Chordates Flashcards
Chordate zoology is what type of biology
whole-organism biology (not just cell, function, form,.. everything)
organismal biology
research at the level of the whole organism, integrated over structure, function, ecology, and evolution
organism structure
anatomy, morphology
organism function
physiology, behaviour
evolution
phylogeny
functional morphology
focuses on the link between form (morphology) and performance
ecological morphology
focuses on the link between performance and ecology (ecomorphs, ecomorphotypes)
ecomorph
species with the same structural habitat/niche, similar in morphology and behavior, but not necessarily close phyletically
ecomorphotype
Any morphological modification caused by, or related to, specific ecological conditions
integrative biology
near synonym for organismal biology; brings different aspects of organisms and their environment together
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
natural history
scientific study of the organism in its natural surroundings
kinds of chordates
Tunicata
Cephalochordata
Vertebrata
Tunicata includes
tunicates and sea squirts
Tunicata was formerly called
Urochordata (also protochordates- not valid taxonomic name)
Cephalochordata includes
lancelets (amphioxus)
Numbers of chordate species
Tunicata - 2150
Cephalochorddata - 25
Vertebrata - 63 600
*numbers are always changing
tunicates
free-living larva, sessile adult
shared characteristics of chordates
notochord dorsal hollow nerve cord pharyngeal gill slits endostyle muscular postanal tail
notochord
incorporated in vertebral column in vertebrates
stiff, flexible rod, runs length of back, provides connections for body muscles, and support
dorsal hollow nerve cord
spinal cord with brain at anterior end in vertebrates
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
endostyle
ciliated groove before larynx
secretes mucus to trap food
homologous with thyroid gland in vertebrates
muscular postanal tail
extension of the body that runs past the anal opening
only present in embryonic stage of humans
paedomorphosis
retaining juvenile characteristics into adult hood
sister group to vertebrates
urochordata (most current theory)
vertebrates and urochordates are sister group to cephalochordates
pattern vs. process
phylogeny vs. scenario
current phylogenetic theory (pattern), process (scenario) yet to be determined
myomeres
blocks of skeletal muscle tissue found commonly in chordates. commonly zig-zag, “W” or “V”-shaped muscle fibers
possible first chordate
Pikaia
aspects of morphology (anatomy)
comparative
functional
transitional
comparative morphology
similarities/differences between groups
ex. heart
functional morphology
how organisms are equipped to deal with different situations
transitional morphology
macroevolutionary change- how we go from one form to another
why study chordates
intrinsic interest vertebrates are us diversity evolutionary record model organisms
chordate diversity
masses range from 10^-4 - 10^5 kg! (larva to blue whale)
altitudes range 58km- deep ocean - over himalayans
why evolutionary record is important for studying chordates
they have the best preservation of all organisms
model organisms
amphibians- developmental biology
birds- population biology
vertebrates characteristics
internal skeleton vertebral column with cranium at anterior spinal nerve cord with brain at anterior neural crest HOX genes
vertebrate internal skeleton
bone and/or cartilage
vertebral column
individual vertebra, skull at end
rudimentary in lampreys
lacking in hagfish
full formed in gnathostomes
gnathostomes
jawed vertebrates
neural crest
cells/tissue- formed during embryonic development and migrate to head
HOX genes
do exist in inverts but more important in vertebrates
vertebrates have the most HOX genes
“grades” of vertebrates
“fishes”
Tetrapods
paraphyletic
composed of some but not all members descending from a common ancestor
fishes
paraphyletic- would be monophyletic if we include all groups that have come from fishes (including us)
31,000 species
mainly Osteichthyes
groups in fishes
Agnathans
Chondrichthyes
Osteichthyes
Agnathans
jawless vertebrates- hagfish and lampreys, Ostracoderms
Chondrichthyes
cartilaginous fish- sharks, rays, ratfish
Osteichthyes
bony fish
Crossopterygii, Actinopterygii
Cossopterygii
Actinistia + Dipnoi (lobe-finned fish- coelacanth + lungfish)
Actinopterygii
ray-finned fish (most fish)
Tetrapods
all the rest of the vertebrates
4 feet, lots of secondary loss examples
Tetrapod examples
Amphibia
Amniota
Aves
Mammalia
Amphibia
7000 species
lissamphibians- apodans salamanders, anurans
Amniota
amniotes– Reptilia
Reptilia
10,000 species- turtles, crocodiles, tuataras, lizards, snakes, amphisbaenians
Aves
10,000 species
directly derived from reptiles, make reptiles a non-taxonomic group
Mammalia
5500 species- mammals
mostly placentals, some monotremes and marsupials
Ectotherms
“fishes”
amphibians
“reptiles”
endotherms
generate own body heat metabolically
birds
mammals
Amniotes
“reptiles”
birds
mammals
(taxonomic group)
Anamniotes
“Fishes”
Amphibians
(not a taxonomic group)
Sauropsids
“reptiles”
birds
synapsids
mammals
lampreys
Agnathan- parasitize other fish
