key to objective classification.
Students who fully comprehend the work of systematics will understand this is a field that integrates all of
biology.
Commentary/Lesson Plan
Background: Most students will have used high school textbooks that addressed classification in a classical manner
and assume it is a matter of memorizing the hierarchy of names. Most secondary biology teachers lack coursework in
systematics and therefore student misconceptions are likely.
Misconceptions: Many students have been confused by biology teachers holding up a specimen and asking students to
“name” (only one scientist gets to “name” a species), “classify,” (group a species with its closest relatives) or
“identify” it, this final word being the correct usage. Identifying an organism by name is not classifying it;
“classifying” is grouping and it takes three organisms to classify so that two are more closely related than they are to a
third. In common life, these terms are used interchangeably, but biology majors must be more careful in their use of
these terms, the process starts now. Students may assume that the number of species to be named is finite, as are the
contents of an auto parts store or a grocery store; the continual evolutionary change dimension will not be evident.
Both cladograms and phylogenies will be assumed to lead to “higher” forms on the right hand side. In practice, each
branching of a phylogeny or node of a cladogram can rotate like an inverted mobile and more recent derived forms can
be placed to the left or right at each branch.
Schedule: If you intend to work through the concepts of “synapomorphy,” “clade,” “monophyly,” etc. and provide
detailed examples of the schools of systematics, add another hour.
10-5
Chapter 10 - Classification And Phylogeny Of Animals
cat-like, and rabbit-like animals but the Australia versions of these mammals have pouches (they are marsupials)
while the U.S. counterparts are placental. Why is the concept of homology important here?
10-6
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CHAPTER XVI.
(1776-1826.)
very fair instance of Jacobite sentiment
A PLEBISCITE
in London, in the year 1777, presents FOR THE
itself in a record by Boswell, in his ‘Life STUARTS.
of Dr. Johnson.’ The doctor, in argument
with the Whig Dr. Taylor, insisted that the popular
inclination was still for the Stuart family, against that of Brunswick,
and that if England were fairly polled, the present king would be sent
away to-night, and his adherents hanged to-morrow!’ Taylor
demurred, and Johnson gave this as the ‘state of the country.’—‘The
people, knowing it to be agreed on all hands, that this king has not
the hereditary right to the crown, and there being no hope that he
who has it can be restored, have grown cold and indifferent upon the
subject of loyalty and have no warm attachment to any king. They
would not, therefore, risk anything to restore the exiled family. They
would not give twenty shillings a piece to bring it about; but if a mere
vote could do it, there would be twenty to one; at least, there would
be a very great majority of voices for it. But, Sir, you are to consider
that all those who consider that a king has a right to his crown, as a
man has to his estate, which is the just opinion, would be for
restoring the king who certainly has the hereditary right, could he be
trusted with it; in which there would be no danger now, when laws
and everything else are so much advanced, and every king will
govern by the laws.’ It was in the same year, 1777, that Johnson
called the design of the young Chevalier to gain a crown for his
father ‘a noble attempt;’ and Boswell expressed his wish that ‘we
could have an authentic history of it.’ More than a generation had
passed away since the attempt had failed, but Johnson thought the
history might be written: ‘If you were not an idle dog, you might write
it by collecting from everybody what they can tell, and putting down
your authorities.’ It was shortly after that, hearing of a Mr. Eld, as
being a Whig, in Staffordshire, Johnson remarked, ‘There are rascals
in all counties.’ It was then he made his celebrated assertion that ‘the
first Whig was the Devil;’ but this Jacobite definition was provoked by
Eld’s coarse description of a Tory as ‘a creature generated between
a nonjuring parson and one’s grandmother.’ Lord Marchmont thought
Johnson had distinguished himself by being the first man who had
brought ‘Whig’ and ‘Tory’ into a dictionary.
‘Nonjuring parsons’ still existed; but the hierarchy was all but
extinguished.
In the last week of November 1779, reverential THE LAST OF
groups were assembled in Theobald’s Road, to THE
witness the passing to the grave of the last nonjuring NONJURING
bishop of the regular succession—Bishop Gordon. BISHOPS.
