diversity, endemism, and extinction in the flora of new ...

Rare, Threatened and Endangered Floras of Asia And the Pacific Rim (C.-I Peng & P.P. Lowry II, eds.), Institute of Botany, Academia Sinica Monograph Series No. 16, pp. 181-206 (1998), Taipei

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DIVERSITY, ENDEMISM, AND EXTINCTION IN THE FLORA OF NEW CALEDONIA: A REVIEW Porter P. Lowry II1 Missouri Botanical Garden, P.O. Box 299, St. Louis, MO 63166-0299, USA. E-mail: [email protected]

ABSTRACT The southwest Pacific island of New Caledonia, covering approximately 17,000 km2, has a remarkably diverse and highly endemic flora for an area its size, with an estimated 3,137 native species of angiosperms and gymnosperms (79% endemic), representing about 763 genera (14% endemic), and 169 families (3% endemic).

Numerous relict endemic lineages occur on the island, such as 14

gymnosperm genera, with 44 species (98% endemic) representing 7% of the world’s total, and members of many primitive angiosperm groups (e.g., Winteraceae, Amborellaceae, Monimiaceae, Annonaceae, Chloranthaceae, etc.). New Caledonia separated from Australia about 65 MYBP and reached its present position around 50 MYBP; during the Paleocene and middle Eocene peridotites covered the island resulting in ultrabasic substrates that still occupy about a third of the surface. Many members of the late Cretaceous-early Tertiary Gondwanan flora survived in New Caledonia’s equable climate but were eliminated in Australia due to increasingly dry conditions. The island’s geography and climate today have resulted in a wide range of vegetation types; recent studies show that specific endemism is highest in maquis vegetation (91%), a characteristic scrub-like heath on ultrabasics, and in rain forests (87%), which occur both on and off these substrates; endemism is lower in the remnant stands of sclerophyllous forest (59%), which are New Caledonia’s most threatened habitat type, covering only 2% of the its original approximately 4,500 km2. An estimated 50 species face impending extinction or have already been lost from the New Caledonian flora. The threat to the flora and vegetation in the other principal habitats is currently much lower than in most tropical areas because of the island’s comparative political and economic stability, but this situation could easily change. Keywords:

Endemism; Extinction; Gymnosperms; New Caledonia; Primitive angiosperms;

Sclerophyllous forest; Threatened habitats. ______________________________________________________________________________________________________ 1

Mailing address: Laboratoire de Phanérogamie, Muséum National d’Histoire Naturelle, 16, rue Buffon, 75005 Paris, France. E-mail: [email protected]

Introduction New Caledonia, a French Overseas Territory located in the southwest Pacific about 1,200 km east of Queensland, Australia and approximately 1,500 km north-northeast of New Zealand, and situated between approximately 19°30' and 22°40' S latitude (Figure 1), has long been of particular interest to botanists, primarily because of its exceptionally rich flora, its remarkable levels of specific and generic endemism, and the presence of extensive areas with ultrabasic substrates where many species have adapted to the very special and highly selective edaphic conditions that they present. The island is also home to numerous apparently relict representatives of the late Cretaceous-early Tertiary Gondwanan flora of Australasia, including members of several primitive angiosperm families and a particularly rich gymnosperm flora, which have survived and radiated on New Caledonia, but are now greatly reduced in numbers or have entirely disappeared from other parts of the region. The origin, evolution, and biogeographic history of New Caledonia’s native plants and vegetation have received much attention over the last 125 years, and as a result the island’s flora is among the best known and most thoroughly documented of any tropical area (Balansa, 1873; Balgooy, 1960, 1971; Baumann-Bodenheim, 1956, 1988, 1989a,b,c, 1990; Bouchet et al., 1995; Brousemiche, 1884; Däniker, 1929, 1931, 1939; Good, 1955; Guillaumin, 1921, 1924, 1928, 1934, 1948, 1953a,b, 1954, 1964; Holloway, 1979; Jaffré, 1974, 1980, 1993, 1995; Jaffré and Veillon, 1991, 1995; Jaffré et al., 1987, 1993; Lowry, 1991; Morat, 1993; Morat et al., 1981, 1984, 1986, 1995; Raven, 1980; Raven and Axelrod, 1972, 1974; Sarasin, 1917; Schlechter, 1905; Schmid, 1987; Thorne, 1963, 1965, 1969; Virot, 1956). New Caledonia has been the subject of intensive botanical exploration throughout much of the 20th century (cf. MacKee, 1964, 1972; Morat, 1993, 1995), which has produced an exceptionally rich base of plant specimens for studies in systematics, ecology, vegetation analysis, phytogeography, and other disciplines. The large amount of high-quality herbarium material now available to botanists also serves as a key resource for the published series Flore de la Nouvelle-Calédonie et Dépendances (Aubréville et al., 1967-present), which to date covers 45 families of angiosperms and gymnosperms, and 26 families of vascular cryptogams, including a total of 1,281 species, representing almost 27% of the approximately 4,780 vascular plant species present in the territory, of which about 1,400 are introduced or cultivated (MacKee, 1994) and 3,380 are regarded as native (Morat, 1993). Moreover, as indicated by Morat (1993), if one also considers manuscripts for the Flore that have been submitted or are now in press, as well as recent articles dealing with the taxonomy and systematics of groups represented on New Caledonia, nearly 70% of the territory’s flora has been the subject of modern treatments. As pointed out by Jaffré (1993), however, the intensity of botanical collecting is uneven within the territory, with the northeast being the least well documented. Within the Territory of New Caledonia, three geographically distinct entities can be recognized, as follows (Figure 2).

1. The principal island, Grande Terre, is approximately 390 km long and about 50 km wide on average, covering an area of 16,890 km2. It is continental in origin, and is oriented along a north-northwest--south-southeast axis, with extensions in both directions to several smaller, geologically related islands to the north-northwest (Iles Bélep, Ile Baaba, Ile Balabio) and the south-southeast (Ile Ouen, Ile des Pins). 2. The Loyalty Islands (Ouvéa, Lifou, Tiga, and Maré), situated approximately 200 km to the northeast of Grande Terre, and covering about 1,970 km2, are geologically distinct, being of much more recent oceanic origin (Dubois et al., 1973). 3. A number of small, isolated, geologically young islands (the Chesterfields, Beautemps-Beaupré, Walpole, Hunter, and Matthew) and reefs (Bellone, d’Entrecasteaux, Astrolabe) are also administratively part of the territory of New Caledonia (Dubois et al., 1981). Essentially all of New Caledonia’s botanical diversity is, however, concentrated on Grande Terre.

