Flashcards in Chordates Deck (469)
1
Chordate zoology is what type of biology
whole-organism biology (not just cell, function, form,.. everything)
2
organismal biology
research at the level of the whole organism, integrated over structure, function, ecology, and evolution
3
organism structure
anatomy, morphology
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organism function
physiology, behaviour
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evolution
phylogeny
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functional morphology
focuses on the link between form (morphology) and performance
7
ecological morphology
focuses on the link between performance and ecology (ecomorphs, ecomorphotypes)
8
ecomorph
species with the same structural habitat/niche, similar in morphology and behavior, but not necessarily close phyletically
9
ecomorphotype
Any morphological modification caused by, or related to, specific ecological conditions
10
integrative biology
near synonym for organismal biology; brings different aspects of organisms and their environment together
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comparative biology
an area of research that attempts to explain biodiversity and its adaptive radiation in a phylogenetic (historical) framework (comparative method)
how phylogeny tests/explains hypotheses
12
natural history
scientific study of the organism in its natural surroundings
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kinds of chordates
Tunicata
Cephalochordata
Vertebrata
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Tunicata includes
tunicates and sea squirts
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Tunicata was formerly called
Urochordata (also protochordates- not valid taxonomic name)
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Cephalochordata includes
lancelets (amphioxus)
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Numbers of chordate species
Tunicata - 2150
Cephalochorddata - 25
Vertebrata - 63 600
*numbers are always changing
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tunicates
free-living larva, sessile adult
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shared characteristics of chordates
notochord
dorsal hollow nerve cord
pharyngeal gill slits
endostyle
muscular postanal tail
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notochord
incorporated in vertebral column in vertebrates
stiff, flexible rod, runs length of back, provides connections for body muscles, and support
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dorsal hollow nerve cord
spinal cord with brain at anterior end in vertebrates
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pharyngeal gill slits
slits in pharynx/throat region through which water passes and food particles are filtered out, involved in filter-feeding and gas exchange
retained in fish- gas exchange
tetrapods- gill slits disappear in adults
gill arches become jaws and other structures in vertebrates
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endostyle
ciliated groove before larynx
secretes mucus to trap food
homologous with thyroid gland in vertebrates
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muscular postanal tail
extension of the body that runs past the anal opening
only present in embryonic stage of humans
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paedomorphosis
retaining juvenile characteristics into adult hood
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sister group to vertebrates
urochordata (most current theory)
vertebrates and urochordates are sister group to cephalochordates
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pattern vs. process
phylogeny vs. scenario
current phylogenetic theory (pattern), process (scenario) yet to be determined
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myomeres
blocks of skeletal muscle tissue found commonly in chordates. commonly zig-zag, "W" or "V"-shaped muscle fibers
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possible first chordate
Pikaia
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aspects of morphology (anatomy)
comparative
functional
transitional
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comparative morphology
similarities/differences between groups
ex. heart
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functional morphology
how organisms are equipped to deal with different situations
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transitional morphology
macroevolutionary change- how we go from one form to another
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why study chordates
intrinsic interest
vertebrates are us
diversity
evolutionary record
model organisms
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chordate diversity
masses range from 10^-4 - 10^5 kg! (larva to blue whale)
altitudes range 58km- deep ocean - over himalayans
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why evolutionary record is important for studying chordates
they have the best preservation of all organisms
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model organisms
amphibians- developmental biology
birds- population biology
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vertebrates characteristics
internal skeleton
vertebral column with cranium at anterior
spinal nerve cord with brain at anterior
neural crest
HOX genes
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vertebrate internal skeleton
bone and/or cartilage
40
vertebral column
individual vertebra, skull at end
rudimentary in lampreys
lacking in hagfish
full formed in gnathostomes
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gnathostomes
jawed vertebrates
42
neural crest
cells/tissue- formed during embryonic development and migrate to head
43
HOX genes
do exist in inverts but more important in vertebrates
vertebrates have the most HOX genes
44
"grades" of vertebrates
"fishes"
Tetrapods
45
paraphyletic
composed of some but not all members descending from a common ancestor
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fishes
paraphyletic- would be monophyletic if we include all groups that have come from fishes (including us)
31,000 species
mainly Osteichthyes
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groups in fishes
Agnathans
Chondrichthyes
Osteichthyes
48
Agnathans
jawless vertebrates- hagfish and lampreys, Ostracoderms
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Chondrichthyes
cartilaginous fish- sharks, rays, ratfish
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Osteichthyes
bony fish
Crossopterygii, Actinopterygii
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Cossopterygii
Actinistia + Dipnoi (lobe-finned fish- coelacanth + lungfish)
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Actinopterygii
ray-finned fish (most fish)
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Tetrapods
all the rest of the vertebrates
4 feet, lots of secondary loss examples
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Tetrapod examples
Amphibia
Amniota
Aves
Mammalia
55
Amphibia
7000 species
lissamphibians- apodans salamanders, anurans
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Amniota
amniotes-- Reptilia
57
Reptilia
10,000 species- turtles, crocodiles, tuataras, lizards, snakes, amphisbaenians
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Aves
10,000 species
directly derived from reptiles, make reptiles a non-taxonomic group
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Mammalia
5500 species- mammals
mostly placentals, some monotremes and marsupials
60
Ectotherms
"fishes"
amphibians
"reptiles"
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endotherms
generate own body heat metabolically
birds
mammals
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Amniotes
"reptiles"
birds
mammals
(taxonomic group)
63
Anamniotes
"Fishes"
Amphibians
(not a taxonomic group)
64
Sauropsids
"reptiles"
birds
65
synapsids
mammals
66
lampreys
Agnathan- parasitize other fish
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Fossil Agnathans
Ostracoderms- coats of bone armour, very different from living fish
68
Gnathostomata
defined by presence of jaws with teeth
usually have limbs
69
thought to be closest living relative of tetrapods
lungfish (3 species)
70
tetrapoda
limbs with digits for terrestrial locomotion
internal nostrils
tympanic membrane and stapes
strong skeleton
71
internal nostrils
breathing through nose
fish do not breath through nose (except lungfish)
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tetrapod tympanic membrane and stapes
to detect airborne sounds- ear drum
in fish stapes supports skull, in tetrapod it is main bone by which sound is transported into inner ear
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tetrapod strong skeleton
support against gravity (adaptation to terrestrial life)
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Lissamphibia
3 major, distinct morphological groups
thin skin
complex life cycle
numerous departures from theme
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Lissamphibia thin skin
highly permeable
respiratory gas exchange- skin covered in mucus to facilitate exchange
water exchange- drink by skin, lose water through skin-- restricted to moist habitats
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Lissamphibia complex life cycle
includes metamorphosis
aquatic egg-- aquatic larva-- aquatic/terrestrial juvenile and adult
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Amphibian groups
salamanders
anurans
apodans
