A survey of floral traits, breeding systems, floral visitors, and pollination systems of the angiosperms of the Juan Fernandez Islands (Chile).

II. Introduction

The extraordinary faunas and floras of oceanic islands--that is, islands located over oceanic plates that have never been connected to continental land masses (Nunn, 1994)--with unique assemblages of species and high levels of endemism have historically been of interest to naturalists (e.g., Sloane, 1707-1725; Shelvocke, 1726; Linnaeus, 1747; Juan & Ulloa, 1813; Hooker, 1853). Biologists paid even more attention to them following the publications of Darwin's (1859) and Wallace's (1880) books that highlighted these fascinating small areas of the world as remarkable laboratories of evolution. Subsequently, a great deal of insightful work has been published on island biology, particularly in the last three decades (e.g., MacArthur & Wilson, 1967; Carlquist, 1974; Bramwell, 1979; Mueller-Dombois et al., 1981; Williamson, 1981; Bowman et al., 1983; Lawesson et al., 1990; Vitousek et al., 1995; Wagner & Funk, 1995; Keast & Miller, 1996; Grant, 1998; Stuessy & Ono, 1998; Whittaker, 1998). Although much on the biota, ecology, and evolution of islands has been published, much remains to be studied, especially in terms of the biology of the plants. There is increasing urgency for these studies (Carlquist, 1998; Raven, 1998), because insular species are being lost at a higher rate than are their continental relatives (Reid & Miller, 1989; Smith et al., 1993). This is not surprising, given the often smaller populations on islands and, for many island species, a relatively low competitive ability (Carlquist, 1974). Human disturbance of fragile island habitats is one of the principal causes. Indeed, anthropogenic alterations of oceanic islands have taken place on a greater scale than on most continental systems (Loope et al., 1988; MacDonald & Cooper, 1995; Mittermeier et al., 1999).

The biota of the oceanic islands often has a different combination of species than do equivalent mainland areas (Carlquist, 1965, 1974; Grant, 1998). Because islands are difficult to reach, and each continental species has different dispersal capability, it is inevitable that islands will possess a nonrepresentative sample of the species from the near continents. In addition, chance must play an important role in determining which species arrive, when, and in what numbers. Difficulties of establishment will further influence the composition of the island communities by favoring some types of colonists over others. Another basic obstacle is reproduction. For angiosperms in particular, hermaphroditic self-compatible (SC) species would be favored by enabling reproduction and establishment after long-distance dispersal (Baker, 1955, 1967; Stebbins, 1957). If pollen transfer is required--for example, when there is temporal or spatial separation of the sexes, or self-incompatibility (SI) or dioecy--then the presenc e of the pollinator fauna must also be a consideration in establishment. For the reasons given above regarding plant colonization, pollinator faunas on islands also are comparatively small, with many kinds of pollinators either absent or poorly represented (Ehrendorfer, 1979; Barrett, 1998). Thus, unless abiotic pollination, in particular wind pollination (Carlquist, 1974; Ehrendorfer, 1979; Anderson et al., 2000a, 2001a), can be a factor, the lack of pollinators can lead to failure of successful colonization.

In spite of its importance for conservation and restoration purposes, studies of the reproductive biology of island plants in general (Carlquist, 1974; Ehrendorfer, 1979; Godley, 1979; Lloyd, 1985; Barrett, 1998), and of their breeding systems in particular, are comparatively uncommon (e.g., Rick, 1966; Carlquist, 1974; Pandey, 1979; Carr et al., 1986; McMullen, 1987, 1990; Connor, 1988; Webb & Kelly, 1993; Anderson et al., 2001a). There are no detailed, comprehensive species-based reviews of these aspects on any oceanic island system. Technical and economic difficulties in reaching islands, in locating species in the commonly inaccessible, steep habitats (for islands of volcanic origin) where they grow, and in finding the plants in the right reproductive stage all seem to have contributed to this paucity of studies. Data on pollination-related floral morphological traits of the species may be used to postulate how pollination is accomplished in these isolated island floras. There has been some analysis of th e general characteristics of flowers in the context of plant reproduction. For instance, some authors have called attention to the high representation of small, white or green, simple-shaped flowers in the floras of oceanic islands and have suggested a correlation with the paucity of native pollinators (Wallace, 1895; Carlquist, 1974; Webb & Kelly, 1993).

