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Encyclopedia of Evolution.pdf - Online Reading Center

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0 island biogeography<br />

Protostomes v. deuterostomes. The bilaterally symmetrical<br />

animals diverged into two lineages that differ primarily in<br />

the manner in which their embryos develop. In protostomes,<br />

the early embryo cell divisions occur in a spiral fashion,<br />

whereas in deuterostomes they occur in a radial fashion. In<br />

protostomes, the developmental fate <strong>of</strong> each cell is determined<br />

from the beginning. In deuterostomes, the fate <strong>of</strong> the cells is<br />

not determined until about the eight-cell stage. Therefore, in<br />

deuterostomes, one fertilized egg cell can become up to eight<br />

identical <strong>of</strong>fspring. Deuterostomes, unlike protostomes, can<br />

have identical twins, triplets, etc.<br />

Protostomes: Ecdysozoans v. others. On the basis <strong>of</strong><br />

DNA evidence, the protostomes are divided into two groups.<br />

The ecdysozoans secrete an external skeleton. As they grow,<br />

they split open the skeleton and squirm out <strong>of</strong> it, secreting<br />

a new one. This process, called ecdysis, characterizes arthropods<br />

and nematodes.<br />

• Arthropods (Phylum Arthropoda) have hard external skeletons<br />

and jointed appendages. Appendages are legs or other<br />

leglike structures (such as the antennae <strong>of</strong> insects). Arthropods<br />

were among the earliest phyla <strong>of</strong> animals to have a<br />

burst <strong>of</strong> evolutionary diversification, in the Cambrian period,<br />

including the trilobites, which evolved into numerous<br />

species, until becoming extinct at the end <strong>of</strong> the Permian<br />

period (see Permian extinction). Today, the major<br />

arthropod groups are the crustaceans (including crabs, lobsters,<br />

shrimps, and terrestrial pillbugs); the centipedes and<br />

millipedes; the chelicerates (including spiders, scorpions,<br />

mites, and ticks); and the insects. In terms <strong>of</strong> sheer weight<br />

and number, as well as species diversity, arthropods rule the<br />

animal world (see biodiversity).<br />

• Nematodes (Phylum Nematoda) are the roundworms. They<br />

are incredibly abundant inhabitants <strong>of</strong> the soil, especially near<br />

plant roots and in wetlands. Experts say that if everything in<br />

the world except nematodes vanished, one would still be able<br />

to see the Earth as a shimmering sphere <strong>of</strong> nematodes.<br />

Among the other protostomes that are not ecdysozoans are:<br />

• Flatworms (Phylum Platyhelminthes) include aquatic and<br />

parasitic forms. The parasitic forms may have extremely<br />

complex life cycles.<br />

• Mollusks and brachiopods. Mollusks (Phylum Mollusca)<br />

have a mantle and guts over a single foot. Bivalve mollusks<br />

(e.g., clams) secrete two hard shells hinged together. They<br />

filter the water in which they live to obtain food particles.<br />

Cephalopod mollusks have a head (cephalo-) over a foot (pod),<br />

with a beaked mouth in the middle <strong>of</strong> the foot, as in<br />

octopus and squid. Gastropod (“stomach-foot”) mollusks<br />

include snails and slugs. Despite their superficial similarity<br />

to bivalves, the brachiopod “lampshells” are a distinct phylum,<br />

because unlike bivalves they have a specialized feeding<br />

apparatus called a lophophore. The present-day abundance<br />

<strong>of</strong> bivalves and relative scarcity <strong>of</strong> brachiopods may be a<br />

leftover effect <strong>of</strong> the Permian extinction, which nearly<br />

destroyed the brachiopods.<br />

• Annelids (Phylum Annelida) are the segmented worms,<br />

such as many <strong>of</strong> the marine worms, the earthworms, and<br />

the leeches. Because annelids and insects both have segments,<br />

they were formerly thought to share a relatively<br />

recent common ancestor, but DNA analysis has forced a<br />

reevaluation <strong>of</strong> this idea.<br />

Deuterostomes. The two phyla <strong>of</strong> deuterostomes are the<br />

echinoderms and the chordates.<br />

• Adult echinoderms (Phylum Echinodermata, which means<br />

“spiny skin”) have spiny or knobby outer layers and move<br />

along the rocks <strong>of</strong> the seashore by means <strong>of</strong> water-powered<br />

tube-feet. They include starfish and sea urchins. The larvae<br />

<strong>of</strong> echinoderms are bilaterally symmetrical, even though<br />

the adults look superficially radial.<br />

• The chordates have a rod, or notochord, along the back,<br />

which strengthens the body and the main nerve cord. Invertebrate<br />

chordates include the lancelet, which looks superficially<br />

like a headless fish. The vertebrates evolved from an ancestor<br />

that resembled a lancelet (see fishes, evolution <strong>of</strong>).<br />

Almost all <strong>of</strong> the animal phyla are represented in the<br />

Cambrian period in at least a rudimentary form. It is not<br />

clear how many <strong>of</strong> them were present before the Cambrian<br />

explosion. Some <strong>of</strong> the Ediacaran organisms from the<br />

period just before the Cambrian resemble jellyfish, while others<br />

resemble colonial cnidarians. All identifications <strong>of</strong> the<br />

Ediacarans, however, are controversial. Fossils <strong>of</strong> animal<br />

embryos are known from the Ediacaran period. DNA studies<br />

suggest that some <strong>of</strong> the earliest evolutionary divergences<br />

<strong>of</strong> animals occurred from 700 million to 1 billion years ago,<br />

although the fossil evidence for these divergences is scarce.<br />

Most invertebrate phyla are completely aquatic. However,<br />

like the vertebrate lineage within the chordate phylum,<br />

several arthropod lineages have diversified into all the habitats<br />

<strong>of</strong> dry land.<br />

Further <strong>Reading</strong><br />

Bottjer, David J. “The early evolution <strong>of</strong> animals.” Scientific American,<br />

August 2005, 42–47.<br />

Knoll, Andrew H., and Sean B. Carroll. “Early animal evolution:<br />

Emerging views from comparative biology and geology. Science<br />

284 (1999): 2,129–2,137.<br />

Museum <strong>of</strong> Paleontology, University <strong>of</strong> California, Berkeley. “Introduction<br />

to the Placozoa: The simplest <strong>of</strong> all known animals.” Available<br />

online. URL: http://www.ucmp.berkeley.edu/phyla/placozoa/placozoa.<br />

html. Accessed April 18, 2005.<br />

island biogeography See biogeography.<br />

isolating mechanisms Isolating mechanisms are biological<br />

processes that cause reproductive isolation, in which populations<br />

cannot successfully interbreed if they are in contact.<br />

If two populations could interbreed if they were in contact<br />

but do not do so because they live in different locations, the<br />

populations are geographically isolated. Geographic isolation<br />

is not an isolating mechanism. Isolating mechanisms keep<br />

populations from interbreeding even if they live in the same<br />

place. <strong>Evolution</strong>ary biologists consider two populations that

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