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Salamanders
mostly north temperate zone
mostly elongate body
mostly limbed- some highly reduced
some permanently aquatic retain larval form, some bipass larval form- direct development, lay eggs on ground, hatch into full formed adult
some pedomorphosis, have gills
79
Anurans
hyper developed back legs
aquatic/terrestrial/arboreal (ex. tree frogs climb w/ suction cups)
no paedomorphic frogs
some cut out larval stage
larval/adult generally radically different
80
unique Anuran forms
one species- female swallows eggs and they develop in her stomach
81
Apodans
(Caecilians)
superficially resemble earthworms/snakes
mostly live hidden in the ground, least familiar order of amphibians, smallest group, carnivores, large mouth
aquatic/burrowing/nocturnal/terrestrial
some have complex lifestyle, some direct development
live bearing- no placenta, hyper developed gills, mostly viviporous
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Amniota
amniotic egg- amniote surrounds embryo
exclusively internal fertilization
83
allantois
waste dump, highly vascularized (blood vessels)
develops to line membrane surrounding the egg- transport system for gas exchange
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amniote shell
holds eggs together, has mating consequences- necessitates internal fertilization, must develop on land (oviparity)
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oviparity
Oviparous animals are animals that lay eggs, with little or no other embryonic development within the mother
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Reptilia
not a taxon group
thicker skin, covered with epidural scutes (keratin)
much more terrestrially adapted, lose water slower
87
scutes
scales
keratin provides mechanical protection, defense against water loss, protection of lipids
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"reptile" groups
turtles
tuataras
lizards
amphisbaenians
snakes
crocodilians
89
turtles
only vertebrate in which ribs are outside
lots of aquatic, some marine, some terrestrial
90
Tuataras
endemic to New Zealand, highly restricted, resemble most lizards, part of a distinct lineage
long, slow life style, last remaining species of diverse group
91
"lizards"
biggest group of "reptiles"
very diverse, worldwide, terrestrial, arboreal
lots of long slender bodies w/o limbs
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Amphisbaenians
derived from lizards, sometimes called worm lizards
burrowing species, swim, mostly tropic, some subtropic, mostly lack limbs, different head shapes for specialization of burrowing
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head types of Amphisbaenians
round head- soft sediment
wedge shaped head- hard substrate
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snakes
mostly have no trace of limbs
many arboreal- long and slender to spread weight out over branches
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rain snake
oval cross section
tail flattened to propel through water
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Crocodilians
~22 species
living forms all fairly similar
body covered with osteoderms
long, elongate skull
extant are only aquatic
tropical/subtropical
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osteoderms
bony deposits forming scales, plates or other structures in the dermal layers of the skin
98
crocodile snout specialization
long, skinny snout- fish eating- quicker swiping through water
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Aves
highly modified reptiles
endothermic
body covered with feathers (keratin)
legs covered with epidermal scutes
wings and other modifications for flight
modified jaw with beak (keratin) and no teeth
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bird feet/beak
tell a lot about overall life ecology
ex. flightless - marine
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bird feathers
aid endothermy
major flight surface of wing
102
Mammalia
endothermic
body covered with hair/fur (keratin, insolation)
highly differentiated dentition
numerous derived traits in skeleton
pinna
103
derived traits in mammal skeleton
squamosal-dentary jaw joint
7 cervical vertebrae (neck vertebrae)- almost all mammals have 7
104
Pinna
the visible part of the ear that resides outside of the head
105
main groups of mammals
monotremes
marsupials
placentals
106
monotremes
lay eggs- platypus, echidna
only a few species
basically same as reptile eggs- hatch then female feeds milk via ducts (not nipples)
107
marsupials
wide diversity of form, mostly in Australia, bears, rodents, carnivores, koala, kangaroo, opossum, wombat
have placenta but not well developed
embryo develops in uterus for short period of time
born at early embryonic stage, crawls up to pouch and attaches until ready to be born
108
marsupial embryo pouch
marsupion
109
placentals
development until birth is in uterus
intimate connection between embryo and mother (nutrients, gas exchange)
110
important parts of evolution
genes, chromosomes, alleles, mutation (many neutral, some lethal), proteins (structural, hormones, enzymes), recombination, dominance, pleiotropy, polygenic traits, epistasis, regulatory genes
111
pleiotropy
single genes code for multiple parts of the body
112
epistasis
interactions between genes
effect of gene depends on presence/expression of other genes
113
regulatory genes
govern expression of genes
114
phenotype =
genotype * environment
115
phenotypic plasticity
ability of an organism to change its phenotype in response to changes in the environment
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heritability
the proportion of observed variation in a particular trait (as height) that can be attributed to inherited genetic factors in contrast to environmental ones
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reaction norms
expression of the phenotype of a particular genotype in different environments
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no genotype environment interaction, reaction norm
phenotype vs. environment graph
variation is constant, parallel for 2 genotypes among different environments
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genotype environment interaction, reaction norm
genotypes are not parallel or constant (lines may cross)
one genotype may produce multiple phenotypes
120
phenotypic flexibility
reversible changes within individuals
ex. plumage variation with season- changeable/reversible phenotype
121
types of selection
artificial selection
(natural) selection
adaptation
directional selection
balancing selection
disruptive selection
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natural selection
variation, heritability, differential survival/reproduction
variation has to be heritable for evolution to occur
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adaptation
a trait that has arisen from natural selection
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three modes of selection
stabilizing
directional
disruptive
125
stabilizing selection
balancing
extreme values are a disadvantage and are 'pruned'
keep population at certain 'optimum'
126
directional selection
one extreme is disadvantageous
leeds to a shift away from one extreme
127
disruptive selection
intermediates are disadvantageous
selection is for the extreme- split down the 'middle'
128
example of directional selection
high proportion of unbounded snakes- selection against banding- population pushed primarily to unbounded
banded snakes remain in population due to migrations from mainland
balance between migration and selection
129
example of balancing selection
UK birth weights- low mortality in mid weight babies
small babies underdeveloped, large babies died during birth
selection for optimal birth size- med size babies were commonest, had best chance of survival, most likely to pass on their characteristics
130
selection isn't the only factor that determines change
genetic drift
founder effect
131
genetic drift
genes that wouldn't normally be passed on are passed due to small population- selection goes in different direction that it normally would
132
founder effect
small founding colony of individuals will have more limited make-up than the population they came from, depends on founding colony being small
133
species/speciation concepts
multiple species concepts
discontinuities
morphospecies
biological species
hybridization
asexual reproduction
134
discontinuities
breaks between species (ex. morphology)
135
multiple species concepts
many definitions of species
the 2 we will focus on are biological species concept (BSC), morphological species concept
136
BSC
depends on sexual breeding, must be able to interbreed, hard to test, depends on species exhibiting sympatry
discontinuity- reproductive isolation
137
sympatry
species that occur in same geographic area
138
why species concept can be hard to test
if they don't come together naturally there is no way of knowing if they naturally mate- bringing them together in the lab is artificial
139
morphospecies
different species should look different
domestic dogs- haven't speciated
140
sibling species
different species that look identical
141
sibling species example
Hyla versicolor- tetraploid (gray tree frog)
Hyla chrysoscelis- diploid (Cope's gray tree frog)
142
tetraploid
Triploid/tetraploid chromosomes are polyploidy
Polyploid organisms are those containing more than two paired (homologous) sets of chromosomes. Most species whose cells have nuclei (Eukaryotes) are diploid, one set inherited from each parent.