We chose the Juan Fernandez Archipelago flora for a comprehensive study for a number of reasons. First, we had the opportunity to do much original fieldwork via several expeditions to the islands, mainly to Robinson Crusoe Island, and have published on the reproductive biology of elements of the flora (Bernardello et al., 1999, 2000; Anderson et al., 2000a, 2000b, 2001a, 2001b). Secondly, the angiosperm flora of the archipelago is, like the area of the islands (100.2 [km.sup.2]; Stuessy, 1995), comparatively small (156 taxa, 152 species; Marti-corena et al., 1998) and fairly well known. The flora also has a high level of endemism (about 63%). At 2.1 native species/[km.sup.2] and approximately. 1 endemic species/[km.sup.2], the autochthonous species density of the native flora and the density of endemics is higher on Robinson Crusoe Island than on any other oceanic island. Finally, this flora is among the most threatened in the world (Allen, 1984; Davis et al., 1995; Mittermeier et al., 1999). The native flora is characterized by low fire tolerance and poor adaptation to herbivore resistance (Skottsberg, 1953). The species are especially vulnerable to human-induced disturbance--that is, to the historical and continued foraging by feral goats and rabbits--and to losses of habitat due to aggressive introduced exotic weeds (Perry, 1984; Wester, 1991; Bourne et al., 1992; Stuessy et al., 1997; Jaksic, 1998). Thus, the flora is of manageable size, of particular interest, and highly vulnerable and threatened.

The Juan Fernandez Archipelago is one of the few regions where there were no permanent human settlements before the sixteenth century (Woodward, 1969; Wester, 1991). It consists of three islands, all of volcanic origin: 667 km W of continental Chile are Robinson Crusoe Island (= Masatierra) and Santa Clara Island; Alejandro Selkirk Island (= Masafuera) is 181 km farther west. Robinson Crusoe Island has been dated at approximately 4 million years old; Alejandro Selkirk Island, at 1-2 million years old; and Santa Clara Island, at 5.8 million years old (Stuessy et al., 1984).

In this article, features of the angiosperm flora of the archipelago are analyzed and statistically compared. Although in progress (Stuessy, in prep.) there is not yet available a comprehensive modern flora of the archipelago; thus, the collection of original data was extensive. Information was gathered primarily from our own fieldwork, and also from the existing literature. The published literature offered relatively little direct information on reproductive biology, but much on floral features. Data recorded include habit, plant sex, several flower features, such as size, shape, and color, and the hypothesized pollination system of the first colonizers. In addition, we summarized the available data on compatibility, presence and type of dichogamy, observed floral visitors, presence of floral rewards, and currently known pollination systems. Our goals were to provide a species-based review for as much of the flora as possible of the features relevant to reproduction and pollination, to identify associations and generalizations, to contribute to the understanding of the colonization and evolution of the angiosperms of the archipelago, and to promote use of these data for conservation, preservation, and restoration.

III. Methods and Materials

The recent catalogue of the vascular flora of the Juan Fernandez Islands by Marticorena et al. (1998) was used for the list of species on the archipelago and their habit. Information on sex and flower features are our own observations, mainly on Robinson Crusoe Island, supplemented by data extracted from Gay (1845-1854), Hemsley (1884), Johow (1896), Reiche (1896-1911), Skottsberg (1921, 1928, 1953), and Moore (1983). Data on breeding system, floral biology, presence of reward in the flowers, floral visitors, and pollination are largely personal observations, supplemented by the published data on visitors extracted from Johow (1896), Skottsberg (1928), Brooke (1987), Meza (1988), and Colwell (1989), and on breeding systems of some species from Ramanna and Hermsen (1981) and Moore (1983). The pollination biology of the hypothesized progenitors or colonizers was estimated after reviewing the literature on the closest continental relatives and publications on other southern Pacific oceanic islands (e.g., Carlqui st, 1974; Porter, 1983; Wagner et al., 1990).

