lecture 21
biological species concept
species are groups of potentially interbreeding populations that are reproductively isolated from other such groups (work pretty well for sexually reproducing animals)
phylogenetic species concept
smallest possible group descending from a common ancestor and recognizable by unique, derived traits (useful for systematics; focuses on phylogenetic history)
general lineage species concept
species are metapopulations that exchange alleles frequently enough to compreise the same gene pool
metapopulation
group of spatially separated populations of the same species that interact at some level
biologist generally agree that
no single definition of a "species" will fit all taxa
speciation
evolutionary process by which new species arise (key step in the evolution of new species is the splitting in two of an original population)
isolating barrier
an aspect of the environment, genetics, behavior, physiology, or ecology of a species that reduces or impedes gene flow from individuals of other species (geographic or reproductive)
geographic
extrinsic properties of landscape that prevent gene flow (allopatry)
reproductive
features of organisms that prevent interbreeding (barriers effective even in sympatry)
prezygotic barriers
pre-mating:
-behavioral isolation
-ecological isolation: habitat, temporal, pollinator
-mechanical isolation
prezygotic barriers
post-mating:
-copulatory behavioral isolation
-gametic isolation
prezygotic barriers
intrinsic:
-hybrid inviability
-hybrid sterility: physiological and behavioral
prezygotic barriers
extrinsic:
-ecological inviability
-behavioral sterility
elk and red deer
two populations are geographically isolated but will reproduce when brought together
total population of free-ranging elk in trans-pecos as estimated in 1995 at
330 but 2014 was 3.5k
premating barriers
timing of reproduction
corals reproduce by releasing gametes into the water, the graph shows
two species of Montastraea corals remain reproductively isolated due to spawning times
some animals have
distinct courtship rituals
some rituals reinforce recent speciation events for
closely related animals
females only respond to
specific courtship displays for mating
premating barriers
pollinator isolation (two species of partially sympatric monkey flowers are pollinated, not hybridized in nature as pollinators aren't interested in the other species)
copulatory behavioral isolation
mating between species results in damage to the female reproductive tract; different species males' copulatory courtship results in eggs not fertilized by deposited sperm
gametic incompability
sperm or pollen from one species fails to penetrate and fertilize the egg of another species
hybrids are produced but have
low fitness
hybrid inviability
the hybrid embryos of sheep and goats, for example, die in the early developmental stages birth
hybrid sterility
mules
ecological inviability (outbreeding depression)
outbreeding depression can lead to coadapted gene complexes being broken apart without the forging of new coadapted gene complexes to take their place
behavioral sterility
e.g., behavioral sterility of hybrid males in acoustically communicating grasshoppers
bateson-dobzhansky-muller incompability
genetic incompatibility in hybrid offspring arising from epistatic interactions at two or more loci
dobzhanksy-muller incompatibility arises if
one population evolves the A1B0 genotype and the other evolves the A0B1 hybrids if the populations were to come into contact
Example of B-D-M incompatibility (epistatic interaction)
QTL was used to identify Hms1 on chromosome 6 and Hms2 on chromosome 13. Hybrids with M. guttatus alleles for Hms1 produce viable pollen if they also inherit at least one M. guttatus allele at Hms2. if they get M. nasutus the pollen is sterile.