There was no demonstration but of respect. Yet there must have
been some Jacobites of the old leaven among the spectators;
though many Nonjurors were not Jacobites at all. To this record may
be added here the fact that in St. Giles’s churchyard, Shrewsbury, lie
the remains of another nonjuring bishop, William Cartwright, who is
commonly called ‘the Apothecary,’ because, like other bishops of the
sturdy little community, he practised medicine. Cartwright (who came
of the ‘Separatists,’ a division which started about 1734, with one
bishop) always dressed in prelatic violet cloth. Hoadley once
surprised a party at Shrewsbury by saying, ‘William Cartwright is as
good a bishop as I am.’ Cartwright hardly thought so himself, for in
1799, in which year he died, he was reconciled to the established
church, at the Abbey in Shrewsbury, by a clergyman who in his old
age revealed the fact to a writer who made it public in 1874, in the
‘Dictionary of Sects, Heresies, and Schools of Thought,’ edited by
the Rev. John Henry Blunt. No reason is given why the alleged fact
was made a mystery of for so long a period.
The very last of all the nonjuring bishops, one of the irregular
succession, died in Ireland in 1805, namely, Boothe. He was
irregularly consecrated by Garnet, who had been consecrated by
Cartwright, who had been consecrated by Deacon. Nonjuring
congregations, in London and elsewhere,—they generally met in
private houses,—diminished and dissolved. Here and there, a family
or an individual might be met with who would use no Prayer Books
but those published before the Revolution of 1688. Probably, the last
Nonjuror (if not the last Jacobite) in England died in the Charter
House, London, in 1875—the late Mr. James Yeowell, for many
years the worthy and well-known sub-editor of ‘Notes and Queries.’
To him, the true church was that of Ken, and his true sovereign was
to be looked for in the line of Stuart; but Mr. Yeowell acknowledged
the force of circumstances, and was as honest a subject of Queen
Victoria as that royal lady could desire to possess.
The Jacobite and Nonjuring pulpits were THE JACOBITE
unoccupied and silent, but the Muses manifested MUSE.
vitality. The tenacity, and one might almost say, the
audacity of Jacobite loyalty was well illustrated in 1779 by the
publication of a collection of songs, under the title of ‘The True
Loyalist, or Chevalier’s Favourite.’ In one of the ballads both Flora
Macdonald and Charles Edward are alluded to:—
Over yon hills and yon lofty mountain,
Where the trees are clad with snow;
And down by yon murm’ring crystal fountain,
Where the silver streams do flow;
There fair Flora sat, complaining
For the absence of our King,
Crying, ‘Charlie, lovely Charlie!
When shall we two meet again?’
At this period, the unhappy Charles Edward was neither lovely nor
loveable. His ballad poet, above, has paraphrased, or parodied, a
popular song, ‘Over Hills and high Mountains,’—but so ill, with
excess or lack of feet, indifferently, as to serve the measure with the
arbitrary despotism with which the Stuarts themselves would have
visited Church and Constitution.
It will be remembered that when Jacobite Johnson JACOBITE
was pensioned, the English language did not suffice JOHNSON.
to give expression to his feelings. He was obliged to
borrow a word from France: he was pénétré with his Majesty’s
goodness. In 1783,—weighing Stuart against Brunswick, Johnson
borrowed a word from the same foreign source, to disparage the
House of Hanover. It must be confessed that Dr. Johnson’s
Jacobitism had become a ‘sentiment,’ in 1783. He could then
indignantly denounce the factious opposition to Government, and yet
account for it on Jacobite principles. He imputed it to the Revolution.
One night, at Mrs. Thrale’s house in Argyle Street, where the
conversation turned on this subject, ‘Sir,’ said he, in a low voice,
having come nearer to me, while his old prejudices seemed to be
fermenting in his mind, ‘the Hanoverian family is isolée here. They
have no friends. Now, the Stuarts had friends who stuck by them so
late as 1745. When the right of the king is not reverenced, there will
not be reverence for those appointed by the king.’
In June of the following year, 1784, Johnson made
BOSWELL, ON
a remark which very reasonably struck Boswell ‘a ALLEGIANCE.
good deal.’—‘I never,’ said Johnson, ‘knew a Nonjuror
who could reason.’ On which observation and on the position of the
Nonjurors and their Jacobite allegiance, generally, Boswell makes
this comment:—‘Surely, he did not mean to deny that faculty to many
of their writers,—to Hickes, Brett, and other eminent divines of that
persuasion, and did not recollect that the seven Bishops, so justly
celebrated for their magnanimous resistance to arbitrary power, were
yet Nonjurors to the new Government. The nonjuring clergy of
Scotland, indeed, who, excepting a few, have lately, by a sudden
stroke, cut off all ties of allegiance to the House of Stuart, and
resolved to pray for our present lawful Sovereign by name, may be
thought to have confirmed this remark; as it may be said that the
divine, indefeasible, hereditary right which they professed to believe,
if ever true, must be equally true still. Many of my readers will be
surprised when I mention that Johnson assured me he had never in
his life been in a nonjuring meeting-house.’—Johnson’s disrespect
for the reasoning powers of the Nonjurors was still less intense than
his detestation of the Whigs. Of some eminent man of the party, he
allowed the ability, but he added, ‘Sir, he is a cursed Whig, a
bottomless Whig, as they all are now.’