143
polyphyletic
does not include the common ancestor of all members of the taxon
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paraphyletic
includes the most recent common ancestor, but not all of its descendents
145
species is used to
describe groups that we recognize
describe what the animals themselves recognize
146
stable hybrid zone
ranges of 2 species come together, where they meet there is a zone that consists of hybrids
hybrids usually not sterile but may be less fit
147
hybrid example
manitoba toads- meet in East manitoba, zone where there are hybrids (separate species)
yellow-rumped warbler (sub-species)
gray wolf/eastern wolf/coyote/red wolf/dogs
European water frogs
148
incipient speciation
evolutionary process in which new species form but are still capable of interbreeding; can be the first part of the larger process of speciation
149
yellow-dumped warbler hybrids
Audubon's warbler / Myrtle warbler
100km hybrid zone
graphs show different traits within/outside of hybrid zone
150
introgression
(introgressive hybridization), movement of a gene (gene flow) from one species into the gene pool of another by repeated backcrossing of an interspecific hybrid with one of its parent species
151
3 known categories of asexual reproduction in vertebrates
recall that BSC requires sexual breeding
hybridogenesis
gynogenesis
parthogenesis
152
hemiclonal
half of females genome is passed on clonally
153
clonal
all of females genome is passed on clonally
154
hybridogenesis
male mates with female but males genetic contribution is discarded at mitosis
egg is fertilized but genetic info. not passed on in next generation
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gynogenesis
mates with male but sperm are not used (salamanders, some fish)
diploid egg, sperm stimulates reproduction but doesn't fertilize
female relies on sperm from heterospecific males to initiate embryogenesis
156
parthenogenesis
males not used at all, diploid, unfertilized egg (clonal)
157
most vertebrates are
bisexual- normal sexual reproduction
158
European water frogs hemiclonal hybridogenesis
Rana ridibunda x R. lessonae = R. esculenta
R. esculenta x R. lessonae = R. esculenta
R. esculenta x R. esculenta = R. ridibunda
R. esculenta x R. ridibunda = R. ridibunda
R. esculenta maintains itself by mating with parent species- at meiosis discards lessonae part of genome and reconstitutes itself
159
usually inviable offspring
R. esculenta x R. esculenta = R. ridibunda
160
origin of R. esculenta
R. ridibunda x R. lessonae = R.esculenta
161
why do males mate with female gynogens (gynogenesis)
selection should favor males that gynogenesis
sexual females increase their preference for males whom they observe consorting with female gynogens
162
elements of speciation
occur in variable orders/rates/ranges/time, process of species formation, multiplication of species, complex process that involves multiple phenomena
elements: reproductive isolation, genetic divergence, phenotypic divergence
163
Bulloch's Oriole and Blatimore Oriole
two morphologically distinct species, but they are interfertile
two morphospecies, but one or two biological species?
164
interfertile
capable of interbreeding
165
cryptic species
morphologically similar, genetically distinct, incapable of interbreeding
166
cryptic species example
African Savanna Elephant, African Forest Elephant
167
speciation can occur
allopatrically, parapatrically/peripatrically, or sympatrically
168
allopatric speciation
geographically separate
169
parapatric speciation
geographically adjacent
170
sympatric speciation
geographically coincident (same area)
171
non-selective speciation
'by accident'- random drift in small founding populations
172
natural selection speciation
mutation-order speciation
ecological speciation
173
ecological speciation
evolution of reproductive isolation between populations by divergent natural selection arising from differences between ecological environments
indirect or direct
174
indirect ecological speciation
by-product speciation
reproductive isolation occurs as an incidental by-product of adaptation to different environments
175
direct ecological speciation
selection directly favours reproductive isolation
ex. if hybrids are less viable
176
allopatric (geographic) speciation
vicariant event- splitting events/barriers cause species to adapt independently to conditions (ex. glaciation)
dispersal- disperse to new area (ex. island), may lead to founder effect (ex. galapagos finches)
177
Possible outcomes of secondary contact after long periods of separation
partial/complete reproductive isolation- basically separate species
hybrid zones
complete introgression- interbreed as if never separated
selection for reproductive isolation- behavioural differences
178
lentic
standing water
179
lotic
running water
180
ecological speciation changes in fishes
marine-freshwater
physio-chemical transitions
lotic-lentic transitions
discrete river habitat
water depth
benthic-open water
benthic substrate shifts
piscivory
durophagy
intrinsic incompatibilities
divergent sexual selection
181
piscivory
eating of fish
182
durophagy
eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs
183
increasing potential for gene flow in absence of differentiation
decreasing spatial scale
allopatric-- parapatric-- sympatric
184
ecological speciation example
repeated parallel speciation in endependent populations of sticklebacks, regulation in gene due to reproductive isolation
deep water- prominent spine, predator protection
shallow water- absence of fin, harder for insect larvae to attach
185
Replicated (parallel) ecological speciation in lizards
habitat matching- different species show the same coloration changes to match light/dark soil habitats
186
punctuated equilibrium
little net evolutionary change for most of geological history, remaining in an extended state called stasis, disrupted by abrupt change
187
phyletic gradualism
speciation is slow, uniform and gradual
When evolution occurs it is usually by the steady transformation of a whole species into a new one
188
microevolution
microphylogeny, microecology
change over short time and small spatial scale
ex. population genetics, shifts in color patterns, polumorphisms
189
macroevolution
historical ecology, macroecology
change over long time scales and large spatial scales
taxonomic and transformational implications- major changes in body form
190
Historical Ecology
taxic-- speciation, cospeciation
transformational- adaptation, coadaptation
191
macroevolutionary phenomena
origin of new structures
discontinuities in 'adaptive space'
rates of evolution
causes and natures of radiations
causes and natures of mass extinctions
192
fundamental to biology
phylogeny, taxonomy, systematics
193
why phylogeny/taxonomy/systematics are important in a world with decreasing biodiversity
different conservational issue depending on whether or not separate species
194
BC spotted frog
found to be 2 species; 1 restricted to coastal Oregon, BC; new species is very restricted and need immediate conservation help
195
Tuatara Sphenodon (lizard) endemic to NZ
one species found to be two, separate conservation plans had to be made
196
folk taxonomy
local and simple