Our observations and experiments were largely confined to natural populations on Robinson Crusoe Island (cf. Bernardello et al., 1999, 2000; Anderson et al., 2000a, 2001 a). In addition, many experimental studies, especially on compatibility, were conducted with native plants grown in the Corporacion Nacional Forestal (CONAF) gardens adjacent to San Juan Bautista, the only permanent settlement on the archipelago. Finally, a limited number of observations were made with material from Alejandro Selkirk Island.

To facilitate the comparisons, flower sizes were placed into five categories based on the flower area (lengthxwidth) as follows: very small = <9 [mm.sup.2], small = 10-25 [mm.sup.2], medium-sized = 30-110 [mm.sup.2], large = 120-400 [mm.sup.2], very large = >400 [mm.sup.2]. We recognized these groups by plotting the average data obtained for every species of the flora and identifying the emergent groups. Shape categories follow the classification of Faegri and van der Piji (1979)--but see our comment in section A.3 below. Color categories were defined by reducing all of the kaleidoscope to just six color categories: yellow, white, green, blue (including purple, lavender, and violet), brown (including straw colored), and red (including pink and orange); orange was considered separately only when comparing flower color with current pollination. When a species had flowers with two or three colors (either multicolored or with different nuances), we coded the most prominent color. All comparisons were made at the species level, except for the hypothesized pollination of colonizers, which we recorded at the genus level.

Statistical comparisons were made using the Statistical Analysis System (SAS for Windows version 8.00, 1999, Cary, NC). The hypothesis tested was that there was no association among the variables. The PROC FREQ was performed using the general association statistic ([Q.sub.CMH], Stokes et al., 1995), which produces both the Pearson chi-square statistic and the Mantel-Haenszel statistic as well as the general association coefficient. The analyses were applied first to the whole series of data in each category. Then, if the result was significant at p <0.01, the same analyses were applied to pairwise combinations of variables to identify significant associations.

IV. Results

All relevant data from our studies and from the literature are summarized in Table I. An analysis of each feature, and pairwise comparisons, where appropriate, follow.

A. HABIT AND REPRODUCTIVE FEATURES

1. Habit

About 85% of the flora comprises perennials: 38% are perennial herbs, 23% are shrubs, 16% are trees, and 9% are shrubs/trees.

2. Flower Size

The majority of the species have small flowers (60%), falling into the very small (46%) or small (14%) ranges. Medium-sized flowers are found in 24% of the species of flora. Large (9%) and very large (7%) flowered species are uncommon.

3. Flower Shape

Most shape descriptors are obvious; the term "inconspicuous" is not. Faegri and van der Pijl (1979) and Bell (1991) use "inconspicuous" basically as a "shape" descriptor. However, this shape descriptor is defined as "with no optical attraction" by Faegri and van der Pijl (1979). Thus, flowers in this category would not seem to have a shape per se. Bowing to convention, we use the term "inconspicuous" as shape descriptor, but we also recognize that all such flowers are very small or small, where shape is essentially not a consideration.

Inconspicuous and dish-shaped flowers are widespread among the angiosperms of the archipelago, constituting 41% and 35% of the species, respectively. Tubular flowers characterize 15% of the species, and bell-shaped flowers only 6%. Finally, species with flag- (2%) and brush-shaped (1%) flowers are rare.

4. Flower Color

Green is the most widespread flower color among the species (41%). White and yellow flowers follow in importance, with 26% and 12% of the species, respectively. The remaining colors are present in much smaller percentages (red 9%, brown 7%, and blue 5%).

