(Compositae) Endemic to the Galapagos Islands!
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Pacific Science (1995), vol. 49, no. 1: 17-30 © 1995 by University of Hawai'i Press. All rights reserved Phytogeography and Ecology of Scalesia (Compositae) Endemic to the Galapagos Islands! 2 SYUZO ITow ABSTRACT: Scalesia (Compositae), a genus endemic to the Galapagos Is lands, consists of 12 shrubby species distributed in the lowland dry zone and three tree species found in the mid-elevation moist zone. They are completely allopatric in distribution. All the species have herbaceous traits: fast growth, soft wood, large pith at the center of trunk, and flowering within 1 yr after germination (in greenhouse). The tree species Scalesia pedunculataHook. f. is shade-intolerant and heliophilous, and predominates as a monoculture in the moist zone of the four larger high-elevation islands. In ecological succession, it functions as pioneer, successor, and climax canopy plant. Even at climax or maturity of this monodominant forest, the canopy is not accompanied by young generations beneath owing to its shade-intolerance. The canopy popula tion of postmature forest dies back nearly synchronously. A new generation then develops to build new forest. The progression from germination to ma turity, and further to senescence and die back, is a self-cyclic succession, with out change of dominant species. Over much of its range, S. pedunculata is en dangered by the effects of past agricultural exploitation or heavy browsing by free-ranging goats, pigs, and donkeys; however, the population on the north side of Isla Santa Cruz has been preserved in good condition in the Galapagos National Park. THE GALAPAGOS ISLANDS are located on the a mew species. Eliasson (1974, 1975) pub equator in the eastern Pacific, about 1000 km lished a comprehensive treatment of the west of the South American coast. The ar genus, and recent nomenclature follows his chipelago is well known as a showcase of or systematic arrangement, though Cronquist ganic evolution. The flora shows a high en (in Wiggins and Porter 1971) treated the demism rate of 51 % in flowering plants same taxa in a different way. Recently, Ha (porter 1979). It contains seven endemic mann and Wium-Andersen (1986) described genera, of which Scalesia (Compositae) is a new species. one of the most important taxa, with 15 spe My experience with and concern about the cies and two subspecies, the largest number genus began in 1964, when I joined the In of species among the endemic genera. All ternational Galapagos Expedition, organized species are woody; 12 are shrubs, 1-2 m high, jointly by the University ofCalifornia, Berke distributed in dry woodlands, and three are ley, and the California Academy of Scien trees, 3 to > 10m high, found in the mid ces. Since then, I visited the islands in 1970, elevation moist zone. The Scalesia species are 1978, 1981, 1986, 1987, and 1991. My major important members of the Galapagos vege concern was the vegetation ecology of the tation. archipelago, in particular the distribution Taxonomically, Stewart (1911) treated the and ecology of Scalesia species in the field. 17 species and subspecies in his monograph Parts of my studies on the vegetation and of the Galapagos flora. Howell (1941) added Scalesia have appeared previously (Itow 1965,1971, 1988, 1990, 1992, Itow and Weber 1974, Itowand Mueller-Dombois 1988). 1 Manuscript accepted 27 April 1994. 2 Plant Ecology Laboratory, Faculty of Liberal Arts, In this paper, I focus on the distribution Nagasaki University, Nagasaki 852, Japan. and ecology of Scalesia, describing pre- 17 18 PACIFIC SCIENCE, Volume 49, January 1995 viously unpublished data and observations west coast of San Cristobal and ca. 1600 rom and citing literature to supplement my earlier at 300 m altitude, and ca. 250 rom for the articles and to draw a comprehensive picture west coast of Floreana and ca. 800 rom at of the genus. Taxonomic treatment follows 300 m altitude. Judging from those records Eliasson (1974) and Hamann and Wium and vegetation zones of those islands, rainfall Andersen (1986). pattern must be similar to that on Santa Cruz. The rainfall pattern must be similar for the southern side of Volcan Azul and Volcan Sierra Negra of Isabela because of the sim ENvmONMENTALSETTING ilarity of their vegetation zones. Physical Environment The southeast trade winds unload mois ture on the southerly located high-elevation The Galapagos Islands are of volcanic islands, especially on their windward sides. origin, created over the Galapagos Hot Spot Therefore, northerly located leeward islands in the Nazca Plate (Hey 1977). Because the and volcanoes receive lesser amounts of rain plate moves toward the southeast, easterly and are drier than the southerly located is located islands are volcanologically older lands. Low-elevation islands are always dry, than westerly located ones, and the oldest regardless of their location, because the trade lava