The

depauperate, relatively species-poor floras of the smaller, younger islands, including the Loyalties (with less than 400 species in 1,970 km2; Jaffré, 1993), contain only a minor sub-set of the diversity occurring on the main island, reflecting the fact that they have become established relatively recently through long-distance dispersal from sources occurring almost exclusively on Grande Terre, and perhaps to a much lesser degree from other areas such as Vanuatu (formerly the New Hebrides). Moreover, the smaller islands contain only a few locally endemic species, such as the palm Cyphophoenix nucele, which is restricted to a single population on Lifou (cf. MacKee et al., 1985; Jaffré and Veillon, 1989) and no unique vegetation types (although a specialized form of moist evergreen forest on limestone is particularly well represented on the Loyalty Islands; see below), and they are therefore of limited interest botanically. For these reasons, only Grande Terre will be taken into consideration in the remainder of this paper.

Geology and Soils The evolution of New Caledonia’s flora and vegetation has been strongly influenced by its geological and tectonic history. Grande Terre (along with its associated islands) represents an exposed section of a complex orogenic belt situated on the Norfolk Ridge, flanked on both sides by deep ocean (Guillon, 1974; Lapouille, 1981; Lillie and Brothers, 1970; Paris, 1981a,b). New Caledonia separated from Australia about 65 million years ago as part of the Rangitata Orogeny, and drifted to the northeast, reaching its present position about 50 million years ago (Coleman, 1980; Raven, 1980). During the Tertiary (in particular the Paleocene and middle Eocene), large parts of New Caledonia underwent a series of submersions, and by the late Eocene nearly all of the island was covered with up to 2,000 m of peridotites, a type of igneous rock formed from ocean crust that was slowly over-thrust during the preceding tectonic movement (Guillon, 1975; Moores, 1973; Paris et al., 1979). Geologists have contended that during at least some of this period all of the land area comprising New Caledonia must have been below the ocean surface. Inference from the modern flora, however, strsongly suggests that at least a portion of the land must have remained exposed throughout this process, serving as a refugium—

although these sites may have been situated to the south and/or west of the present-day Grande Terre in areas that are now submerged. Many attributes of New Caledonia’s flora, such as its high generic and familial diversity, and the presence of numerous primitive groups, would be particularly difficult to explain by invoking long-distance dispersal over the large water barrier that separated the island from the Australian continent throughout the Paleogene.

Moreover, many groups, including some that have diversified and radiated extensively in New

Caledonia, appear to be poorly adapted to wind or animal dispersal. The most parsimonious interpretation is that a substantial component of today’s flora comprises the descendants of pre-Eocene Australasian lineages that were able to survive while being rafted on one or more continually exposed portions of the New Caledonian land mass as it separated and drifted away from Australia. The extensive ultramafic substrates (peridotites, and to a lesser extent serpentinites) that once covered nearly all of the Grande Terre and its associated islands have now been reduced substantially by erosion (Guillon, 1969, 1975; Guillon and Routhier, 1971; Trescases, 1969, 1975), but today still cover about 5,500 km2, or about one third of the total land area (Jaffré et al., 1987). These substrates have given rise to New Caledonia’s characteristic ultrabasic soils (Latham, 1975a,b,c, 1981, 1986; Latham et al., 1978), which have exceptionally high levels of Fe, Mg, and of several heavy metals such as Ni, Cr, Co, and Mn, elements that are generally toxic to plants, along with very low levels of N, P, K, Ca, and Al (Jaffré, 1976; Jaffré et al., 1987). These chemical imbalances, and especially the ratio of Ca to Mg, result in highly specialized edaphic conditions that have had a profound influence on the evolution and diversification of New Caledonia’s flora and vegetation (Jaffré, 1980; Jaffré and Latham, 1974; Jaffré et al., 1970, 1997b; Lee et al., 1977a; see also below). Elsewhere the soils are more typical of tropical environments, including vertisols, alluvial and brown tropical soils, as well as ferrsiallitic and non-ultrabasic ferrallitic soils, among others (Latham, 1981).

Climate and Geography New Caledonia’s climate is similar to that found on other tropical islands situated at comparable latitudes and which have substantial geographic relief. During the warm season (mid-November to mid-April), frequent tropical depressions and cyclones produce large amounts of precipitation and can cause localized damage to the native vegetation. After a brief transition, during which the Intertropical Convergence Zone shifts to the north, the cool season (mid-May to mid-September) begins, during which rainfall amounts are lower, resulting from cold fronts associated with polar low pressure areas. This is followed by a transition period (mid-September to mid-November) with generally clear weather and increasing southeasterly trade winds (Service Métérologique de la NouvelleCalédonie et Dépendances, 1981). Average annual temperatures at the territory’s numerous weather stations (all of which are situated below 600 m elevation) vary between 22°C and 24°C; substantially cooler temperatures occur at higher elevations, although official records are not available. The warmest month is February, during which average temperatures range from about 25°C to 27°C; the coolest period is July and August, when averages are from about 18°C to 22°C (Section d’Hydrologie de l’ORSTOM and Service Territorial de la Métérologie, 1981).

The geography of Grande Terre is dominated by a chain of high mountains that run along its entire length, comprising numerous massifs above 1,000 m, and five summits that exceed 1,500 m (Mt. Panié, 1,628 m; Mt. Humboldt, 1,618 m; Mé Maoya, 1,508 m; Mt. Colnett, 1,505 m; and Mt. Kouakoué, 1,501 m). The central mountain chain has a profound influence on precipitation patterns, generating high levels of orographic rainfall when the dominant winds coming from the east-northeast to southeast sector are uplifted as they encounter the island’s relief. As a consequence, precipitation is highest along the windward, eastern side, where annual totals range from nearly 2,500 mm to over 4,000 mm in some areas. Rainfall is also high in the central mountain massifs, especially on their eastern slopes, and levels well in excess of the recorded maxima probably occur on most of the highest peaks. By contrast, the leeward western side of the island lies within a marked rain shadow formed by the central chain, and annual precipitation levels are generally well below 1,500 mm, reaching only 1,000 mm in many areas, and averaging below 800 mm at Ouaco (Section d’Hydrologie de l’ORSTOM and Service Territorial de la Métérologie, 1981; Danloux, 1993); in the driest years rainfall can be as low as 250–300 mm at some sites (Danloux, 1987). Thus, many of the physical and climatological conditions occurring on New Caledonia are similar to those found on numerous other moderately sized tropical islands.

However, New Caledonia’s geological and

phytogeographic history, combined with its large areas of highly selective ultrabasic soils, present unique dimensions that have resulted in the evolution of an impressively diverse flora and exceptionally high levels of endemism.

