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Evolution__3rd_Edition

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462 PART 4 / <strong>Evolution</strong> and Diversity<br />

(a) (b)<br />

Sivapithecus<br />

Orang-utan Gorilla Chimp Ramapithecus Homo Orang-utan (incl. “Ramapithecus” ) Gorilla Chimp Homo<br />

>9–12 million<br />

years ago<br />

Figure 15.25<br />

Relations of Homo, other great apes, and Ramapithecus,<br />

according to (a) original paleontological and morphological<br />

Molecular evidence suggested a<br />

more recent origin for humans<br />

>9–12 million<br />

years ago<br />

About 5 million<br />

years ago<br />

evidence, and (b) molecular (and revised paleontological and<br />

morphological) evidence. (The dashed lines imply uncertainty<br />

in the order of the human–chimpanzee–gorilla split.)<br />

In the early 1960s, Goodman (1963) first demonstrated the molecular similarity of<br />

humans and other great apes; but the molecular argument for a recent human–ape split<br />

was most influentially made in a paper by Sarich & Wilson (1967). Sarich and Wilson<br />

used an immunological distance measure. The method is similar in philosophy to DNA<br />

hybridization, but differs in the exact molecule used.<br />

To measure immunological distance, Sarich and Wilson first made an antiserum<br />

against human albumin by injecting human albumin into rabbits (albumin is a common<br />

protein which circulates in the blood). They then measured how much that antiserum<br />

cross-reacted with the albumin of other species, such as chimps, gorillas, and<br />

gibbons. The antiserum recognizes the albumins of closely related species, because they<br />

are similar to human albumin; but it does not recognize them quite as efficiently as it<br />

does human albumin. The degree of cross-reactivity gives a measure of the immunological<br />

distance (ID) between a pair of species. ID increases among phylogenetically<br />

more distant relatives, and the relative rate test (Box 7.2, p. 166) suggests that ID<br />

increases at a constant rate through time; immunological distance is a sort of molecular<br />

clock. The clock can be calibrated using the fossil record for some of the studied species,<br />

and the ID can then be used to estimate the divergence time for other pairs of species.<br />

The results of this method suggest that Homo and the other great apes have too short<br />

an ID to fit with a pre-Ramapithecus divergence: Sarich and Wilson suggested humans<br />

and chimps diverged only about 5 million years ago. Subsequent molecular work has<br />

supported them. The DNA hybridization results that we looked at earlier suggest a similar,<br />

if perhaps slightly older, figure (see Figure 15.12), and other molecules suggest a<br />

figure of 3.75–4 million years. The corollary is that if Homo diverged from chimps and<br />

gorillas 5 million years ago, it cannot be more closely related to Ramapithecus than to<br />

the living great apes. The phylogeny must be more like Figure 15.25b.<br />

So the molecular and fossil evidence disagreed. A controversy began, in which both<br />

the molecular and morphological evidence was challenged (often by experts in the<br />

..

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