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Chapter 24 The Origin of Species Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings • Fruit flies • Horizontal gene flow Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.1 The flightless cormorant (Nannopterum harrisi), one of many new species that have originated on the isolated Galápagos Islands Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.2 Two patterns of evolutionary change Anagenesis is evolution within a lineage. Cladogenesis is evolution that results in the splitting of a lineage. (a) Anagenesis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Cladogenesis Figure 24.3 The biological species concept is based on the potential to interbreed rather than on physical similarity (a) Similarity between different species. The eastern meadowlark (Sturnella magna, left) and the western meadowlark (Sturnella neglecta, right) have similar body shapes and colorations. Nevertheless, they are distinct biological species because their songs and other behaviors are different enough to prevent interbreeding should they meet in the wild. (b) Diversity within a species. As diverse as we may be in appearance, all humans belong to a single biological species (Homo sapiens), defined by our capacity to interbreed. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.4 Reproductive Barriers Prezygotic barriers impede mating or hinder fertilization if mating does occur Habitat isolation Behavioral isolation Temporal isolation Individuals of different species Mechanical isolation Mating attempt HABITAT ISOLATION TEMPORAL ISOLATION BEHAVIORAL ISOLATION (b) MECHANICAL ISOLATION (g) (d) (e) (f) (a) (c) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Gametic isolation Reduce hybrid fertility Reduce hybrid viability Hybrid breakdown Viable fertile offspring Fertilization REDUCED HYBRID VIABILITY GAMETIC ISOLATION REDUCED HYBRID FERTILITY HYBRID BREAKDOWN (k) (j) (m) (l) (h) (i) Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.5 Two main modes of speciation (a) Allopatric speciation. A (b) Sympatric speciation. A small population becomes a new species population forms a new without geographic separation. species while geographically isolated from its parent population. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.6 Allopatric speciation of antelope squirrels on opposite rims of the Grand Canyon A. harrisi Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings A. leucurus Figure 24.7 Can divergence of allopatric fruit fly populations lead to reproductive isolation? EXPERIMENT Diane Dodd, of Yale University, divided a fruit-fly population, raising some populations on a starch medium and others on a maltose medium. After many generations, natural selection resulted in divergent evolution: Populations raised on starch digested starch more efficiently, while those raised on maltose digested maltose more efficiently. Dodd then put flies from the same or different populations in mating cages and measured mating frequencies. Initial population of fruit flies (Drosphila Pseudoobscura) Some flies raised on starch medium Mating experiments after several generations Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Some flies raised on maltose medium RESULTS 22 9 8 20 Male Male Starch Maltose Female Starch Maltose Mating frequencies in experimental group CONCLUSION Different Same populations population When flies from “starch populations” were mixed with flies from “maltose populations,” the flies tended to mate with like partners. In the control group, flies taken from different populations that were adapted to the same medium were about as likely to mate with each other as with flies from their own populations. Female Different Same population populations 18 15 12 15 Mating frequencies in control group The strong preference of “starch flies” and “maltose flies” to mate with like-adapted flies, even if they were from different populations, indicates that a reproductive barrier is forming between the divergent populations of flies. The barrier is not absolute (some mating between starch flies and maltose flies did occur) but appears to be under way after several generations of divergence resulting from the separation of these allopatric populations into different environments. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.8 Sympatric speciation by autopolyploidy in plants Failure of cell division in a cell of a growing diploid plant after chromosome duplication gives rise to a tetraploid branch or other tissue. Gametes produced by flowers on this branch will be diploid. Offspring with tetraploid karyotypes may be viable and fertile—a new biological species. 2n 2n = 6 4n = 12 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4n Figure 24.9 One mechanism for allopolyploid speciation in plants Unreduced gamete with 4 chromosomes Hybrid with 7 chromosomes Species A 2n = 4 Unreduced gamete with 7 chromosomes Viable fertile hybrid (allopolyploid) Meiotic error; chromosome number not reduced from 2n to n 2n = 10 Normal gamete n=3 Species B 2n = 6 Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Normal gamete n=3 Figure 24.10 Does sexual selection in cichlids result in reproductive isolation? EXPERIMENT Researchers from the University of Leiden placed males and females of Pundamilia pundamilia and P. nyererei together in two aquarium tanks, one with natural light and one with a monochromatic orange lamp. Under normal light, the two species are noticeably different in coloration; under monochromatic orange light, the two species appear identical in color. The researchers then observed the mating choices of the fish in each tank. Monochromatic Normal light orange light P. pundamilia P. nyererei RESULTS CONCLUSION Under normal light, females of each species mated only with males of their own species. But under orange light, females of each species mated indiscriminately with males of both species. The resulting hybrids were viable and fertile. The researchers concluded that mate choice by females based on coloration is the main reproductive barrier that normally keeps the gene pools of these two species separate. Since the species can still interbreed when this prezygotic behavioral barrier is breached in the laboratory, the genetic divergence between the species is likely to be small. This suggests that speciation in nature has occurred relatively recently. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.11 Long-distance dispersal Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.12 Adaptive radiation Dubautia laxa 1.3 million years MOLOKA'I KAUA'I MAUI 5.1 million years O'AHU LANAI 3.7 million years Argyroxiphium sandwicense HAWAI'I 0.4 million years Dubautia waialealae Dubautia scabra Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Dubautia linearis Figure 24.13 Two models for the tempo of speciation Time (a) Gradualism model. Species descended from a common ancestor gradually diverge more and more in their morphology as they acquire unique adaptations. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings (b) Punctuated equilibrium model. A new species changes most as it buds from a parent species and then changes little for the rest of its existence. Figure 24.14 A range of eye complexity among molluscs Pigmented cells (photoreceptors) Pigmented cells Epithelium Nerve fibers Nerve fibers (b) Eyecup. The slit shell (a) Patch of pigmented cells. The mollusc Pleurotomaria limpet Patella has a simple has an eyecup. patch of photoreceptors. Cornea Cellular Fluid-filled cavity fluid Epithelium (lens) Optic nerve Pigmented layer (retina) (c) Pinhole camera-type eye. The Nautilus eye functions Cornea like a pinhole camera (an early type of camera lacking a lens). Lens Optic nerve Optic nerve (d) Eye with primitive lens. The marine snail Murex has a primitive lens consisting of a mass of crystal-like cells. The cornea is a transparent region of epithelium (outer skin) that protects the eye and helps focus light. Retina (e) Complex camera-type eye. The squid Loligo has a complex eye whose features (cornea, lens, and retina), though similar to those of vertebrate eyes, evolved independently. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.15 Allometric growth (a) Differential growth rates in a human. The arms and legs lengthen more during growth than the head and trunk, as can be seen in this conceptualization of an individual at different ages all rescaled to the same height. Newborn 2 5 15 Age (years) Chimpanzee fetus (b) Comparison of chimpanzee and human skull growth. The fetal skulls of humans and chimpanzees are similar in shape. Allometric growth transforms the rounded skull and vertical face of a newborn chimpanzee into the elongated skull and sloping face characteristic of adult apes. The same allometric pattern of growth occurs in humans, but with a less accelerated elongation of the jaw relative to the rest of the skull. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Human fetus Adult Chimpanzee adult Human adult Figure 24.16 Heterochrony and the evolution of salamander feet in closely related species (a) Ground-dwelling salamander. A longer time peroid for foot growth results in longer digits and less webbing. (b) Tree-dwelling salamander. Foot growth ends sooner. This evolutionary timing change accounts for the shorter digits and more extensive webbing, which help the salamander climb vertically on tree branches. Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.17 Paedomorphosis Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Figure 24.18 Hox genes and the evolution of tetrapod limbs Chicken leg bud Zebrafish fin bud Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Region of Hox gene expression Figure 24.19 Hox mutations and the origin of vertebrates Hypothetical vertebrate ancestor (invertebrate) with a single Hox cluster First Hox duplication 1 Most invertebrates have one cluster of homeotic genes (the Hox complex), shown here as colored bands on a chromosome. Hox genes direct development of major body parts. 2 A mutation (duplication) of the single Hox complex occurred about 520 million years ago and may have provided genetic material associated with the origin of the first vertebrates. 3 In an early vertebrate, the duplicate set of genes took on entirely new roles, such as directing the development of a backbone. Hypothetical early vertebrates (jawless) with two Hox clusters Second Hox duplication Vertebrates (with jaws) with four Hox clusters Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings 4 A second duplication of the Hox complex, yielding the four clusters found in most present-day vertebrates, occurred later, about 425 million years ago. This duplication, probably the result of a polyploidy event, allowed the development of even greater structural complexity, such as jaws and limbs. 5 The vertebrate Hox complex contains duplicates of many of the same genes as the single invertebrate cluster, in virtually the same linear order on chromosomes, and they direct the sequential development of the same body regions. Thus, scientists infer that the four clusters of the vertebrate Hox complex are homologous to the single cluster in invertebrates. Figure 24.20 The branched evolution of horses Recent (11,500 ya) Equus Pleistocene (1.8 mya) Hippidion and other genera Nannippus Pliohippus Hipparion Neohipparion Pliocene (5.3 mya) Sinohippus Megahippus Callippus Archaeohippus Miocene (23 mya) Merychippus Hypohippus Anchitherium Parahippus Miohippus Oligocene (33.9 mya) Mesohippus Paleotherium Epihippus Propalaeotherium Eocene (55.8 mya) Pachynolophus Orohippus Key Hyracotherium Copyright © 2005 Pearson Education, Inc. publishing as Benjamin Cummings Grazers Browsers