Vegetation and Chorology A wide range of vegetation types are present on New Caledonia, reflecting the various climatic and edaphic conditions found on the island, which have a profound influence on the vegetation’s structure and its distribution, as elsewhere in the world. The work of Morat et al. (1981; see also Morat et al., 1995), expanding on previous studies (Aubréville, 1965; Balansa, 1873; Guillaumin, 1921; Schlechter, 1905; Virot, 1956), describes and maps broad categories of natural and anthropogenic vegetation based primarily on physiognomy (see also Lowry et al., 1997, for additional comments on vegetation classification). These include three types of moist evergreen forest (Figures 3 and 4): 1) low- and mid-altitude forests, and 2) high altitude forests (both of which occur on ultrabasic as well as other substrates such as basalt, schist, etc.), and 3) an edaphic variant of these forests occurring on limestone. Together the moist evergreen forests, which have a closed canopy comprised of moderately sized trees (up to 20 m at lower elevations; about 3–8 m at higher altitudes), cover approximately 22% of the territory, including about 3,000 km2 on Grande Terre, found principally in the east and the mountainous center of the island where rainfall is highest, and an additional approximately 1,000 km2 on limestone substrates primarily in the Loyalty Islands and on a portion of the Ile des Pins. Sclerophyllous forests (Figure 5) occur on Grande Terre and some of its associated islands, mostly on various sedimentary substrates (chert, sandstone, limestone, etc.), and in the past on Cretaceous basalts, where they have now been completely removed. This dense, closed canopy formation, found between sea level and about 300 m

elevation, is dominated by semi-deciduous trees about 10–12 m in height, with a woody evergreen, sclerophyllous middle stratum and abundant vines (Jaffré et al., 1993; Morat et al., 1995). Sclerophyllous forests were once widespread in the drier west of Grande Terre, where their presence is determined primarily by climatic rather than edaphic factors. Some of the elements that are characteristic of this formation also occur in the moist evergreen forests on limestone in the Loyalty Islands (Veillon, 1993). Today, however, pristine stands of sclerophyllous forest are extremely restricted, covering only about 100 km2, or just over 2.2% of their estimated original area of about 4,500 km2, with an additional approximately 250 km2 of variously degraded and secondarized sclerophyllous forests also remaining (Jaffré et al., 1993; Bouchet et al., 1995). Maquis vegetation (Figure 6), a characteristic low, sclerophyllous, evergreen, heath-like formation that is largely restricted to ultrabasic substrates at low, middle and high altitudes on Grande Terre and its associated islands, covers around 30% (4,500 km2) of the territory (smaller areas of maquis also occur on acidic substrates in the north of the island). Mangrove vegetation, covering about 200 km2) is found in its typical habitat, in areas with saline water subject to tidal influence, with somewhat different floristic composition along the east and west coasts (Veillon, 1993; Morat et al., 1995). Swamp vegetation occurs in localities with periodic standing water, including some in areas with ultrabasics (e.g., the Plaine des Lacs), and littoral vegetation, with widespread Indo-Pacific species, is found in close proximity to the ocean. Secondary vegetation now covers a little over 50% of New Caledonia’s land area, and is recognized by Morat et al. (1981) to comprise various degraded forms of the natural vegetation types. These anthropogenic formations include secondary forests and “savannas” (see Lowry et al., 1997, and references therein regarding the use of this term), which often occur together in a mosaic, as well as thickets, which are considered to represent the most degraded form of vegetation resulting from human action (fire, overgrazing, land clearing, etc.). Variants of these types are found in both wet and dry parts of the island, and on various substrates, each reflecting to at least some degree the native vegetation type that was originally present. No comprehensive chorological analysis of the New Caledonian flora has yet been made to assess phytogeographic patterns within the territory, despite the availability of an exceptionally large and well documented specimen base in numerous groups that have been the subject of modern systematic revisions. Jaffré and Veillon (1989), however, recognized five phytogeographic sectors using distributional data from New Caledonia’s 32 native palm species (see also Jaffré, 1993). It would be interesting and informative to expand this approach to include additional taxa (see, for example, Lowry et al., 1997; White, 1993).

Diversity and Endemism The most recent published estimates indicate that New Caledonia has a total of approximately 4,780 vascular plant species, about 1,400 of which are introduced or cultivated (MacKee, 1994), with the remaining 3,380 being regarded as indigenous (Jaffré et al., 1993; Morat, 1993). Among the native species, 243 are vascular cryptogams representing 75 genera in 26 families (as treated by Brownlie, 1969), 44 are gymnosperms totaling 15 genera in 5

families (de Laubenfels, 1972; Jaffré, 1995), and the remaining approximately 3,093 species are flowering plants (around 80% of which are dicotyledons) in about 750 genera and 165 families. According to Morat (1993), however, these figures, which are based on currently recognized taxa, substantially underestimate the true size of the flora. Given the rate at which new taxa are being discovered (whether in the herbarium or the field) as individual groups are studied in detail, Morat has suggested that an additional 5–10% of the species remain to be described, which would bring the total native flora to about 3,550–3,700 species in all. The richness of the New Caledonian flora can be more fully appreciated when its density, as measured by the number of native angiosperm species per 1,000 km2 of land area, is compared with similar values for other tropical and subtropical areas noted for their botanical diversity (Table 1).

While several of the areas cited have

substantially more total species than New Caledonia (e.g., Peru, Madagascar, and the Cape Region of Southern Africa), each of them is considerably larger in size, resulting in much lower density values. Among the areas presented in Table 1, only Puerto Rico and Jamaica have more species per unit area than New Caledonia. The flora of New Caledonia is, however, somewhat disharmonic, as one might expect for an island that has been isolated for such a long period of time. Several typically tropical families are remarkably speciose, while others are poorly represented or entirely lacking. For example, the four largest families in the New Caledonian flora (Myrtaceae, Rubiaceae, Orchidaceae, and Euphorbiaceae) comprise between 193 and 223 species each, for a total of 831 species, or 26.9% of the entire flowering plant flora (Morat, 1993). Another seven families contain about 80 or more species each (Poaceae, Apocynaceae, Cyperaceae, Cunoniaceae, Rutaceae, Araliaceae, and Sapotaceae), representing an additional 660 species in all (21.3% of the angiosperm flora). By contrast, only one native species of Melastomataceae is present, and no Begoniaceae or Ochnaceae (sensu stricto) occur in the territory. Likewise, several other families, such as Araceae, Boraginaceae, Brassicaceae, Commelinaceae, Gesneriaceae, and Zingiberaceae, are substantially under-represented in New Caledonia. Recent compilations also indicate that fully 76.4% of New Caledonia’s native angiosperm and gymnosperm species are strictly endemic to the territory, and in particular to Grande Terre, while slightly more than 14% of the native genera are also endemic (Jaffré et al., 1993). Moreover, depending on the system of classification, as many as five families can be recognized as occurring only on New Caledonia:

Amborellaceae, Oncothecaceae,

Paracryphiaceae, Phellinaceae (often included in Aquifoliaceae), and Strasburgiaceae (usually placed in Ochnaceae). These data clearly indicate that New Caledonia’s native flora is among the most highly endemic in the world. Comparison with recent estimates for other areas with exceptionally high proportions of endemic angiosperm species (Table 2) shows that only a few have higher figures, including Hawaii (89.2%), New Zealand (81.9%), and Madagascar (about 80%), while several others, such as the Cape Region (68%), Juan Fernández (67.9%), Fiji (61.4%), Peru (31.2%), and Taiwan (approximately 25–30%), fall well below the proportion found on New Caledonia.