5. Sex

As expected, the majority of the species on the archipelago are hermaphroditic (about 70%). Some 9% of the species are dioecious, and 9% monoecious. Dioecious species are included in the dicot genera Coprosma, Fagara, Pernettya, and Robinsonia; Juania is the only dioecious monocot in the archipelago. Monoecious dicots are in the genera Boehmeria, Dysopsis, and Urtica, and there are several monoecious Cyperaceae. The remaining sexual systems are found exclusively among the dicots: Gynomonoecious species include 7% of the flora and are in Abrotanella, Cuminia, Chenopodium, Erigeron, and Lactoris; the one gynodioecious species is in Rhaphithamnus; polygamous taxa are scarce and included in Empetrum and Parietaria, as are andromonoecious taxa included in Cyperaceae and Asteraceae.

6. Separation of the Sexes

Because field studies of the floral biology are particularly meager, data on such characteristics as dichogamy and herkogamy are very limited. The combination of our direct observations and Skottsberg's observations (1928), together with generalizations from published data for some continental genera or families, indicate that around 30% of the species are or should be protandrous and 7% protogynous. Although these data are not complete, they do show that temporal separation of the sexes is not uncommon. Protandry was found mainly among members of Asteraceae, Campanulaceae, and Gunneraceae, with protogyny among the genera Drimys, Eryngium, Loctoris, Myrceugenia, and Plantago. The only orchid known in the archipelago, which grows on Alejandro Selkirk Island, is herkogamous.

7. Compatibility

Only about 14% (21 species) of the flora has been studied sufficiently to determine compatibility. Some 85% of the species studied (18) are SC, and only three are SI.

8. Floral Rewards

This information is also not exhaustive. Our direct observations (Bernardello et al., 2000; Anderson et al., 200la), combined with our general knowledge of related South American genera, indicate that at least 55% of the species do or are likely to offer nectar but that only about 2% offer pollen as a reward.

9. Floral Visitors

Overall, floral visitors are rare to uncommon. Combined, our direct observations and those of Skottsberg (1928) cover about 35% of the species. These data indicate that 29 species (54% of those studied) never had any floral visitors, even though those species have been carefully observed in the field for considerable periods of time. Both sexes of two hummingbird species (Sephanoides fernandensis, an endemic, and S. sephaniodes, native, but also found in Chile south of the Atacama Desert [Roy et al., 1998]) were recorded as regular visitors for 14 species of the total flora. We presume that they are regular pollinators for these, chiefly woody species in the genera Centaurodendron, Cuminia, Dendroseris, Eryngium, Escallonia, Greigia, Lobelia, Nicotiana, Notanthera, Ochagavia, Rhaphithamnus, and Sophora. Skottsberg (1928) reported pollen from three of those species (Rhaphithamnus venustus, Cuminia eriantha, Escallonia callcottiae) on the heads of captured hummingbirds.

Native insects (flies, moths, and beetles) have only rarely been recorded on flowers. Sporadic insect visitors were recorded for only 7% of the total flora; that is, 11 species in nine genera. Decreasing this total even further is the fact that two of the species visited by native insects, Escallonia callcottiae and Eryngium bupleuroides, are hummingbird pollinated. Furthermore, the insect visits we have documented only very doubtfully constitute effective pollination because of the casual nature of the visits and the lack of fidelity of the visitors (Anderson et al., 200la). More recent discoveries indicate that there are, in addition, an introduced ant (Linepithema humile, E. O. Wilson, pers. comm.; Anderson et al., 2001a) and an (doubtfully) endemic bee in the genus Lasioglossum (Anderson et al., 2000b; Engel, 2000.). Neither the bee nor the ants are yet important to the reproduction of the native flora.