classifying animals based on known plants, local
used in everyday language- 'bugs' 'shrubs'
197
world exploration
disruption to local order
198
Linnaeus
1700s, Systema Naturae binominal system
subjective, based on intuition, experience, 'umwelt', mix of art and science
species names have 2 parts, first subjective system
199
evolutionary taxonomists
1800s, influenced by Darwin
classification should reflect evolutionary relationships
shift in point of view but not methods
a species is what a good taxonomist says it is
200
making taxonomy a science
1950s, injecting objectivity
201
numerical taxonomy
unweighted characters, quantitative
take as many characters as can, try to find similarities to make unbiased connections
202
molecular taxonomy
proteins, RNA, DNA, comparison of apples and oranges
differences between species in relation to proteins and DNA
first time characters could be compared in vastly different organisms
203
Cladistic taxonomy
phylogenetic taxonomy
shared evolutionary novelties
not all traits are useful, some are more important than others- shared derived traits
204
fungi are closely related
to animals (more than to plants)
205
classification
ordering of organisms into sets based on relationships
relationships defined in various ways- phylogeny, resemblance
206
identification
allocation of previously unidentified specimens to the correct sets, as in 'keying'
207
taxonomy
the theoretical study of classification, including principles and procedures (rules)
208
taxon
general term for any taxonomic group of any rank
209
systematics
study of diversity and interrelationships of organisms, causes and origins of relationships including zoogeography
broader than taxonomy
210
nomenclature
very narrow term, having to do with actual naming
211
types of classification
cross-tabulation
linearly ordered list- alphabetical, numerical (not a good classification, just a list)
hierarchy- by some criterion of similarity
212
cross-tabulation
take 2 traits and compare them in 2x2 grid
ex. endothermy/ectothermy, eyes/no eyes
213
biological classifications
hierarchical - groups nested within groups
based on similarity- morphological, genetic
usually reflect evolutionary relationships
214
taxon ranks in Linnean classification system
Kingdom- Phylum- Subphylum- Class- Subclass- Order- Suborder- Family- Genus- Species (binomen)- Subspecies (trainmen)
215
plural of taxon
taxa
216
order names end in
a
Order Squamata - not the Squamata order
or we can talk about squamates, or squamate reptiles
217
family names end in
idea
Family Viparidae- not the Viperidae family
or "viperirds" or "viperid snakes"
218
subfamily names end in
inae
Subfamily Viperinae
or "viperines" or "viperine snakes"
219
nomenclature rules are found
in International Code of Zoological Nomenclature
220
kinds of similarity
similar origin (developmental or evolutionary)
similar function
similar structure
221
similarity in origin example
articulation of jaw in mammals- squamosal and dentary
in all tetrapods except mammals- articular and quadrate
222
articular and quadrate in mammals
middle ear- different fn and structure but similar origin
articular = malleus
quadrate = incus
223
middle ear ossicles
3 in mammals- incus, malleus, stapes
1 in other tetrapods- stapes (trace back to fish)
224
similarity in structure example
3 different kinds of wings- bird, pterosaur, bat
forelimbs of tetrapods- similar features; ulna, radius, humerus
parts shaped differently with same basic elements
225
similarity in function example
noise-making in snakes- rattling, tail-vibrating, hissing, stridulation, cloacal popping
226
rattling in snakes
Crotalus: series of interlocking segments make up rattle, special shaker muscle vibrates at rapid rate for long time,
add segments to rattle every time they shed their skin, babies only have one segment and can't rattle
227
tail-vibrating and hissing in snakes
Pituophis: shake tail to make noise in dry leaves (confused with rattlers)
specialized keel increases loudness of hissing (~10 decibels higher with keel- decibels are log scale)
228
stridulation in snakes
Echis: (scale-rubbing) specialized scale on side of body, move body segments against each other to make noise
229
cloacal popping
Micruroides, Gyalopion: force air out of cloacal (opening for intestine, reproductive, urinary tract) and make a popping noise
230
kinds of similarity
homology
homoplasy
analogy
231
homoplasy
"false" evolutionary resemblance
parallelism, convergence, reversal, loss, mimicry
can confound construction of phylogenies
but can be used to make inferences about adaptation
232
"false" evolutionary resemblance
phenotypic similarities mislead us to relationship
233
reversal
loss of trait back to original state
234
confound construction of phylogenies
sorting out homoplasies from other similarities is key
235
Convergent evolution (this is the term we will use)
convergence- evolution of similar phenotypic features independently in different lineages, usually from different features and developmental pathways
236
Parallel evolution
parallelism- evolution of similar/identical features independently in related lineages, usually based on similar modification of the same developmental pathways
237
convergence in African Cichlids
fish in two different lakes look similar- mimic each other
fish in same lake look different but are more similar to each other than to fish in other lake
238
convergence in mammals
similar body forms through time
many marsupials look similar to eutherians (placentals)
mole-- marsupial mole
flying squirrel-- sugar glider
woodchuck-- wombat
239
convergence of echolocation
different kinds of echolocation arise independently multiple times (bats, dolphins)
240
homology
"true" evolutionary resemblance, a relative term
similarity of origin is important, to some extent
taxic or transformational
241
homology is determined from
comparative anatomy, fossil record, developmental biology, distribution of character states among taxa (using parsimony)
242
reptile to mammal transformation homology
jaw joint to middle ear ossicle
polarity of the trait = direction of change over time
243
taxic homology
shared evolutionary novelties that help define taxa
244
tetrapod wings
show how the level of analysis changes the definition, relative terms
homologous as tetrapod limbs
homoplasious as wings (bird, pterosaur, bat all independently derived wings)
245
cladistics
phylogenetic systematics
grouped based on 1+ shared unique characteristics from the group's LCA
246
monophyletic group
ancestor plus all descendants
247
paraphyletic group
monophyletic group with some descendants missing
ex. "reptiles" birds missing, would be a correct taxonomic group if we included birds
248
polyphyletic group
group composed of members separated by two or more ancestors
usually constructed by homoplasies
249
homoplasy
character shared by a set of species but not present in their common ancestor
250
plesiomorphies
ancestral, 'primitive' traits
251
apomorphies
derived, 'advanced' traits
252
symplesiomorphy
shared ancestral trait
253
synapomorphy
shared derived trait
254
we want to construct phylogenies with only
synapomorphies (shared derived traits), only clades
ex. all vertebrates have hearts does not set them apart from other taxon
