EvoluTion And dEvEloPmEnT 373
body. Heterotopic differences among species are very common in plants (FIGURE
15.4A,B). In vertebrates, many phylogenetically new bones have arisen in tendons
or other connective tissues subject to stress. Many dinosaurs had bony tendons in
the tail, and the giant panda (Ailuropoda melanoleuca) has a novel “thumb” that is
not a true jointed digit (see Fig ure 20.9). Bony elements are conspicuous in the skin
of armadillos and crocodiles (FIGURE 15.4C,D).
The bodies of many organisms consist of modules—distinct units that have
distinct genetic specifications, developmental patterns, locations, and interactions
with other modules [57]. Some such modules are repeated at various sites on the
body and are termed serially homologous. In some cases, serially homologous fea-
tures lack distinct individual identities (e.g., leaves of many plants, teeth of most
amphibians and reptiles) and may be considered representatives of a single charac-
ter. An important evolutionary phenomenon is the acquisition of distinct identities
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(A) (B)
FIGURE 15.3 Neoteny in salamanders.
(A) The tiger salamander (Ambystoma tigri-
num), like most salamanders, undergoes
metamorphosis from an aquatic larva (left;
note the gills) to a terrestrial adult (right).
(B) The adult axolotl (Ambystoma mexica-
num), with gills and tail fin, resembles the
larva of its terrestrial relative. The axolotl
remains aquatic throughout its life.
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(B)
(C)
(D)
(A)
Stem
Roots
FIGURE 15.4 Examples of heterotopy. (A) The vine Monstera deliciosa
has evolved exposed roots that grow from an aerial stem. (B) Entire
plants develop on the leaf margins of Bryophyllum; they eventually drop
off and take root. (C, D) Platelets of bone (osteoderms) develop in the
skin of diverse vertebrates, such as armadillos (C) and crocodiles (D).
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