Walpole was satisfied that the Stuart race was effete, and that the
family was incapable of exciting the smallest sensation in England.
He could not, however, pass over an incident in ‘the other family.’
In allusion to the Prince of Wales and the Roman Catholic widow
(of two husbands) whom he married,—Mrs. Fitzherbert, he says:
1786, ‘We have other guess matter to talk of in a higher and more
flourishing race; and yet were rumour;—aye, much more than
rumour, every voice in England—to be credited, the matter,
somehow or other, reaches from London to Rome.’ Happily, no new
‘Pretender’ arose from this extraordinary union.
In this year, in the month of July, the comedy of
A JACOBITE
‘The Provoked Husband’ was played at the ACTRESS.
Haymarket, ‘Lady Townley, by a Lady, her 1st
appearance in London.’ The lady and the incident had some interest
for those who held Jacobite principles. They knew she was the
daughter of an old Scotch Jacobite, Watson, whose participation in
the ’45 had perilled his life, ruined his fortune, and caused him to fly
his country. He died in Jamaica. His widow returned to Europe, and
brought up the family, creditably. In course of time; Miss Watson
married a paper-manufacturer, or vendor, named Brooks. His early
death compelled her to go on the stage; her success, fair in the
metropolis, was more brilliant in Dublin, Edinburgh, and other
important cities, especially where Jacobite sympathy was alive. It is
curious that in Boswell’s account of the tour to the Hebrides with
Johnson, under the date, September 7th, 1773, when they were at
Sir Alexander Macdonald’s, at the farm of Corrichattachin, in Skye,
among the things which he found in the house was ‘a mezzotinto of
Mrs. Brooks, the actress, by some strange chance in Skye.’ The
portrait, in 1773, was not that of an actress; nor was the lady then
Mrs. Brooks; but that was her name, and such was her profession
when Boswell published his Life of Dr. Johnson, in 1791; at which
time, however, he was not aware of her Jacobite descent. Some
persons, unpleasantly advanced in years, recollect old Mrs. Brooks’s
powerful delineation of Meg Murdockson, in T. Dibdin’s ‘Heart of Mid
Lothian,’ about the year 1820, at the Surrey Theatre, and they
suggest that she was the old Jacobite’s daughter.
In the year in which the Jacobite’s daughter made
BURNS’S
her first appearance in London, as ‘Lady Townley,’ ‘DREAM.’
Burns wrote the verses which he called ‘A Dream,’
with this epigraph:—
Thoughts, words, and deeds the Statute blames with reason,
But surely Dreams were ne’er indicted Treason.
The poet then dreams of being at St. James’s on the king’s birthday,
and addressing George III. in place of the Laureate. The feeling
expressed was no doubt one that had come to be universal,—
namely, of respect for a monarch and his family, about whom,
however, the poet could see nothing of that divinity which was
supposed of old to hedge such supreme folk. But Burns recognised
a constitutional king, from whom he turned, to attack his responsible
ministers:—
Far be’t frae me that I aspire
To blame your legislation,
Or say ye wisdom want, or fire,
To rule this mighty nation.
But, faith! I muckle doubt, my Sire,
Ye’ve trusted ’Ministration
To chaps who, in a barn or byre,
Wad better fill’d their station
Than courts, yon day.
In the following year, Burns still more satisfactorily BURNS ON
illustrated the general feeling as being one of loyalty THE
to the accomplished fact in the person of the king at STUARTS.
St. James’s, but with no diminution of respect for the
royal race that had lost the inheritance of majesty. This the Scottish
bard expressed in the ‘Poetical Address’ to Mr. W. Tytler. He
lamented indeed that the name of Stuart was now ‘despised and
neglected,’ but, he adds:—
My fathers that name have revered on a throne;
My fathers have fallen to right it.
Those fathers would spurn their degenerate son,
That name should he scoffingly slight it.
Still, in pray’rs for King George, I must heartily join
The Queen and the rest of the gentry:
Be they wise, be they foolish, is nothing of mine;
Their title’s avow’d by my country.