is 3-5 million years old (Bailey 1976, winds are intercepted only slightly by the low Cox 1983). The highest peak is Volcan Wolf, topography. the northwesternmost volcano of the archi pelago, 1707 m above sea level. Volcanic Vegetation Zonation ejecta in the archipelago are basaltic lava, scoria, pumice, and ash, either old or new, There are basically four vegetation zones. and these are the principal substrates that They range from the dry zone, through the support plant life. transition and moist zones, to the highland The rainfall pattern is another important zone (Stewart 1911, 1915, Bowman 1961, factor affecting the plant life. Because the ar How 1965, 1971, 1990, 1992, Wiggins and chipelago lies in the southeast trade wind Porter 1971, How and Weber 1974, van der zone, the climate and weather are strongly Werff 1978, 1979, 1980, Hamann 1979, influenced by the moisture-bearing trade 1981). The dry zone on an island is much winds and the topography of the islands. In wider on the leeward north side (Fosberg general, southeasterly located islands, partic 1967, How 1971, 1992) and on northerly ularly their windward sides, receive much located islands than the other zones (Stewart more rainfall than northwesterly located is 1911, 1915), reflecting the above-mentioned lands and their leeward sides. A good wind rainfall pattern. ward/leeward example can be seen on Isla In the dry zone, Bursera graveolens Santa Cruz, which is a centrally located is (HBK.) Trian. & Planch. is most prevalent land rising to an elevation of 864 m above over the archipelago. There are many white sea level. The rainfall on this island ranges barked trees such as Acacia species, Cordia from 372 rom at the south coast, through species, Croton scouleri Hook. f., Piscidia 1070 rom at 200 m altitude, to 1845 mm at carthagensis Jacq., Prosopis juliflora (Sw.) 620 m altitude on the southern side; low-ele DC, and Tournefortia species. Giant cacti, vation Isla Baltra, located next to the north Opuntia and Jasminocereus, are prominent coast of Santa Cruz, receives only 76 rom on some islands (Islas Santa Fe and Plaza (How 1992). [see Figure 8], the southern sides ofIsla Santa For the southerly located high-elevation Cruz, and Volcan Sierra Negra of Isabela). islands, San Cristobal (715 m above sea level) Shrubby species of Scalesia are found among and Floreana (640 m), records of annual those plants in the dry zone. Transition-zone rainfall (Charles Darwin Research Station, vegetation is truly transitional between xero unpublished data) give ca. 400 rom for the phytic and mesophytic. Major substrates in Phytogeography and Ecology of Scalesia-ITow 19 the dry and transition zones are volcanic treeless vegetation prevails. (For details on ejecta, almost unweathered because of low Santa Cruz, see How 1990 and 1992.) rainfall. In the moist zone, the ejecta are well weathered, and the soils are deep and fertile. BIOGEOGRAPHY OF Scalesia SPECIES Abundant growth of terrestrial ferns and epiphytic bryophytes and ferns indicates that There are 15 species, four subspecies, and the atmospheric moisture and precipitation two varieties in the genus Scalesia (Table 1). regimes are high enough to support lush Based on my botanical explorations in seven plant life here. Prominent in the moist zone visits to all islands and volcanoes (except re are trees of Scalesia, Psidium galapageium mote northernmost Islas Darwin and Wolf, Hook. f., and Zanthoxylumfagara (L.) Sarg., and Volcan Wolf of Isabela) and of all spe and shrubs of Acnistus ellipticus Hook. f., cies but one (80 atractyloides Amott), to Psychotria rufipes Hook. f., and Tournefortia gether with previous studies (Wiggins and rufo-sericea Hook. f. In the highland zone, Porter 1971, Elliason 1974, 1975, Hamann TABLE I DISTRIBUTION OF Scalesia SPECIES ON INDIVIDUAL ISLANDS AND VOLCANOES OF THE GALAPAGOS ISLAND AND VOLCANO' TAXA Tree taxa pedunculata x X X X cordata XX microcephala var. microcephala XXX var. cordifolia X Shrub taxa incisa X divisa X gordilloi X villosa X affinis subsp. affinis X subsp. brachyloba X subsp. gummifera X X X X X X helleri subsp. helleri X subsp. santacruziana X retrojlexa X crockeri XX aspera X X stewartii X X atractyloides var. atractyloides X var. darwinii X baurii subsp. baurii X subsp. hopkinsii XX Nwnber of taxa 4 3 6 4 2 2 2 2 2 2 1 • SC, San Cristobal; FL, Floreana; SF, Santa Fe; SZ, Santa Cruz; BL, Baltra (off NE coast of SZ); ED, Eden (off NW coast of SZ); ST, Santiago; BA, Bartolome (off E coast of ST); PZ, Pinzon; FE, Fernandina; PT, Pinta; WL, Wolf. Volcanoes of Isabela: az, Azul; ng, Sierra Negra; aI, Alcedo; dw, Darwin; wf, Wolf. 20 PACIFIC SCIENCE, Volume 49, January 1995 and Wium-Andersen 1986), all the species Figures I and 2 show the distribution of are nearly completely allopatric in distribu Scalesia species on individual islands. Islas tion, with a wide distance between their indi Santa Cruz and San Cristobal, which are vidual ranges in the archipelago (Figure 1).