The Origin of Species

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LECTURE PRESENTATIONS For CAMPBELL BIOLOGY, NINTH EDITION Jane B. Reece, Lisa A. Urry, Michael L. Cain, Steven A. Wasserman, Peter V. Minorsky, Robert B. Jackson Chapter 24 The Origin of Species Lectures by Erin Barley Kathleen Fitzpatrick

Overview: That Mystery of Mysteries In the Galápagos Islands Darwin discovered plants and animals found nowhere else on Earth

Speciation, the origin of new species, is at the focal point of evolutionary theory Evolutionary theory must explain how new species originate and how populations evolve Microevolution consists of changes in allele frequency in a population over time Macroevolution refers to broad patterns of evolutionary change above the species level

Animation: Macroevolution Right-click slide / select Play

Concept 24.1: The biological species concept emphasizes reproductive isolation Species is a Latin word meaning kind or appearance Biologists compare morphology, physiology, biochemistry, and DNA sequences when grouping organisms

The Biological Species Concept The biological species concept states that a species is a group of populations whose members have the potential to interbreed in nature and produce viable, fertile offspring; they do not breed successfully with other populations

Figure 24.2 (a) Similarity between different species (b) Diversity within a species

Reproductive Isolation Reproductive isolation is the existence of biological factors (barriers) that impede two species from producing viable, fertile offspring Hybrids are the offspring of crosses between different species Reproductive isolation can be classified by whether factors act before or after fertilization

Figure 24.3_a Prezygotic barriers Postzygotic barriers Habitat Isolation Temporal Isolation Behavioral Isolation Mechanical Isolation Gametic Isolation Reduced Hybrid Viability Reduced Hybrid Fertility Hybrid Breakdown Individuals of different species MATING ATTEMPT FERTILIZATION VIABLE, FERTILE OFFSPRING (a) (c) (e) (f) (g) (h) (i) (l) (d) (j) (b) (k)

Prezygotic barriers block fertilization from occurring by: Impeding different species from attempting to mate Preventing the successful completion of mating Hindering fertilization if mating is successful

Habitat isolation: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers

Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes Eastern Spotted Skunk mating March April 7-14 day implantation Western Spotted Skunk delayed implantation mating in Sept Oct fertilized eggs free float until March April

Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers

Mechanical isolation: Morphological differences can prevent successful mating ex: bushbaby

Mechanical isolation: Morphological differences can prevent successful mating plants, too. black sage white sage

Gametic Isolation: Sperm of one species may not be able to fertilize eggs of another species important in broadcast spawners isolation at the molecular level

Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult: Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown

Reduced hybrid viability: Genes of the different parent species may interact and impair the hybrid s development

Reduced hybrid fertility: Even if hybrids are vigorous, they may be sterile

Hybrid breakdown: Some first-generation hybrids are fertile, but when they mate with another species or with either parent species, offspring of the next generation are feeble or sterile

Limitations of the Biological Species Concept The biological species concept cannot be applied to fossils or asexual organisms (including all prokaryotes) The biological species concept emphasizes absence of gene flow However, gene flow can occur between distinct species For example, grizzly bears and polar bears can mate to produce grolar bears

Figure 24.4 Grizzly bear (U. arctos) Polar bear (U. maritimus) Hybrid grolar bear

Other Definitions of Species Other species concepts emphasize the unity within a species rather than the separateness of different species The morphological species concept defines a species by structural features It applies to sexual and asexual species but relies on subjective criteria

The ecological species concept views a species in terms of its ecological niche It applies to sexual and asexual species and emphasizes the role of disruptive selection Scrub Jay insects, fruit in summer nuts in winter Pinyon Jay social primarily pine nuts

Concept 24.2: Speciation can take place with or without geographic separation Speciation can occur in two ways: Allopatric speciation Sympatric speciation

Allopatric ( Other Country ) Speciation In allopatric speciation, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations

The Process of Allopatric Speciation The definition of barrier depends on the ability of a population to disperse For example, a canyon may create a barrier for small rodents, but not birds, coyotes, or pollen Harris Antelope Squirrel White Tailed Antelope Squirrel

Figure 24.8 A. formosus A. nuttingi Atlantic Ocean Isthmus of Panama Pacific Ocean A. panamensis A. millsae

Regions with many geographic barriers typically have more species than do regions with fewer barriers Geographic isolation is followed by reproductive isolation. Reproductive isolation between populations generally increases as the distance between them increases For example, reproductive isolation increases between salamanders that live further apart

Figure 24.10 EXPERIMENT Some flies raised on starch medium Initial population of fruit flies (Drosophila pseudoobscura) Mating experiments after 40 generations Some flies raised on maltose medium RESULTS Starch Female Maltose Female Starch Starch population 1 population 2 Male Maltose Starch 22 9 8 20 Male Starch population 1 Starch population 2 18 15 12 15 Number of matings in experimental group Number of matings in control group

Figure 24.10a EXPERIMENT Some flies raised on starch medium Initial population of fruit flies (Drosophila pseudoobscura) Mating experiments after 40 generations Some flies raised on maltose medium

Figure 24.10b RESULTS Female Female Starch Maltose Starch population 1 Starch population 2 Male Maltose Starch 22 9 8 20 Male Starch population 1 Starch population 2 18 15 12 15 Number of matings in experimental group Number of matings in control group

Increased separation (time and/or distance) increases reproductive isolation and results in increasingly distinct speciation.

In sympatric speciation, speciation takes place in geographically overlapping populations

Polyploidy Polyploidy is the presence of extra sets of chromosomes due to accidents during cell division Polyploidy is much more common in plants than in animals An autopolyploid is an individual with more than two chromosome sets, derived from one species

An allopolyploid is a species with multiple sets of chromosomes derived from different species

Figure 24.11-4 Species A 2n = 6 Normal gamete n = 3 Species B 2n = 4 Meiotic error; chromosome number not reduced from 2n to n Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Normal gamete n = 3 Unreduced gamete with 7 chromosomes New species: viable fertile hybrid (allopolyploid) 2n = 10

Speciation can occur rapidly or slowly and can result from changes in few or many genes Many questions remain concerning how long it takes for new species to form, or how many genes need to differ between species

Patterns in the Fossil Record The fossil record includes examples of species that appear suddenly, persist essentially unchanged for some time, and then apparently disappear Niles Eldredge and Stephen Jay Gould coined the term punctuated equilibria to describe periods of apparent stasis punctuated by sudden change The punctuated equilibrium model contrasts with a model of gradual change in a species existence

The interval between speciation events can range from 4,000 years (some cichlids) to 40 million years (some beetles), with an average of 6.5 million years