As pointed out by Morat (1993), however, levels of diversity and endemism vary substantially among New Caledonia’s three principal vegetation types (Table 3). The moist evergreen forest formations are the richest, with 2,009 species (Jaffré et al., 1997b), of which 87.2% are endemic to the territory (Morat et al., 1984; updated by Jaffré et al., 1993), and with 484 genera, including 20.0% endemics. The maquis vegetation has a less diverse flora, with 1,082 species and 306 genera, but its levels of endemism are slightly higher for species (91.2%) and nearly as high for genera (19.0%) (Morat et al., 1986; Jaffré et al., 1993). Figures for the sclerophyllous forests are comparatively lower, with only 379 species (58.8% endemic), and 227 genera (4.8% endemic), although this very likely reflects at least in part the fact that this formation has been reduced to a number of small, isolated stands, many of which have been subjected to varying levels of degradation (Jaffré et al., 1993; Bouchet et al., 1995; see also below). When these figures are considered in light of the estimated original area that each of the main vegetation types was thought to occupy on Grande Terre prior to being impacted by humans, the richness of the moist evergreen forests stands out even further (Table 3). The higher number of species occurring in this formation is compounded by the fact that it only covered about 3,000 km2 of land area, or a little over half the estimated figure for the maquis, and two thirds that of the sclerophyllous forest. Thus, the moist evergreen forests contain about 670 species per 1,000 km2 of estimated original cover, whereas maquis has only 240 species, and sclerophyllous forests about 84 species per 1,000 km2. This indicator probably underestimates the original richness of the sclerophyllous forests, as suggested above. Nevertheless, even prior to human caused degradation, it seems unlikely that the floristic diversity originally present in this formation equaled that of the maquis, and it almost surely did not approach that of the moist evergreen forests. The distribution of floristic within New Caledonia’s moist evergreen forests differs according to substrate (Jaffré et al., 1997b). This type of forests on ultrabasic substrates covers about 1,800 km2 and contain an estimated 1,360 species representing 400 genera, whereas on acidic soils (ca. 2,200 km2) it has about 1,367 species in 431 genera, and on calcareous substrates (ca. 900 km2) only 225 species in 155 genera. Levels of endemism also differ between ultrabasic and non-ultrabasic substrates, regardless of vegetation (Jaffré et al., 1987, 1997; Morat, 1993). For example, while the total number of angiosperm and gymnosperm species occurring in the two main formations that are present on both kinds of substrate (moist evergreen forest and maquis) is essentially identical, with 1,844 species on ultrabasics versus 1,840 on non-ultrabasics (of which 668 are found on both), 90.6% of the species on ultrabasic substrates are endemic to the New Caledonian flora, while only 68.1% of those on other substrates are restricted to the territory (Morat, 1993). Similarly, of the 440 genera represented on ultrabasics, 20.7% are endemic, as compared to only 9.4% endemism among the 170 genera on non-ultrabasic substrates. Perhaps even more remarkable is the fact that among the 1,176 species that occur exclusively on ultrabasics (37.5% of the total flora), fully 98.0% are endemic to New Caledonia. Likewise, 72 genera (9.4% of those present in the territory) are restricted to these highly selective substrates, of which 52.8% (38 genera in all) are also New Caledonian endemics. Recent plot studies conducted in several areas of moist evergreen forest further demonstrate the floristic diversity of this vegetation type in New Caledonia. A total of 131 species with stems

10 cm in diameter at breast

height (dbh) were found in a 1.25 hectare plot established at approximately 160–250 m elevation on ultrabasic substrate located on a gentle slope in the Rivière Bleue Reserve, and 103 species were recorded in a similar plot situated in a nearby area of alluvial forest at about 160 m elevation (Jaffré and Veillon, 1990). In another study, between 93 and 103 species with a dbh

10 cm were identified in three 1.0 hectare plots located between 400 and

500 m elevation in forest on schist near the Col d’Amieu (Jaffré and Veillon, 1995). These figures are substantially lower than those for certain forests in New Guinea (116–147 species per 0.8 hectare), Sarawak (123–223 species per 1.0 hectare), Kalimantan (128–149 species per 1.0 hectare; 205–239 species per 1.6 hectare), and Amazonian Brazil (179 species

15 cm dbh per 1.0 hectare), but are equal to or higher than the numbers for many other areas

such as Amazonian Venezuela (63–83 species per 1.0 hectare), Surinam (59–94 species per 1.49 hectare; 135 species per 1.25 hectare), Nigeria (23–47 species per 1.49 hectare), Cameroon (73 species per 1.49 hectare), and other sites in Amazonian Brazil (60–87 species per 1.0 hectare) (references cited in Jaffré and Veillon, 1990, 1995). The species data from plots in New Caledonia are also similar to those obtained from recent studies in Madagascar, in which for example approximately 110 species with a dbh

10 cm were recorded in a 1.0 hectare

plot in moist evergreen forest at 950 m in Ranomafana National Park (Schatz and Malcomber, in press), and 112 to 116 species were identified in two 1.0 hectare plots in littoral forest on sand at 0–10 m altitude in the Sainte Luce forest (Rabevohitra et al., 1996). The diversity in New Caledonian moist evergreen forests is substantially higher, however, than in several other littoral forest areas examined in Madagascar (some of which were partially degraded), where only 38 to 82 species were found per 1.0 hectare plot (Rabevohitra et al., 1996).

