Chapter 24 Origin of Species

Speciation 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 © 2011 Pearson Education, Inc.

24.1 – reproduction isolation Biologists compare morphology, physiology, biochemistry, and DNA sequences when grouping organisms © 2011 Pearson Education, Inc.

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 Gene flow between populations holds the phenotype of a population together © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Habitat isolation: Two species encounter each other rarely, or not at all, because they occupy different habitats, even though not isolated by physical barriers © 2011 Pearson Education, Inc.

Temporal isolation: Species that breed at different times of the day, different seasons, or different years cannot mix their gametes © 2011 Pearson Education, Inc.

Behavioral isolation: Courtship rituals and other behaviors unique to a species are effective barriers © 2011 Pearson Education, Inc.

Mechanical isolation: Morphological differences can prevent successful mating © 2011 Pearson Education, Inc.

Figure 24.3f (f) Figure 24.3 Exploring: Reproductive Barriers

Gametic Isolation: Sperm of one species may not be able to fertilize eggs of another species © 2011 Pearson Education, Inc.

Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown Postzygotic barriers prevent the hybrid zygote from developing into a viable, fertile adult: Reduced hybrid viability Reduced hybrid fertility Hybrid breakdown © 2011 Pearson Education, Inc.

Reduced hybrid viability: Genes of the different parent species may interact and impair the hybrid’s development © 2011 Pearson Education, Inc.

Reduced hybrid fertility: Even if hybrids are vigorous, they may be sterile © 2011 Pearson Education, Inc.

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 © 2011 Pearson Education, Inc.

Other Definitions of Species The morphological species concept defines a species by structural features © 2011 Pearson Education, Inc.

The ecological species concept views a species in terms of its ecological niche The phylogenetic species concept defines a species as the smallest group of individuals on a phylogenetic tree- shared genetic history © 2011 Pearson Education, Inc.

Concept 24.2: Speciation can take place with or without geographic separation Speciation can occur in two ways: Allopatric speciation Sympatric speciation © 2011 Pearson Education, Inc.

Allopatric speciation. A population forms a new species while Figure 24.5 Figure 24.5 Two main modes of speciation. (a) Allopatric speciation. A population forms a new species while geographically isolated from its parent population. (b) Sympatric speciation. A subset of a population forms a new species without geographic separation.

Allopatric (“Other Country”) Speciation In allopatric speciation, gene flow is interrupted or reduced when a population is divided into geographically isolated subpopulations © 2011 Pearson Education, Inc.

A. harrisii A. leucurus Figure 24.6 Figure 24.6 Allopatric speciation of antelope squirrels on opposite rims of the Grand Canyon.

Evidence of Allopatric Speciation 15 pairs of sibling species of snapping shrimp (Alpheus) are separated by the Isthmus of Panama These species originated 9 to 13 million years ago, when the Isthmus of Panama formed and separated the Atlantic and Pacific waters © 2011 Pearson Education, Inc.

A. formosus A. nuttingi Atlantic Ocean Isthmus of Panama Pacific Ocean Figure 24.8 A. formosus A. nuttingi Atlantic Ocean Isthmus of Panama Pacific Ocean Figure 24.8 Allopatric speciation in snapping shrimp (Alpheus). A. panamensis A. millsae

Regions with many geographic barriers typically have more species than do regions with fewer barriers Reproductive isolation between populations generally increases as the distance between them increases © 2011 Pearson Education, Inc.

Sympatric (“Same Country”) Speciation In sympatric speciation, speciation takes place in geographically overlapping populations Polyploidy Habitat differentiation Sexual selection © 2011 Pearson Education, Inc.

Polyploidy Polyploidy is the presence of extra sets of chromosomes due to accidents during cell division more common in plants than in animals autopolyploid is an individual with more than two chromosome sets, derived from one species Allopolyploid – a hybrid that reproduces asexually or may later become fertile and reproduce with other allopolypoids. © 2011 Pearson Education, Inc.

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

Many important crops (oats, cotton, potatoes, tobacco, and wheat) are polyploids © 2011 Pearson Education, Inc.

Habitat Differentiation appearance of new ecological niches For example, the North American maggot fly can live on native hawthorn trees as well as more recently introduced apple trees © 2011 Pearson Education, Inc.

Sexual selection can drive sympatric speciation Sexual selection for mates of different colors has likely contributed to speciation in cichlid fish in Lake Victoria © 2011 Pearson Education, Inc.

Allopatric and Sympatric Speciation: A Review In allopatric speciation, geographic isolation restricts gene flow between populations Reproductive isolation may then arise by natural selection, genetic drift, or sexual selection in the isolated populations Even if contact is restored between populations, interbreeding is prevented © 2011 Pearson Education, Inc.

In sympatric speciation, a reproductive barrier isolates a subset of a population without geographic separation from the parent species Sympatric speciation can result from polyploidy, natural selection, or sexual selection © 2011 Pearson Education, Inc.

Concept 24.3: Hybrid zones reveal factors that cause reproductive isolation A hybrid zone is a region in which members of different species mate and produce hybrids Hybrids are the result of mating between species with incomplete reproductive barriers © 2011 Pearson Education, Inc.

B. variegata-specific allele Figure 24.13 EUROPE Fire-bellied toad range Fire-bellied toad, Bombina bombina Hybrid zone Yellow-bellied toad range 0.99 Hybrid zone 0.9 Figure 24.13 A narrow hybrid zone for Bombina toads in Europe. B. variegata-specific allele Frequency of 0.5 Yellow-bellied toad range Fire-bellied toad range Yellow-bellied toad, Bombina variegata 0.1 0.01 40 30 20 10 10 20 Distance from hybrid zone center (km)

Pundamilia pundamilia Figure 24.16 Pundamilia nyererei Pundamilia pundamilia Figure 24.16 Fusion: The breakdown of reproductive barriers. Pundamilia “turbid water,” hybrid offspring from a location with turbid water

Patterns in the Fossil Record The fossil record includes examples of species that appear-persist- disappear 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 © 2011 Pearson Education, Inc.

(a) Punctuated pattern Time (b) Gradual pattern Figure 24.17 Figure 24.17 Two models for the tempo of speciation.

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 © 2011 Pearson Education, Inc.

From Speciation to Macroevolution Macroevolution is the cumulative effect of many speciation and extinction events © 2011 Pearson Education, Inc.