Fossil Agnathans
Ostracoderms- coats of bone armour, very different from living fish
Gnathostomata
defined by presence of jaws with teeth
usually have limbs
thought to be closest living relative of tetrapods
lungfish (3 species)
tetrapoda
limbs with digits for terrestrial locomotion
internal nostrils
tympanic membrane and stapes
strong skeleton
internal nostrils
breathing through nose
fish do not breath through nose (except lungfish)
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
tetrapod strong skeleton
support against gravity (adaptation to terrestrial life)
Lissamphibia
3 major, distinct morphological groups
thin skin
complex life cycle
numerous departures from theme
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
Lissamphibia complex life cycle
includes metamorphosis
aquatic egg– aquatic larva– aquatic/terrestrial juvenile and adult
Amphibian groups
salamanders
anurans
apodans
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
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
unique Anuran forms
one species- female swallows eggs and they develop in her stomach
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
Amniota
amniotic egg- amniote surrounds embryo
exclusively internal fertilization
allantois
waste dump, highly vascularized (blood vessels)
develops to line membrane surrounding the egg- transport system for gas exchange
amniote shell
holds eggs together, has mating consequences- necessitates internal fertilization, must develop on land (oviparity)
oviparity
Oviparous animals are animals that lay eggs, with little or no other embryonic development within the mother
Reptilia
not a taxon group
thicker skin, covered with epidural scutes (keratin)
much more terrestrially adapted, lose water slower
scutes
scales
keratin provides mechanical protection, defense against water loss, protection of lipids
“reptile” groups
turtles tuataras lizards amphisbaenians snakes crocodilians
turtles
only vertebrate in which ribs are outside
lots of aquatic, some marine, some terrestrial
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
“lizards”
biggest group of “reptiles”
very diverse, worldwide, terrestrial, arboreal
lots of long slender bodies w/o limbs
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
head types of Amphisbaenians
round head- soft sediment
wedge shaped head- hard substrate
snakes
mostly have no trace of limbs
many arboreal- long and slender to spread weight out over branches
rain snake
oval cross section
tail flattened to propel through water
Crocodilians
~22 species living forms all fairly similar body covered with osteoderms long, elongate skull extant are only aquatic tropical/subtropical
osteoderms
bony deposits forming scales, plates or other structures in the dermal layers of the skin
crocodile snout specialization
long, skinny snout- fish eating- quicker swiping through water
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
bird feet/beak
tell a lot about overall life ecology
ex. flightless - marine
bird feathers
aid endothermy
major flight surface of wing
Mammalia
endothermic body covered with hair/fur (keratin, insolation) highly differentiated dentition numerous derived traits in skeleton pinna
derived traits in mammal skeleton
squamosal-dentary jaw joint
7 cervical vertebrae (neck vertebrae)- almost all mammals have 7
Pinna
the visible part of the ear that resides outside of the head
main groups of mammals
monotremes
marsupials
placentals
monotremes
lay eggs- platypus, echidna
only a few species
basically same as reptile eggs- hatch then female feeds milk via ducts (not nipples)
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
marsupial embryo pouch
marsupion
placentals
development until birth is in uterus
intimate connection between embryo and mother (nutrients, gas exchange)
important parts of evolution
genes, chromosomes, alleles, mutation (many neutral, some lethal), proteins (structural, hormones, enzymes), recombination, dominance, pleiotropy, polygenic traits, epistasis, regulatory genes
pleiotropy
single genes code for multiple parts of the body
epistasis
interactions between genes
effect of gene depends on presence/expression of other genes
regulatory genes
govern expression of genes
phenotype =
genotype * environment
phenotypic plasticity
ability of an organism to change its phenotype in response to changes in the environment
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
reaction norms
expression of the phenotype of a particular genotype in different environments
no genotype environment interaction, reaction norm
phenotype vs. environment graph
variation is constant, parallel for 2 genotypes among different environments
genotype environment interaction, reaction norm
genotypes are not parallel or constant (lines may cross)
one genotype may produce multiple phenotypes
phenotypic flexibility
reversible changes within individuals
ex. plumage variation with season- changeable/reversible phenotype
types of selection
artificial selection (natural) selection adaptation directional selection balancing selection disruptive selection
natural selection
variation, heritability, differential survival/reproduction