255
grade vs clade
grade- group of species united by morphological/ physiological traits, gives rise to another group differing from ancestral condition- not considered part of the ancestral group, same grade can be in different clades
ancestral group- not phylogenetically complete, doesn't form a clade, represents a paraphyletic taxon
256
Linnaean classification of Vertebrates
Class Chondrichthyes- sharks, rays
Class Osteichthyes*- bony fish
Class Amphibia- lissamphibians
Class Reptilia*- reptiles
Class Aves- birds
Class Mammalia- mammals
* paraphyletic groups
257
anagenesis
change in form within a lineage
258
cladogenesis
splitting events
259
grades =
paraphyly, group which does not include all its descendents
"fish" and "birds" are paraphyletic
260
clades =
monophyly, taxon (group of organisms) which forms a clade, meaning that it consists of an ancestral species and all its descendants
261
grades can be attributable to
synapomorphy- monophyletic groups (grade = clade, mammals are a grade and a clade)
symplesiomorphy- paraphyletic groups (reptile, fish)
convergence- polyphyletic
262
symplesiomorphies
characters shared by everybody, everybody's got these traits so they don't help characterize one group
263
synapomorphies
unique traits that maybe can be used to categorize one group vs. another
264
autapomorphies
derived traits that are unique to a particular taxon
265
each person is, in certain respects like all other people, like some other people, and like no other person
all other people- symplesiomorphies
some other people- synapomorphies
no other people- autapomorphies
266
in phylogeny we are trying to recognize sister taxa
a synapomorphy is present only in 2 sister-taxa and is the inherited autapomorphky from their LCA. ID'ing a character as a synapomorphy is essential for recognizing sister taxa
267
phylogenetic systematics
taxa (clades) defined by shared derived traits only
evolutionary relationship (phylogeny) among taxa represented by a branching diagram (cladogram)
sister clades receive equal rank (no taxon labels)
all taxa are monophyletic
groups defined by shared ancestral traits have no taxonomic value (may be important in other ways)
268
dichotomous branching
each branch is a sister group
269
fully resolved phylogeny
each point has only two branches
>2 - polytomy
270
phylogenetically defined taxa are always
monophyletic
taxon = monophyletic group = lineage = clade
271
groups defined by shared ancestral traits have no taxonomic value (may be important in other ways)
ectotherms- not a natural grouping, has eco-physiological meaning
272
fossil problem
incomplete specimens
incomplete record through time
mostly hard parts, few soft tissues
no molecular data
maybe unique derived traits that we can not see
273
types of clades
node-based
stem-based
apomorphy-based
274
node-based
crown group
based on extant forms (more information)
just below node of sister group
275
stem-based
total group
consists of crown group + extinct forms (stem) that are part of the ancestral lineage of crown group
just above node below sister group
276
apomorphy based
based on first known appearance of a particular derived trait
ex. first appearance of limbs in fossil record
somewhere in between sister group node and node below
277
crown group defined
by extant groups but can contain extinct groups
278
why phylogeny is more useful than Linnean system
more complete and informative classification
phylogeny recoverable from taxonomy and vice versa
can use a list to create cladogram (isomorphic, equal rank get equal indentation)
279
phylogeny limitations
polarity and/or homology of many traits is not unambiguously established (whats the direction of change)
evidence is evidence not proof, many disagreements on 'correct' phylogeny (still much debate, where do turtles go)
molecules vs. morphology
everything is provisional
280
molecules vs. morphology
modern phylogenies usually derived from molecular data
we then can map other traits onto the phylogeny and interpret them (ex. convergences)
mostly, now, start with molecular phylogeny and then map morphological
281
ingroup
group of taxa whose relationships we want to resolve
282
outgroup
comparison taxon closely related to the taxa of interest
sometimes 2 or more outgroup taxa are used, one more closely related and one more distantly related
283
outgroups used to determine
how the cladogram should be rooted
ex. help determine polarity (direction) of changes in traits
284
calibrating phylogeny so branch lengths give estimate of time since separation
fossil record
base phylogeny on genes and calibrate via molecular clock
use ancient DNA
285
ghost lineage
incomplete history, missing fossil record implied by sister group
286
NGS
next-generation DNA sequencing (paleogenomics)
some contamination problems
can bring evidence on questions such as time since divergence
can sequence hundreds of thousands of years back
can give an idea of process rather than just pattern
287
Phylogeny
evolutionary chronicle
branched chronological series of character state changes along lineages (what)
clear methodology (cladistics)
288
Genealogy
historical narrative
casual statements, explanation, interpretations, evolutionary scenarios (how and why)
no clear methodology- gaps between taxa filled by fossils, extant taxa, imagination, intuition
289
the comparative method
using comparisons across species that have evolved independently
290
why should we care about systematics
systematics are fundamental to biology and can tell us alot
291
problems comparing characteristics
shared history- lineage specific effects (non independence)
phylogenetic effects/differences- history matters
292
independent comparison method for 2 characters in a single phylogeny
sister taxa are averaged to give node value
differences are taken between sister taxa for d1 and d2
differences are taken between nodes d3
differences compared for characters and then the 2 characters differences are plotted against each other
293
number of rooted, bifurcating, labeled trees for n species
(2n-3)! / (2^n-2)(n-2)!
294
problems constructing phylogenies
large number of possible tree topologies (15 trees for n=4)
different analytical methods give different answers
numerous different trees for same group
fossil forms incomplete
295
analytical methods of phylogeny
parsimony and variants
compatibility (character "cliques")
distance matrix methods
likelihood methods
others, all quantitative
296
consensus tree
constructed when we have 2 or more different trees for the same group
297
parsimony
Occam's Razor
the simplest, most plausible hypothesis is the obvious place to start
if you hear hoofbeats, think horses not zebras
cladogram w/ fewest evolutionary steps is best starting hypothesis
298
inferring characters in poorly known taxa
modern analogues
extant phylogenetic bracket
299
extant phylogenetic bracket
if a character is exhibited by bracketing extant species, then most likely exhibited by internal branches (extinct species)
ex. parental care seen in crocodilians and birds, so probably in the dinosaurs 'in between'
may have evolved independently but less parsimonious- more evolutionary steps
300
endothermy in dinosaurs?