Origin of the New Caledonian Flora The flora of New Caledonia is characterized by the presence of a large number of lineages that appear to represent remnants of the late Cretaceous-early Tertiary Gondwanan flora that is thought to have once covered large parts of Australasia. As New Caledonia separated from Australia and drifted to its present position, it almost surely carried with it a sample of this early flora, the descendants of which have been able to survive in the island’s relatively equable climate, while most of their relatives in Australia and other parts of the region were lost as a result of the well-documented changes in climatic conditions (in particular drying) that took place in Neogene times (Holloway, 1979; Kershaw, 1984; Raven, 1980; Raven and Axelrod, 1972, 1974). There are many remarkable and well-known examples of such relictual endemics in New Caledonia, and they are widely regarded as an indication of the ancient origins of the island’s original flora. For example, 43 of the 44 native species of gymnosperms (Figures 7–9) are endemic to the territory (de Laubenfels, 1972), representing almost 7% of the world’s total gymnosperm flora (see also Jaffré, 1995). Similarly, New Caledonia has a high concentration of endemic taxa with vesselless wood, a feature regarded by many as primitive within the flowering plants; these include the single species of Amborella (Amborellaceae), and 18 species of Winteraceae (Figures 10 and 11), currently placed in a single genus, Zygogynum (Vink, 1993; although some authors have recognized Bubbia, Belliolum, and Exospermum as distinct). Other examples of early lineages that have survived on New Caledonia include species of Annonaceae and Chloranthaceae, as well as primitive members of such families as

Arecaceae, Cunoniaceae, Menispermaceae, Monimiaceae (including Atherospermataceae and Trimeniaceae), Piperaceae, Pittosporaceae, Rutaceae, and Sapindaceae (Morat et al., 1984; Raven and Axelrod, 1972), and of Araliaceae (Lowry, 1986; Oskolski et al., 1997; Oskolski and Lowry, submitted), among others. While an important portion of the New Caledonian flora is derived from the primarily Gondwanan lineages that were almost surely present at the time the island separated from Australia, many other groups appear to have reached the territory more recently as part of a general and widespread movement of Indo-Malesian elements that expanded into Australasia during the early and middle Tertiary (Holloway, 1979; Jaffré et al., 1993; see also below). Some of these newer components of the flora have undergone intensive speciation, and today are among the largest genera on the island. Examples include Phyllanthus, with 111 species (Schmid, 1991), Psychotria (about 85 species), and Eugenia (around 37 species), among others (cf. Morat, 1993). New Caledonia’s ultrabasic substrates have long been considered by many authors to have played a central role in the evolution and diversification of the island’s flora (e.g., Holloway, 1979; Jaffré, 1980; Thorne, 1965; Virot, 1956). For example, Thorne (1965) hypothesized that the ultrabasics allowed many older Australasian groups present in the island’s original flora to survive in a sort of local, ecological refugium. By adapting to the special edaphic conditions of the ultrabasics shortly after their deposition in the Paleocene and Eocene, he argued that many of the primitive groups were able to persist on New Caledonia because these substrates subsequently provided a degree of ecological separation and protection from more competitive rain forest taxa that must have continually arrived from other tropical areas, primarily those to the northwest. As indicated above, the flora occurring in the various vegetation types present on New Caledonia’s ultrabasic substrates, in particular the moist evergreen forests and maquis, is especially rich in species and has a remarkably high degree of endemism (90.6% at the species level). Conversely, very few of the island’s non-endemics are found on the extensive ultrabasic substrates. In fact, of the 740 native species New Caledonia shares with other areas, only 173 (23.4%) occur on ultrabasics, while 587 are found on other substrates, with only 20 of these species occurring on both ultrabasics and non-ultrabasics (data derived from Morat, 1993). Jaffré et al. (1987) presented a series of ideas regarding the changes that are thought to have accompanied the establishment of New Caledonia’s ultrabasics, and the processes by which the modern flora occurring on these substrates was established. They suggested that a certain number of species present in the original, pre-Eocene flora almost inevitably must have been eliminated because they were not able to adapt to the extreme conditions of the ultrabasics. Other taxa were able to survive, however, perhaps largely in forest islands on non-ultrabasic substrates that are postulated to have persisted in parts of the northeast in areas that were not effected by the processes of peridotite deposition. Based on a comprehensive analysis of the floristic composition and vegetation of the ultrabasic substrates, Jaffré et al. (1987) further suggested several ways in which various components of the flora were able subsequently to occupy and adapt to the highly selective conditions that they present. First, the authors envisioned an early colonization of the vast unoccupied areas created by the newly deposited ultrabasics involving a small number of

pre-adapted species. These plants were pioneer species that occurred naturally in open habitats within the largely forested areas that are thought to have survived, and which were physiologically able to tolerate the extreme conditions of the ultrabasics (Jaffré, 1980). Today the presumed descendants of these species are found primarily in open climax vegetation on the ultrabasics (i.e., most forms of maquis), and include species of Callistemon, Uromyrtus, Beaupreopsis, Grevillea, Corbassona, Costularia, and Hibbertia, among many others. Second, Jaffré et al. (1987) hypothesized that during the long process by which the ultrabasics were deposited, local conditions in some places favored the development of certain highly weathered, poor, acidic soils whose chemical characteristics resembled those of some soils formed on non-ultrabasic substrates such as schists. These new soils were colonized by forest species that had long been adapted to such conditions, including for example many gymnosperms, Myrtaceae, Sapotaceae, and Proteaceae. Third, the authors suggested that over time many species present in the original, primarily forest flora eventually evolved tolerance to the extreme edaphic conditions of highly basic soils. These might have included plants whose habitat preference is determined more by climatological factors relating to altitude, rainfall, and temperature than to soil conditions, examples of which include several paleo-endemics such as Libocedrus yateensis Guillaumin, and Parasitaxus ustus (Vieill.) Laubenf., the world’s only known parasitic gymnosperm. It seems clear that the initial vegetation occupying the ultrabasic substrates must in any case have been very depauperate, and that these areas consequently presented ecological conditions that were highly favorable for subsequent speciation among the components of the original flora that were able to colonize and survive in them. Jaffré et al. (1987) concluded that a new wave of diversification and adaptive radiation gave rise to an important neo-endemic component of the flora based on these early colonizers, which today accounts for a large portion of the species-level diversity seen on the ultrabasics, especially in maquis vegetation (e.g., Alyxia, Phyllanthus, Morinda, Psychotria, among others; cf. Morat et al., 1986), but also in the moist evergreen forests (e.g., Araucaria, Argophyllum, Geissois, Nothofagus, Phyllanthus, Psychotria, Stenocarpus, Xanthostemon, etc.; Morat et al., 1984). Speciation was particularly active among sclerophyllous groups that are considered to be pre-adapted to poor soils (Beadle, 1966), such as many Myrtaceae, Cunoniaceae, Epacridaceae, Proteaceae, and others. This process also resulted in the evolution of taxa that are able to limit the uptake of Ni or Mg, or to tolerate high concentrations of heavy metals (Brooks et al., 1974, 1981; Jaffré, 1977a,b, 1979; Jaffré and Schmid, 1974; Jaffré et al., 1976, 1979a,b; Kersten et al., 1979; Lee et al., 1977b). There is general agreement that species (and in many instances entire lineages) that are adapted to ultrabasic substrates have benefited, and continue to benefit today, from some degree of protection against allochthonous floristic elements reaching New Caledonia from other areas (e.g., Holloway, 1979; Jaffré et al., 1987; Morat, 1993; Morat et al., 1984, 1986). However, the evidence in support of the idea that this process provided a special, differential protection to the older Australasian elements comprising the island’s original late Cretaceous-early Tertiary flora is by no means unequivocal. If this type of selective protection had taken place on ultrabasic substrates, for example, one would expect to find a generally higher proportion of species belonging to older groups

