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The Geography of Phytochemical Races

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250 6 Oceanic Islands<br />

levels <strong>of</strong> species endemism (for such a limited land area) anywhere in the world.<br />

According to Wagner et al. (1990, 1999), writing in the Manual <strong>of</strong> the Flowering<br />

Plants <strong>of</strong> Hawaii, there are 32 endemic genera (31 dicot, 1 monocot), accounting<br />

for 15% <strong>of</strong> island genera, 850 endemic species (89%), and 1094 endemic taxa overall<br />

(91%). From a slightly different perspective, there is 94% endemism in dicot<br />

taxa and 73% in monocot taxa. <strong>The</strong> fl ora consists <strong>of</strong> 146 families (none endemic),<br />

649 genera, and 1817 species. Fosberg (1948) estimated that the affi nities <strong>of</strong> the<br />

Hawaiian fl ora were 40.1% Indo-Pacifi c, 16.5% Austral, 18.3% American, 12.5%<br />

Pantropical, 2.6% Boreal, and 10.3% obscure.<br />

Three major contributions dealing with the Hawaiian fl ora have appeared in<br />

recent years to which the interested reader is referred for detailed descriptions <strong>of</strong> the<br />

fl ora and discussions <strong>of</strong> evolutionary relationships. <strong>The</strong> two volume fl ora by Wagner<br />

et al. (1990, 1999) was referred to above. Volume 1 includes a long introduction that<br />

deals with many aspects <strong>of</strong> the islands, including geology, climate, and vegetation.<br />

Hawaiian Biogeography: Evolution on a Hot Spot Archipelago, edited by Wagner<br />

and Funk (1995), provides detailed summaries <strong>of</strong> relationships and evolutionary<br />

history <strong>of</strong> several <strong>of</strong> the major groups <strong>of</strong> vascular plants on the islands. This volume<br />

also features an up-to-date treatment <strong>of</strong> the islands’ geology. <strong>The</strong> third is a book<br />

dealing specifi cally with the silversword alliance (Argyroxiphium, Dubautia, and<br />

Wilkesia) edited by Carlquist et al. (2003).<br />

<strong>The</strong> Hawaiian archipelago can be thought <strong>of</strong> as consisting <strong>of</strong> two groups <strong>of</strong><br />

islands, the high islands that comprise the “tourist group,” and a series <strong>of</strong> lesser<br />

islands extending roughly northwestward from Kauai. <strong>The</strong> islands have resulted<br />

from the movement <strong>of</strong> the Pacifi c Plate over a “hot spot” located at approximately<br />

19°N, 155° 30′W (Clague and Dalrymple, 1987). <strong>The</strong> main group <strong>of</strong> islands<br />

(maximum estimated ages in parentheses in millions <strong>of</strong> years) consists <strong>of</strong> Hawaii<br />

Island (0.43); Maui (1.32); Kahoolawe (1.03); Lanai (1.28); Molokai (1.9); Oahu<br />

(3.7); Niihau (4.89); and Kauai (5.1) (Carson and Clague, 1995). <strong>The</strong> complex consisting<br />

<strong>of</strong> Maui, Kahoolawe, Lanai, and Molokai is frequently referred to as “Maui<br />

Nui,” or big Maui, since these four formed a single landmass during Pleistocene<br />

glaciation. Hawaii Island (known locally as the Big Island) is actively volcanic;<br />

volcanoes on the other islands range from dormant to extinct awaii. Beyond Kauai,<br />

the oldest <strong>of</strong> the extant tall islands, lies a chain <strong>of</strong> degraded volcanic islands (e.g.,<br />

Nihoa, Necker, Gardner Pinnacles, Laysan, and Lisianski) and reefs stretching<br />

over 3000 km to Midway Island (28°12′N, 177°24′W). Beyond Midway lies<br />

the Emperor Chain, a series <strong>of</strong> seamounts that extends some 2400 km in a more<br />

northerly direction from the Daikakuji Seamount to Meiji Seamount (Carson and<br />

Clague, 1995).<br />

A closer look at the age <strong>of</strong> the Hawaiian Islands is useful. In attempting to estimate<br />

the time <strong>of</strong> arrival <strong>of</strong> propagules on the islands, it is important to realize that there<br />

have been receptive high islands for a much longer time than is indicated by the age<br />

<strong>of</strong> Kauai (ca. 5.1 million). As Clague and Dalrymple (1987) point out, the Leeward<br />

Islands range from about 7 to 28 million years old. Attempts to address the age <strong>of</strong><br />

arrival <strong>of</strong> a propagule and rate <strong>of</strong> subsequent genetic divergence <strong>of</strong> new species

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