crocodilians- 4 different dinosaurs- birds
we know that crocodilians are not endothermic, birds are
there are 5 equally parsimonious trees to explain the evolution of endothermy in to birds, so we can't infer where it evolved
301
non-therian synapsids
all synapsids other than mammals
302
nocturnality in non-therian synapsids
study finds synapsids were nocturnal ancestrally
carnivores are predominantly nocturnal or cathemeral
303
cathemeral
irregularly active at any time of night or day, according to prevailing circumstances
304
myrmecophagy
feeding behavior defined by the consumption of termites or ants
305
convergence example
myrmecophagy in Dendrobatids (frogs)
echolocation in bats and dolphins
flightlessness in birds
endothermy in Scombroidean Fishes
306
species that overlap geographically are sister taxa
sympatric speciation
307
sister species do not overlap geographically
allopatric speciation
308
sister species do not overlap and the range of one is smaller than the range of the other
peripatric speciation
309
polymorphism
occurrence together in same habitat of 2 or more discontinuous forms, or "phases", of a species in proportions that the rarest of them cannot be maintained merely by recurrent mutation (genetically determined variants)
310
polyphenism
phenomenon where two or more distinct phenotypes are produced by the same genotype
Multiple phenotypes in population (polymorphic), not based on genotype
phenotypic variation induced by an environmental cue
form of phenotypic plasticity
311
example of polyphenism
amphibian larvae – shifts in morphological phenotype
due to predators, competitors, and prey
cannibalism is induced when high density of conspecifics- leads to larger mouth and teeth
312
why nervous system is useful in phylogenetic research
lots of data to compare
lots of genetic markers
313
origin of chordates, old system and new system
old- vertebrates sister group to Amphioxus
new system- Vertebrates sister group to Tunicates
314
why were tunicates 'moved' in the phylogeny?
found to have a number of fast-evolving genes
very derived, even though they appear basal
no longer a kew organism in understanding
315
mosaic development
something happened early in tunicate development that allowed them to rapidly evolve in a different direction
316
serial mesoderm (segmented) evolution
'hairy' gene in Arthropods and Annelids
homolog of 'hairy' in chordates
~same gene, evolved once
317
chordate characters
pharyngeal gill slits
dorsal nerve cord
segmental muscles/mesoderm
notochord
postanal tail
318
pharyngeal gill slits
probably first to evolve, seen in hemichordates and maybe echinoderms
amphioxus doesn't use as gills- filters food
large surface area for gas exchange
319
segmental muscles/mesoderm
serial myotoes/somites
other inverts. typically circular/longitudinal muscles
in vertebrates only around guts
320
postanal tail
anus not terminal, body extends over primary anus to form tail, new anus is formed (non-terminally)
321
dorsal nerve cord
ventral in proterostome
evolves separately in new place OR inversion, mouth migration (moves down in chordates)
322
mouth formation
delayed, area btw mouth and nerve cord expands
ventral mouth is a derived character-- lead to face evolution
as mouth 'moves' pituatary 'chases' it- moves to oral cavity, then up to underside of nerve cord
323
model organisms for developmental biology of vertebrates
zebrafish, mouse, chick, frog
324
why frog?
easy to obtain large # of eggs, embryogenesis occurs outside body- eggs can be manipulated
325
frog cell cycle
rapid cell cycle- no G1, G2-- increases # of cells in embryo (large cell #)
no zygotic transcription
326
blastula parts
animal pole (AP)- top, blastocoel, superficial cells
Vegetal pole- cells more dense, comprise gut structures
327
gastrulation
series of coordinated cell rearrangements leading to germ layers
equatorial region forms dorsal blastopore lip- bottle cells 'crawl' and involute- displaces blastoceal
328
germ layers
ectoderm
mesoderm
endoderm
329
ectoderm
skin, nervous system
330
mesoderm
muscle, bone
331
endoderm
gut
332
results of gastrulation
endoderm cells were on outside before gastrulation
archenteron formed
AP moved equatorial
blastopore formed (future anus)
333
protostome gastrulation
starts at mouth
334
deuterostome gastrulation
starts at anus
335
upon completion of gastrulation
3 germ layers are evident and layered (endo, meso, ecto)
A-P and dorsal-ventral poles established
preliminary notochord present, terminating in dorsal blastopore lip
336
next stage after gastrulation
neurulation - dorsal ectoderm becomes CNS (central nervous system)
337
mammalian, chick embryo proper
flat sheet of cells, epiblast, prior to gastrulation
cells migrate through primitive groove to give rise to 3 germ layers
338
neural plate stage
neural tube formation
neuroectoderm-- neural groove formation-- neural fold-- neural tube pinches off-- neural crest cells disperse and notochord forms-- epidermis on top
gives rise to brain and spinal cord
339
neural crest
one of defining features of vertebrates- novel
migrate to different regions- even non neural cells
340
neural crest derivatives
PNS- peripheral nervous system
Endocrine and paraendocrine derivatives
pigment cells
facial cartilage and bone
connective tissue
341
sometimes referred to as the 4th germ layer
neural crest
distinguishes vertebrates from other chordates
342
a9.49 cell
always gives rise to pigment- overexpression of 'twist' = loss of melanocyte-- migration is similar to neural crest cell (may be a precursor)
343
closure of neural tube
---neurula stage- formation of somites, tailed, expansion of forebrain, gill area
failure of closure = spina bifida (.7/1000 births, folic acid deficiency)
344
notochord remnants
retained in lancelet, larval tunicate
supports vertebrae in mammals
345
notochord implicated in different cell type generation
signals ectoderm to form CNS
implicated in different cell type generation
move/remove notochord- induce cell diversity in dorsal part of neural tube, in a graded manner (1 22 1)
346
shh
sonic hedgehog secreting cells- gradient directs TF expression- 3TF = 5 domains
gradient of morphogen- cells respond to density = expression of different genes
one signalling factor gives rise to multiple factors
347
tadpole with yellow in head
remnants of yolk, starts feeding after thats gone
348
coqui
'direct developers'
hatch as live frogs- no larval stage
much larger embryo and yolk, quicker maturation, nutritional endoderm, less reliant on water (eggs can be laid on land)
parallel evolution with amniotes
349
frog metamorphosis controlled by
thyroid hormone, T3 tri-iodothyrodine
350
nutritional endoderm
novel cell type, single cell containing yolk, doesn't integrate into frog, connected for feeding, similar to amniote
351
allantois
stores waste
gas exchange
352
chorion
outer most membrane
gas exchange
mammals = placenta
353
polyembryony
more than one embryo from one egg
armadillos always give rise to 4 identical twins, all share same amnion and chorion (very rare in humans- 1% of human twins)
354
to make a phylogenetic tree start with
homologous DNA sequences that you have collected in the lab or downloaded from a database search
align sequences
355
BLAST
basic local alignment search tool - takes query, finds pairwise match, 'hit'
356
Clustal W
gives multiple sequence alignments- doesn't discriminate between non-homologous sequences- this is up to the biologist
357
correctly aligned, non-homologous
ACCTCATC
_C____T_
358
used to manipulate Clustal W results
BioEdit
359
distance matrix
how many differences between sequences
start with columns = sites, rows = sequences, count how many differences between sequences
make new grid, ex. sequences 234 vs. 123 with # differences
360
distance tree
tree branches add up to differences, sized based on differences
works best if you have same amount of data for each sequence- less similarities if comparing a short sequence- (not b/c the similarities don't exist)
361
if you don't have same amount of data for each sequence
use p-distances (proportions)
362
problems with number of differences/p-distances
unobserved mutations
A--G--T = 2 substitutions but only 1 is observed
A--G--A = 2 substitutions, none are observed
what we see is not always the whole story
363
attempt to correct for unobserved mutations
Jukes and Cantor formula
364
Jukes and Cantor
distantly related sequences likely to have experienced more substitutions than visible
mutations are more likely to occur at the same place given a large number of mutations
365
without Jukes and Cantor correction p-distance over time graph
asymptotes
366
Jukes and Cantor formula
gets rid of asymptote
distance = -(3/4)ln(1-(4/3)p)
p = proportion of changed sites, p-distance
367
if p= 0.18, JC distance =
0.206
368
influence of JC formula
distance btw similar sequences increases very little
distance btw distant sequences increases more
increases branch lengths in distant sequences
369
Actinopterygii
almost all fish
370
Sarcopterygii
lungfish, coelocanth
forgs, reptiles, mammals,..