such as gymnosperms and comparatively primitive angiosperm families in these areas than in zones with nonultrabasic substrates, where competition from more recently arrived, paleo- and pan-tropical elements is almost surely greater. Data presented by Jaffré et al. (1987) show that each of the three main gymnosperm families, Araucariaceae, Podocarpaceae, and Cupressaceae, indeed has more species occurring on ultrabasics than on other substrates (15:6, 18:8, and 6:1 species, respectively), and that the same situation is found among Lauraceae (36:22). However, they also indicate that approximately equal numbers of Winteraceae (11:9) and Annonaceae (8:5) occur on and off of ultrabasics, and that the single species of Amborellaceae is limited to non-ultrabasic substrates. Similarly, equal numbers of Chloranthaceae occur on both kinds of substrate (2:2), while the figures for Monimiaceae (3:8), Atherospermataceae (0:1), and Trimeniaceae (0:1) show more species on non-ultrabasics (Jérémie, 1982). In a more recent study (Jaffré et al., 1997b) examining only the flora of New Caledonia’s moist evergreen forests, a similar pattern was found among the gymnosperms, Winteraceae, Annonaceae and Monimiaceae, whereas Lauraceae were equally represented on ultrabasics (25 species) and other substrates (23 on acidic soils and 2 on calcareous substrates) Moreover, many groups that probably reached New Caledonia well after its separation from Australia, but prior to or during the deposition of the ultrabasic rocks in the Paleocene and middle Eocene, show precisely the same kind of distribution pattern between ultrabasic and non-ultrabasic substrates as the presumably older taxa (Jaffré et al., 1987). Thus, while the presence of ultrabasics may well have provided a kind of ecological isolation and protection against elements that arrived in post-Eocene times, it appears for the most part to have done so indifferently with respect to the primitive and more advanced lineages that were then present on New Caledonia. The various groups occurring in the sclerophyllous forests do not appear to have undergone the same kind of intense speciation that is thought to have resulted in the large numbers of neo-endemics present in the moist evergreen forests and the maquis. A very limited number of genera have more than a few species represented in the sclerophyllous forests (e.g., Eugenia, 13 species; Diospyros, 11 species; Austromyrtus, 8 species), and as indicated above, the species level diversity in this formation is substantially lower than in the two other major vegetation types—although a total of 59 of the endemic species are none the less restricted to the sclerophyllous forests, along with one endemic genus of Rubiaceae (Captaincookia). Also, several groups that are well represented in the moist evergreen forests are completely absent from this formation, including gymnosperms, Arecaceae, Epacridaceae, Fagaceae, Pandanaceae, and Winteraceae, among others (Jaffré et al., 1993; Bouchet et al., 1995). While again this may in part be an artifact of the relictual nature of the remaining stands, it also very likely reflects the relatively young age of this formation, the flora of which is considered to comprise elements largely dating from the Miocene and Quaternary, which were added to a few older, pre-Eocene groups that had been able to survive after the deposition of the ultrabasic substrates (Jaffré et al., 1993).

Phytogeographic Affinities The phytogeographic relations of the New Caledonian flora have interested botanists for many years (e.g., Balansa, 1873; Balgooy, 1960, 1971; Baumann-Bodenheim, 1956; Guillaumin, 1921, 1924, 1934, 1953; Thorne,

1963, 1965, 1969), and have recently been analyzed in detail for the moist evergreen forests (Morat et al., 1984), the maquis (Morat et al., 1986), and the sclerophyllous forests (Jaffré et al., 1993). Together these formations include nearly 82% of the native species (almost 70% of the genera) and cover about 45% of Grande Terre (including nearly all of the remaining primary vegetation), and consequently can be considered as a reasonable and representative sample of the flora for the purpose of quantitative analysis of its affinities with other areas (Lowry, 1991).

Following Balgooy (1971), these studies used the genus as the fundamental unit for assessing

phytogeographic relationships, for the following reasons: a) more homogeneous comparisons can be made at the generic level; b) genera are relatively more stable taxonomic entities through time as knowledge improves and classifications are revised; and c) there is generally less divergence of opinion among systematists regarding the definition of what constitutes a genus. In each study (Morat et al., 1984, 1986; Jaffré et al., 1993), a correlation coefficient was calculated that is proportional to the number of genera common to each of the 15 regions analyzed and New Caledonia (Table 4), and inversely proportional to the total number of these phytogeographic territories in which each of the genera occurs. The relative importance of the floristic relationship between New Caledonia and each of the 15 regions was then expressed as a percentage of the total of the correlation coefficients. The results of these quantitative analyses are very similar for each of the three principal vegetation types (Table 4), strongly suggesting that despite the many differences in the details of their physiognomy and floristic composition, these formations have a fundamentally similar history and a common set of origins, as well as a shared set of mechanisms largely responsible for their establishment, as suggested by Jaffré et al. (1993). Phytogeographic affinities for each vegetation type are by far strongest with Australia, New Guinea, and Malesia—in that order for moist evergreen forests and maquis, and with the order of the last two reversed for the sclerophyllous forests. This finding is consistent with the hypothesis, mentioned earlier, that the New Caledonian flora comprises two principal elements:

the ancient Australasian (Gondwanan) component that was already present on the island when it

separated from Australia, the derivatives of which have survived to this day; and an element originating primarily in New Guinea and Malesia, whose ancestors reached New Caledonia relatively more recently by long-distance dispersal. By contrast, comparatively few groups present in New Caledonia exhibit phytogeographic affinities with other tropical Pacific islands such as Fiji, Vanuatu, the Solomons, and Samoa-Tonga, or with areas farther to the south including Norfolk Island, Lord Howe Island, and New Zealand (Table 4). In the case of the tropical Pacific islands, as well as Norfolk and Lord Howe, this is for the most part probably a reflection of the fact that they constitute rather small source and target areas for long-distance dispersal, coupled in some cases with their young age (i.e., for the volcanic islands of recent origin) and their generally less diversified climatic and edaphic conditions. The low phytogeographic affinities of the more southern, largely temperate floras, especially New Zealand’s, with that of New Caledonia, most likely result primarily from the strong climatic differences between the territories.