allopatric speciation
result of geographic isolation, after a geographic boundary separate them, the populations become increasingly divergent
reinforcement refers to
the increase of reproductive isolation between populations through selection against hybrid offspring
consequences of secondary contact
-fusion of incipient populations back into one species
-reinforcement
-stable or transient hybrid zone formation
-speciation of hybrids
reinforcement increases
prezygotic reproductive isolation
reinforcement (definition)
natural selection favors prezygotic isolation mechanisms that prevent the formation of hybrids with reduced fitness (postzygotic isolation must evolve in allopatry)
reinforcement outcomes
-extinction of one of the initial populations
-the initial populations can hybridized
-the initial populations can speciate, are determined by both initial divergence and level of fitness of the hybrids
sympatric speciation
reproductive isolation evolves without geographic isolation (requires nonrandom mating based on genetic or phenotypic factors )
parapatric speciation
geographic barrier only partially separates populations (some gene flow possible), for example, grass Agrostis tenuis, individuals are tolerant to heavy metals, a heritable trait, survive well on contaminated soil, but poorly on non-contaminated soil (the reverse occurs for intolerant populations)
isolation by distance
populations tend to breed with those in close proximity (causes variation across the range of the species)
ring species
is a connected series of neighboring populations, each of which interbreed with closely sited related populations, but for which there exist at least two "end" populations
ecological speciation
selection for different ecological traits in different niches creates reproductive barriers (can lead to pre- ad postzygotic isolation
magic traits
traits that simultaneously confer divergent local adaptation and act as reproductive barriers
a species population of pink-petal flowers becomes isolated in allopatry or parapatry. novel pollinators drive the selection of new traits (purple petals) in the isolated populations eventually leading to
speciation of the flower populations
phylogentic evidence for allopatric speciation
a land bridge from north america to south america only arose 3mya. allopatric speciation resulted.
palm trees evolved due to soil types (one liked basic while the other liked acidic), therefore
there was also a shift in flowering times
islands provide
opportunity for allopatric speciation
colonization leads to
speciation
(Laupala crickets only found in hawaii) QTL mapping suggests that
loci associated with male pulse rate also maps to female song preference
populations on the same island often differ in male songs and female preferences for the songs
polar bears evolved into a distinct species beginning
500,000 years ago
ecological speciation and interspecies gene flow between polar bears and brown bears
polar bears and brown bears
genes under strong selection in polar bears linked to ecological niche
-APOB proteins (to handle high levels of cholesterol from diet rich in fatty acids)
-heart function (long-distance swimming)
-hair pigmentation
(interbreeding has occurred since divergence)
when D=1 Drosophila spp. can
no longer interbreed, using molecular clocks, it takes hundreds of thousands of years for them to diverge into new biological species.
allopolyploidy can lead to
extremely rapid speciation
allopolyploid
an individual having two or more complete sets of chromosomes derived from different species
cryptic species
compromise two or more taxa that are grounded under a single name because they are more-or-less indistinguishable morphologically (important for detailed biodiversity assessments)
skippers in costa rica has caterpillars with different patters, and were apparantly
cryptic species
cryptic meadowlarks (eastern meadowlark, western meadowlark, and chihuahuan meadowlark) in
texas
16S rRNA genes vary by up to
3% in microbial species
is microbes are 97% similar to 16S rRNA, they are the
same species
single spoonful of soil may contain
10,000 different species of bacteria (soil has one trillion species by locey and lennon 2016)
horizontal gene transfer in
E.coli, however, core genome is only 3188 genes, but there are 90,000+ gene across the entire E.coli genome
horizontal gene transfer makes classification difficult
the mixing genes makes it difficult, if not impossible, to delineate species boundaries
horizontal gene transfer is
common
bacterial species often defined by
specific adaptations
stable ecotype model
species concept for microbes, species is long-lived population of genetically related individuals that share a stable set of adaptations for hte same ecological niche, yet scientist criticize this method
taking horizontal gene transfer into account
shapiro et al. suggests that gene acquired from other lineages can be important for the emergence of lineages with new adaptations
geographic barriers to gene flow
physically separate populations (important for all species concepts)
reproductive barriers are
intrinsic features of organisms that reduce likelihood of interbreeding
separation of populations over time will
reduce gene flow
divergent floral traits can cause
pollinator isolation and reduce interbreeding
genetic incompabilities that reduce hybrid fitness can be
important barriers to gene flow
specitation can occur in symatry when
populations rarely interbreed
identifying cryptic species is important for
measures of biodiversity
cryptic species may diverge from ancestral populations without
easily distinguishable morphologies
microbial organisms present challenges to species concepts
developed eaukaryotes