371
relationship btw lungfish, lobe-fin, tetrapod, ray-fin
LF and Coelocanth (lobe fin) sister taxa
LF, C and T sister taxa
LF, C, T and RFF sister taxa
372
exons and introns
exons encode a.a.'s, introns dont
changes in interns less likely to have effect on proteins
we compare exons- more likely to be similar
373
exon vs. intron graph
more distance between families than within
more distance in introns (x-axis) then exons (y-axis)
374
red pandas
sister group of raccoons, badger, otter
more closely related to seals and sea lions than other bears
375
giants panda
at the base of the bear tree
other bears are more closely related to one another than to giant pandas
376
heyenas
are much more closely related to cats than to dogs
377
OLD convergence and parallelism
C- evolution of similar traits in unrelated taxa
P- evolution of similar traits in closely related taxa
378
NEW convergence and parallelism
C- evolution of similar traits in different lineages, involving different developmental pathways
P- evolution of similar traits in related lineages, involving same developmental pathways
379
new convergence and parallelism definitions based on
closeness of relatedness and molecular pathway
but don't really need to say anything about relationship
C- results from different genetic mechanisms
380
toothlessness evolution
same genes are involved- parallelism
if we look at the relationships it would be convergence
381
evolution of body elongation
in sister taxa- same mechanism (increasing size of vertebrae) - parallelism
in more distantly related species- different mechanism (increasing # of vertebrae) - convergence
382
Pattern
the "what"?
ex. phylogeny
simple description of what we see
383
Process
the "how"?
genetic development/mechanism
process by which pattern arose
ex. giraffes have long necks b/c they expand size of vertebrae
and the "why"?
adaptive/selective scenario
assumed to be for feeding at higher levels
384
The integrated organism
diverse physiological processes all affect each other (ability, rate) one imposes trade-off on another
ex. water absorption through skin affects gas loss through skin, heat exchange, etc.
385
fluxes of mass
respiratory gases, organic and inorganic substances, water
386
fluxes of energy
heat, charge, potential energy
387
physiologic trade-off in Darwin's finches
negative trade-off btw velocity and bite force- 2 separate muscles, can't maximize both at the same time
388
endothermy trade-off
can live in cold parts of the world
must eat all the time to conserve thermoregulation processes
389
interacting organ systems of vertebrates
skin, skeleto-muscular, nervous, endocrine, respiratory, digestive, excretory, circulatory
all interacting
390
major transitions in evolutionary history of vertebrates
water to land (origin of tetrapods)
land to water (origin of whales)
origin of amniotes
ectothermy-endothermy
evolution of flight (origin of birds)
origin of mammals
origin of turtles
origin of snakes
all involve multiple systems at once
391
adaptations in the transition between land and water
density/viscosity changes - locomotion and support
O2/CO2 content - gas exchange
thermal capacity - thermoregulation
refractive index - visual system
velocity of sound - hearing apparatus'
392
evolution of endothermy
insulation - controlling heat loss/gain
circulatory system
respiratory system (endo. high O2 demand)
metabolic rate
musculoskeletal system
393
evolution of birds
endothermy
flight
wings
pneumatic bones (air filled bones)
feathers
rhamphotheca (bill)
394
evolution of snakes
elongation of body
reduction/loss of limbs
repackaging of internal anatomy (long and skinny)
modified locomotion
modified feeding apparatus
395
correlated progression assumptions
incremental changes
natural selection
systems evolve in parallel
--traits can only change by one unit at a time, can be no more than one unit different than connecting traits, no obvious starting point
396
incremental changes in correlated progression
characters are highly integrated with each other, no one can evolve by more than a small amount at a time, without losing functional integration within organism as a whole
ex. can't just grow a leg and become terrestrial
397
natural selection in correlated progression
natural selection tests the fitness of an organism as a whole, not any individual characteristics
ex. fitness of leg is not the issue, fitness of the organism is
398
systems evolve in parallel, correlated progression
all structures and functions evolve by respective sequences of small steps in loose correlation with each other to maintain continuous functional integration
399
your inner fish
all the parts mammals have are modified parts from fish ancestors
no parts are new parts, history matters
400
co-option
new use for old parts
401
gill arches
co-opted for other functions
arches 1-4 --- jaws, ears, larynx, throat
bones, muscles, nerves, arteries develop inside these gill arches
402
every bone in our head can be traced to
plates, blocks, and rods (from fish)
plates- dermatocranium
blocks- chondrocranium
rods- splanchonocranium (jaws)
403
distinguishing feature of mammals (bone)
lower jaw only one bone- dentary
404
macroevolution
changes above or at a species level (speciation)
organisms are highly integrated
constrained by ancestral history- if we could build a new one it would be much different
405
linking micro and macroevolution
phenotypic plasticity- variation within individual species raw material for evolution of new species
developmental biology (evo-devo)
406
graph of phenotypic plasticity in reaction norm
phenotype vs. environment is a linear graph where phenotype is able to vary with different environments
407
genetic assimilation
occupying on enviro.---unexpressed capacity for plasticity-- enviro. changes-- rxn norm allows population to persist-- novel phenotype, no initial genetic change-- phenotype becomes fixed-- rxn norm loses plasticity
408
why reaction norm may lose plasticity
drift, cost associated w/ maintaining plasticity when it is not favoured by natural selection- old environment no longer favoured
409
macroevolutionary phenomena
origin of new structures
discontinuities in "adaptive space"
rates of evolution
causes and natures of radiation
causes and natures of mass extinctions
410
therapsids
mammal like reptiles
411
cause and nature of radiation example, therapsids
different lineages, showing increasingly mammalian characteristics, parallel changes in lineages
412
phylogenetic baggage
we are constrained by our evolution
413
exaptation
and the related term co-option describe a shift in function of a traits evolution
ex. trait can evolve because it served 1 function, but comes to serve another
414
preadaptation
large change in function is accomplished with little change in structure
415
mammals and vertebrae
7 cervical vertebrae almost always in mammals
exception- manatees have 6/7; sloths have recruited thoracic vertebrae to lengthen neck
416
why are mammals constricted to 7 cervical vertebrae?