Threats, Extinction, and Conservation Of the various native vegetation types that occur on New Caledonia, only one, sclerophyllous forest, currently faces serious threats to its continued existence. The moist evergreen forests are subjected to some logging and occasional land clearing (ca. 3-5% between 1975 and 1995, according to Morat et al., 1995), as well as destruction from mining activities, which is primarily concentrated at higher elevations in a number of massifs (although mining was far more extensive during the nickel “boom” of the 1960s and early 1970s), but the impacts are nevertheless localized, and affect only a small portion of the approximately 3,000 km2 that this formation occupies on Grande Terre. The maquis vegetation is substantially more vulnerable to fires, which are widespread in New Caledonia (often set by humans), especially during the dry season and primarily in areas with lower levels of precipitation, and which have resulted in varying degrees of degradation to an undefined (but apparently rather small) portion of the estimated 4,500 km2 occupied by this formation. Mining activities also have taken their toll on maquis at a number of sites, resulting in degradation or total destruction (Morat et al., 1995), but because this vegetation type is almost exclusively restricted to ultrabasic substrates, which are essentially otherwise useless to humans, especially for agriculture (cf. Bonzon et al., 1997), it is thus only very slightly impacted (but see Jaffré et al., 1997a). Mangrove and swamp vegetation, as well as the other native formations, most of which do not contain high numbers of locally endemic species, also appear at present to be relatively unthreatened in general, although individual sites are of course vulnerable to a wide range of destructive activities. A few species have been effected by over-exploitation (Morat et al., 1995). The situation for New Caledonia’s sclerophyllous forests, however, stands in sharp contrast, and their conservation status is far more serious. The history of destruction that has ravaged this unique vegetation type, and the plight of the few remaining stands present in the territory, are the subject of a recent article in which an urgent call is made for immediate conservation action (Bouchet et al., 1995). As indicated above, only about 100 km2 (2.2%) of the estimated 4,500 km2 of original sclerophyllous forest cover remains in a more-or-less undisturbed state, with another approximately 250 km2 (5.6%) that are moderately to heavily degraded, and are in most cases highly secondarized. Moreover, the remaining stands are very small (mostly covering less than 5 hectares, and never more than 200 hectares), and are highly fragmented and scattered along the west coast of Grande Terre and on a few of its associated islands (Jaffré et al., 1993), which presents additional problems for their continued viability. The causes of the wholesale destruction of such large areas of sclerophyllous forest (large at least on the scale of New Caledonia) are essentially the same as for many other dry tropical forests (cf. Humbert, 1935), which are often regarded as among the world’s most threatened habitats (e.g. Janzen, 1988). Much of the vegetation cover was apparently badly degraded or completely removed by fires set by humans since their first arrival about 3,500 years ago, but the process was intensified following European colonization in the mid-1800s as lowlands were cleared and burned for various agricultural activities, including cattle grazing. The introduction of the highly successful Indonesian deer in the 1880s, whose numbers grew very rapidly, further accelerated the destruction of the sclerophyllous forests because these animals, along with cattle and goats, impede or prevent the regeneration of

many plant species due to trampling and grazing (Bouchet et al., 1995). More recently, continued clearing for agriculture, and the regular and uncontrolled spread of huge wildfires, often intentionally and illegally set during the dry season as a form of political protest or to facilitate new spring growth of herbaceous species for hunting, have increased the pressure on and vulnerability of the few remaining stands of sclerophyllous forest to an alarming point. The impact of these destructive forces is exacerbated by the fact that flora of the sclerophyllous forests is manifestly insular in nature. Unlike the other principal vegetation types present in New Caledonia, this formation, especially when it has been even slightly degraded and opened up, is highly susceptable to invasion by aggressive, exotic species, such as Cryptostegia grandiflora R. Br., Lantana camara L., Leucaena leucocephala (Lam.) de Wit, and Psidium guajava L., and even a few native taxa, most notably Acacia spirorbis Labill. (Bouchet et al., 1995). Rapid colonization by these and other weedy species, in conjunction with the effects of burning, clearing, and damage from introduced hoofed animals, essentially precludes the possibility that the original sclerophyllous forest can re-establish itself, at least within the time frame of hundreds or perhaps even thousands of years. Because such a high proportion of New Caledonia’s native plant species are endemic, the actual and potential effects of habitat destruction on global biodiversity are correspondingly large. As in many tropical environments, this is further compounded by the fact that a large number of these endemics have highly restricted distributions, occurring in only one or a few localities, and often in a very limited number of small, localized populations (Morat et al., 1995). Many of the 59 endemic flowering plant species that are cited by Bouchet et al. (1995) as occurring only in the sclerophyllous forests (to which an undescribed Polyscias restricted to the Uitoé Peninsula and an area near Pouembout can be added; Lowry, unpubl. data) certainly fit this description, and they are clearly among the island’s most vulnerable taxa. But the risk is by no means limited to these species. Bouchet et al. (1995) have documented the recent extinction of one species endemic to sclerophyllous forests on Leprédour Island (Pittosporum tanianum Veillon and Tirel), and based on a survey of the published volumes of the Flore de la Nouvelle-Calédonie et Dépendances, which cover about 32% of the native angiosperm and gymnosperm flora, they list 15 other species growing in various vegetation types that have not been seen in the last 80 years, many of which are presumed to be extinct. Perhaps twice as many can be assumed to occur in the groups that have not yet been treated in the Flore, which would bring the total to nearly 50 species. Certainly others still must have been lost even before their existence could be documented by botanists, including many that were confined to the formerly widespread sclerophyllous forests, and especially those occurring on basaltic substrates, which have now been completely destroyed (Jaffré et al., 1993). Moreover, essentially all of New Caledonia’s numerous, highly restricted micro-endemics (which probably number in the hundreds) are potentially vulnerable to local disturbance, regardless of the vegetation type or the substrate on which they grow. Clearly the situation regarding New Caledonia’s sclerophyllous forests and the species (including both plants and animals; cf. Chazeau, 1993) that depend on the habitats they provide is critical and of unquestionable

conservation importance, requiring immediate action (see Bouchet et al., 1995, for suggested steps to be taken). Furthermore, considering the richness of the island’s flora and its high level of endemism, New Caledonia— including all of its principal native vegetation types—certainly deserves recognition as one of the world’s biodiversity “hot spots” (Myers, 1988). Although the territory currently has 23 officially recognized terrestrial parks and reserves, covering 54,149 hectares (about 2.9% of the territory), the lack of strong protection, and more importantly the absence of a clearly articulated environmental policy and appropriate regulations that are consistently enforced in each of New Caledonia’s three semi-autonomous provinces, constitute major obstacles to effective long-term conservation (Bouchet et al., 1995). Veillon (1993) points out that several vegetation types are not sufficiently represented in the network of protected areas in order to ensure the viability of the biodiversity they contain. In particular, he lists five formations that require additional protection: 1) sclerophyllous forest, 2) mangrove vegetation, whose floristic composition differs between the east and west coasts, 3) moist evergreen forest primarily on non-ultrabasic substrates in the north and center of Grande Terre, 4) forests in the Loyalty Islands, and 5) maquis on various substrates, mostly in the north and northeast of the island. Despite these facts, however, when viewed in the global context, and compared with the vast majority of tropical areas that are facing far more acute threats from the all too frequent combination of rapid population growth, extreme poverty, and dysfunctional or non-existent government institutions, New Caledonia’s native flora has to be regarded as relatively well off—especially those parts that are found in the island’s moist evergreen forests and its distinctive maquis vegetation. The comparative wealth of the territory (per capita income is similar to that in Australia and New Zealand), coupled with its present political and economic stability, currently preclude many of the highly destructive activities that result each year in the degradation and loss of millions of hectares of native vegetation in other parts of the tropics. New Caledonia is, however, by no means isolated from serious threats to its rich biodiversity through habitat destruction and species extinction. The situation is already critical for the tiny fragments of sclerophyllous forest that remain, and could easily become so for the island’s other formations in the relatively near future, especially if the political environment changes substantially following the re-consideration of possible independence that is scheduled for 1998, and/or if the territory’s economic base is substantially eroded. New Caledonia does not qualify as a resource-rich territory (despite the presence of huge deposits of nickel ore and other potentially valuable minerals), and its current prosperity must be regarded as vulnerable. If appropriate action is taken now by responsible leaders in conjunction with scientists and the conservation community, an exceptional opportunity exists to protect the remarkable botanical diversity of this island for future generations; failure to do so, however, may condemn it to the same irreversible fate that so many other tropical floras have regrettably suffered.