variation has to be heritable for evolution to occur
adaptation
a trait that has arisen from natural selection
three modes of selection
stabilizing
directional
disruptive
stabilizing selection
balancing
extreme values are a disadvantage and are ‘pruned’
keep population at certain ‘optimum’
directional selection
one extreme is disadvantageous
leeds to a shift away from one extreme
disruptive selection
intermediates are disadvantageous
selection is for the extreme- split down the ‘middle’
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
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
selection isn’t the only factor that determines change
genetic drift
founder effect
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
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
species/speciation concepts
multiple species concepts discontinuities morphospecies biological species hybridization asexual reproduction
discontinuities
breaks between species (ex. morphology)
multiple species concepts
many definitions of species
the 2 we will focus on are biological species concept (BSC), morphological species concept
BSC
depends on sexual breeding, must be able to interbreed, hard to test, depends on species exhibiting sympatry
discontinuity- reproductive isolation
sympatry
species that occur in same geographic area
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
morphospecies
different species should look different
domestic dogs- haven’t speciated
sibling species
different species that look identical
sibling species example
Hyla versicolor- tetraploid (gray tree frog)
Hyla chrysoscelis- diploid (Cope’s gray tree frog)
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.
polyphyletic
does not include the common ancestor of all members of the taxon
paraphyletic
includes the most recent common ancestor, but not all of its descendents
species is used to
describe groups that we recognize
describe what the animals themselves recognize
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
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
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
yellow-dumped warbler hybrids
Audubon’s warbler / Myrtle warbler
100km hybrid zone
graphs show different traits within/outside of hybrid zone
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
3 known categories of asexual reproduction in vertebrates
recall that BSC requires sexual breeding
hybridogenesis
gynogenesis
parthogenesis
hemiclonal
half of females genome is passed on clonally
clonal
all of females genome is passed on clonally
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
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
parthenogenesis
males not used at all, diploid, unfertilized egg (clonal)
most vertebrates are
bisexual- normal sexual reproduction
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
usually inviable offspring
R. esculenta x R. esculenta = R. ridibunda
origin of R. esculenta
R. ridibunda x R. lessonae = R.esculenta
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
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
Bulloch’s Oriole and Blatimore Oriole
two morphologically distinct species, but they are interfertile
two morphospecies, but one or two biological species?
interfertile
capable of interbreeding
cryptic species
morphologically similar, genetically distinct, incapable of interbreeding
cryptic species example
African Savanna Elephant, African Forest Elephant
speciation can occur
allopatrically, parapatrically/peripatrically, or sympatrically
allopatric speciation
geographically separate
parapatric speciation
geographically adjacent
sympatric speciation
geographically coincident (same area)
non-selective speciation
‘by accident’- random drift in small founding populations
natural selection speciation
mutation-order speciation
ecological speciation
ecological speciation
evolution of reproductive isolation between populations by divergent natural selection arising from differences between ecological environments
indirect or direct
indirect ecological speciation
by-product speciation
reproductive isolation occurs as an incidental by-product of adaptation to different environments
direct ecological speciation
selection directly favours reproductive isolation
ex. if hybrids are less viable
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)
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
lentic
standing water
lotic
running water
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
piscivory
eating of fish
durophagy
eating behavior of animals that consume hard-shelled or exoskeleton bearing organisms, such as corals, shelled mollusks, or crabs
increasing potential for gene flow in absence of differentiation
decreasing spatial scale
allopatric– parapatric– sympatric
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
Replicated (parallel) ecological speciation in lizards
habitat matching- different species show the same coloration changes to match light/dark soil habitats
punctuated equilibrium
little net evolutionary change for most of geological history, remaining in an extended state called stasis, disrupted by abrupt change
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