morphological integration with other body parts, evo-devo- environmental biology evolved along with neck
each vertebra is restricted by a somite that produces myoblasts
C3-C5 migrated further back in body, responsible for formation of diaphragm
C6-C7 associated with muscles in shoulder area
417
largest known mammal necks
increase size of vertebrae not number (giraffe)
418
Annolis inovations
lizard- 'pave the way' for future evolutions
feet w/ hooks for climbing vertical surfaces- success of the species
key innovations can only really be determined to be key in hind sight
419
adaptation
feature that performs a specific function, arises as a result of selection
420
exaptation
one function taking over old function
new use will arise for a structure
421
in trade-offs be aware of
other variables that affect trade-offs
do simple correlation value and semipartial correlation values change
422
scaling relationships
how different characteristics are related to body size
423
allometry
statistical shape analysis in biology for differential growth rates of the parts of a living organism's body
morphometrics, multivariate stats, homologous landmarks
424
morphometrics
quantitative study of shape, heavily statistical
425
landmarks
changes in position give change in shape
knowing distances between points to determine shape and convert it to data
426
example of landmarks as a beneficial study
determining landmarks in endangered species to determine sex (M/F) for rearing eggs
427
PCA
principle component analysis
428
using landmarks in salamanders
2 salamanders with overlapping ranges, occurring in allopatry and sympatry
IDing landmarks on heads to determine how they differ in allopatry and sympatry
429
isometric vs. allometric growth
Iso- no change in shape proportions of body parts- stay the same
Allo- body proportions change
430
ontogenetic change in shape
change in shape during growth, through life
431
cohort
born at the same time
432
longitudinal study
following a cohort, following a group through life
433
cross-sectional study
slice in time, harder to determine age
434
measuring and testing allometry, bivariate case H_o
H_o : isometry, b = 1
435
testing bivariate allometry, equation
Y = aX^b
ln(Y) = ln(a) + b*ln(X)
436
testing for area
H_o: b = 2
437
testing for volume or mass
H_o: b = 3
438
results of Natrix natrix allometry regression
M have longer tails in absolute measurements, not in relative = isometry (M are smaller than F)
2 sexes are different but that is established right at birth, no change during life time- isometric
longer for copulation, reproductive organs
439
growth rate of head in humans
negatively allometric to body size in early life before head reaches full size
brain growth declines with age, relatively speaking
440
growth rate of heart in humans
about isometric to body size
441
proportionate difference in skull shapes of dogs and cats
dogs undergo greater shape changes than do cats
greater diversity of skull shapes in different breeds of dogs
dogs are tremendously allometric in skull growth
cats are close to isometric in skull growth
442
chameleon characteristics
highly derived- lots of apomorphves
zygodactyly (2 fingers opposing other 2)
highly projectile tongue
independently moving eyes
prehensile tail (to anchor)
443
slope of log(mass) vs. log(length)
mass = volumetric = 3 if isometric
< 3 is negatively allometric
>3 is positively allometric
444
allometric tail growth in chameleons
negatively allometric at posterior end, positively allometric at anterior end- prehensile tail- able to curl tightly at end because vertebrae are very small
445
adaptive significance of allometry in hatchling sea turtles
highly vulnerable to predation, swallowed whole by dolphinfish (mahimahi)- grow wider shell (positive allometry)
after week 5, probability of lethal encounter decreases quicker with wider shells
turtles- caretta caretta, chelonia mydas, lepidochelys kempii
446
intraspecific
same species
447
interspecific
different species
448
phylogenetic studies require
interspecific comparisons, representative measurements from different species
449
most important evolutionary force of overall color
camoflauge
450
patches of color
intraspecific signalling
451
types of concealment
crypsis, disruptive coloration/obliterative shading, irregular marks to break up bodies outline, lighter ventral surface
452
spotted species are generally found
arboreal (forests)
453
striped species are generally found
grasslands (blend in to blades of grass)
454
aposematism
prey advertise noxiousness/pugnacity
ex. skunk
455
discontinuous color variation
white or black
albinism- likely no adaptive significance, rare
melanism- common in tropical forests (black panther)
456
sexual dichromatism
rare in most mammals, common in primates
457
physiological coloration
reflet/absorb sunlight for thermoregulation, enhance/reduce evaporation, reduce glare (dark eyes)
458
thermoregulation coloration
pale species found in deserts- reflect sunlight
dark species found in tropics- enhance H2O evaporation, protect against UV
459
If plasticity was acting on lizard field experiment
would expect immediate difference in hindlimb length
not what was seen
460
problems inferring behaviour of extinct species
underlying assumption have to be made
have to rely on modern analogues when there may not be
behaviours exhibited were documented in crown groups, little know of stem taxa
461
wing skeleton and migration
migratory, semi-migratory taxa have proportionally longer wing skeleton than non migratory taxa
462
large crushing force with bill
exceeds force needed for herbivory-- likely predation, scavenging
463
fossilized footprints
many found oriented in same direction along shoreline-- moving in parallel, gregarious behaviour
464
modern avian behaviours exhibited by ancestor of crown birds
vegetative nests, extensive pre and post-hatching parental care, egg brooding
465
loss of hearing represents
regressive evolution
466
reduction of hair cell density indicates
involvement in high-freq hearing loss
fewer hair cells = fewer sites for signal transduction
467
how selection against production of unfit hybrids can lead to differentiation
ex. female choice selecting for a divergence in male secondary sexual traits that facilitates species recognition
468
reinforcement hypothesis
the ultimate explanation for a sympatric character divergence is that it reduces the probability of hybridization
469