Acknowledgments. Work on New Caledonia was funded in part by a Doctoral Dissertation Improvement Grant from the U. S. National Science Foundation (BSR 83-14691), the Division of Biology and Biomedical Sciences of Washington University, St. Louis, and the Missouri Botanical Garden. Additional support provided by the W.

Alton Jones Foundation, the John D. and Catherine T. MacArthur Foundation, and the Liz Claiborne and Art Ortenberg Foundation is gratefully acknowledged. Courtesies in New Caledonia were generously extended by the ORSTOM Center, Nouméa, the former Service des Forêts et du Patrimoine Naturel, the Service de l’Environnement et de la Gestion des Parcs et Réserves de la Province Sud, and the Service Forêt-Bois-Environnement de la Province Nord. I thank Ching-I Peng for inviting me to participate in the symposium, Ph. Morat, C. Taylor, and two anonymous reviewers for critical comments on an earlier version of the manuscript, and P. Hoch, D. Neill, P. Jørgensen, and J.-M. Veillon for valuable assistance.

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Figure 1. Geographic position of New Caledonia in the South Pacific. Figure 2. New Caledonia: Grande Terre and associated islands (Iles Bélep, Ile des Pins); Loyalty Islands. Figure 3. Moist evergreen forest on Mt. Mov (1,160 m). Figure 4. Moist evergreen forest with Araucaria columnaris at Port Boisé. Figure 5. Remnant sclerophyllous forest on private land southeast of Pouembout. Figure 6. Maquis vegetation on the Plaine des Lacs (ca. 260 m). Figure 7. Araucaria muelleri (Araucariaceae) in the Haute Ouinné basin. Figure 8. Dacrydium guillauminii (Podocarpaceae) along the Rivière des Lacs; a rare, locally endemic species. Figure 9. Callitris sulcata (Cupressaceae) along the Tontouta River. Figure 10. Zygogynum baillonii (Winteraceae) from the Montagne des Sources. Figure 11. Zygogynum stipitatum (Winteraceae) on Mt. Panié.

Table 1. Number of angiosperm species and density per 1,000 km2 for selected areas of high plant diversity. Area

Native angiosperm species

Total area (km²)

Species per 1,000 km²

Puerto Rico Jamaica New Caledonia (Grande Terre only) Cape Region (S. Africa) Taiwan

2,8091 2,8882 3,2503

9,104 10,990 16,890

308.5 262.8 192.4

8,493 3,3504

90,000 35,980

94.4 93.1

Hawaii Madagascar Peru New Zealand

980 10,000 17,119 2,066

16,887 587,040 1,285,220 268,680

58.0 17.0 13.3 7.7

Source Gentry (1986) Adams (1972) Jaffré et al. (1993); Morat (1993) Bond and Goldblatt (1984) Peng et al. (1994); Peng (pers. comm.). Wagner et al. (1990) Phillipson (1994) Brako and Zarucchi (1993) Sohmer (1994)

Includes adventive species. Includes fully naturalized species. 3 Based on Morat’s (1993) conservative estimate that 5% of the native species remain to be described. 1 2

4

Includes species and infraspecific taxa.

Table 2. Number of native angiosperm species and percentage of endemism in selected tropical floras. Area

Native angiosperm species

Hawaii New Zealand Madagascar New Caledonia Cape Region (S. Africa) Juan Fernández Fiji Peru Taiwan 1 2

980 2,066 10,000 3,2501 8,493 157 1,302 17,119 3,3502

Endemism (%) 89.2 81.9 ca. 80 76.4 68 65.0 61.4 31.3 20-30

Source Wagner et al. (1990) Sohmer (1994) Humbert (1959) Jaffré et al. (1993); Morat (1993) Bond and Goldblatt (1984) Stuessy et al. (1997) Sohmer (1994) Brako and Zarucchi (1993) Peng et al. (1994); Peng (pers. comm.)

Based on Morat’s (1993) conservative estimate that 5% of the native species remain to be described. Includes species and infraspecific taxa.

Table 3. Number of native angiosperm and gymnosperm species in New Caledonia, percent endemism, and species density by principal vegetation type1. Vegetation type Moist evergreen forest (all altitudes and substrates) Maquis (all substrates) Sclerophyllous forest

Species (% endemic)

Genera (% endemic)

Estimated original area (km²) (Grande Terre only)

Species per 1,000 km2

2,009 (87.2)

484 (20.0)

3,000

670

1,082 (91.2) 379 (58.8)

306 (19.0) 227 (4.8)

4,500 4,500

240 84

1Adapted in part from Morat (1993) and Jaffré et al. (1993, 1997).

Table 4. Phytogeographic affinities of the New Caledonian flora based on correlation coefficients of floristic territories. Region

Moist Evergreen Forest1

Maquis2

Sclerophyllous Forest3

27.2 20.2 11.7 9.8 7.6 6.8 4.3 3.2 2.8 1.6 1.6 1.4 1.0 0.5 0.3

31.3 16.8 13.3 5.5 6.9 4.2 5.6 5.8 4.3 1.6 1.0 0.6 1.0 1.2 0.9

23.6 14.3 15.2 3.6 7.5 5.2 2.5 10.4 6.3 3.6 3.5 0.0 0.9 1.0 2.4

Australia New Guinea Malesia Fiji Vanuatu Solomon Islands New Zealand Asia Africa Lord Howe America Samoa-Tonga Norfolk Island Polynesia North Pacific From on Morat et al. (1984). From Morat et al. (1986). 3 From Jaffré et al. (1993), corrected. 1 2