10. Pollination

From the data above, we can confidently conclude that about 9% of the total flora is bird pollinated. Given the length of study and the conspicuousness of these pollinators, we have reasonable confidence that this is, indeed, an accurate figure for the flora in toto. On the other hand, given the scant, irregular, and imprecise nature of native insect visitation, there is no certainty that any of the plant species any truly insect pollinated (with the possible exception of Libertia; see below). Based on floral morphology, floral biology, compatibility, and effective seed set in the absence of pollinators, we conclude that about 47% of the flora is likely wind pollinated. The pollination for the remaining 44% of the flora is unknown. Among the anemophilous species, some have most of the typical features, not surprisingly among the Chenopodiaceae, Cyperaceae, Juncaceae, Poaceae, Urticaceae, and the genera Coprosma, Dysopsis, Empetrum, Gunnera, Haloragis, Peperomia, Plantago, and Lactoris (Bernardello et al., 199 9). However, other species show combinations of adaptations to wind pollination and other features that would indicate biotic pollination (e.g., Azara, Drimys, Pernettya, and Robinsonia; Anderson et al., 2000a). A special case of wind as an indirect force of pollen transfer was found in Wahienbergia berteroi (Anderson et al., 2000b). A summary of the pollination is presented in section V, Discussion.

11. Hypothesized Pollination of Colonizers

Based on the pollination biology known for continental relatives, we estimate that about half of the colonizer species were primarily insect pollinated. Wind-pollinated colonizers likely constituted a comparatively high proportion as well (40%), whereas bird-pollinated colonizers and colonizers with mixed pollination account for much lower percentages (7% and 4%, respectively).

B. COMPARISONS OF HABIT AND FLORAL FEATURES

1. Habit and Flower Size

Species that are perennial herbs usually have diminutive flowers (70% very small, p <0.0001, Table II; 14% small). Tree species are at the extremes, in that they have mostly very large (36%) or very small (21%) flowers, but all size classes are represented. Other species with a perennial habit have mainly medium-sized flowers (62%).

2. Sex and Habit

Hermaphroditic species show all of the possible habits. Monoecious species are mostly perennial herbs or trees; all dioecious species are woody.

3. Sex and Flower Color

Most species are hermaphroditic, and there is not a dominant color, although the relatively dull white (38%, p = 0.0003, Table II) and green (31%) hermaphroditic flowers are the most frequent. More than 93% of the monoecious taxa have green flowers (p<0.0001, Table II), and green and yellow flowers are predominant in both gynomonoecious and dioecious species.

4. Flower Shape and Flower Color

Species with inconspicuous flowers possess predominantly dull colors: 71% are green (p <0.0001, Table II), and 10% are brown (Fig. 1A). Species with dish-shaped and tubular flowers show the full range of colors, but in both types white is most common (36% and 32%, respectively, Fig. 1A). Bell-shaped flowers are also most often white (85%, p 0.0005, Table II); a few species have either red or green flowers (Fig. 1A).

5. Flower Size and Flower Color

Most of the species (Fig. 1B) with very small flowers have green flowers (67%, p<0.000 1, Table II). Similarly, small-flowered species generally are not brightly colored: 56% are white, 19% brown, and 19% green (Fig. 1B). The species with very large flowers (Fig. 1B) are exclusively white (66%) or red (33%). The species with medium-sized flowers show the full range of colors, with white (41%) and yellow (33%) most common (Fig. 1B).

6. Flower Size and Sex

Hermaphroditic species show all of the possible flower sizes: About half (45%) have very small, 16% small, and 18% medium-sized flowers. Monoecious taxa are characterized almost exclusively (92%) by very small flowers. Dioecious species possess mainly medium-sized flowers (84%), and most gynomonoecious species bear medium-sized (55%) or very small (36%) flowers.

7. Flower Shape and Size

Not surprisingly, all of the substantial percentage (41%) of the flora (Fig. 2) that is characterized by the "inconspicuous" shape has flowers that are either very small (94%, p<0.0001, Table II) or small (6%). Species with dish-shaped flowers are commonly medium-sized (44%, Fig. 2). Species with tubular flowers have flowers more or less equally distributed by size: small (37%), medium-sized (37%), or large (21%, Fig. 2).

8. Flower Shape and Sex

Given that hermaphroditic flowered species are the majority in the flora, it is also not surprising that their flowers show all of the possible shapes. Similarly, species with inconspicuous and dish-shaped flowers are distributed over all of the sexual systems. Monoecious species bear primarily inconspicuous flowers (85%), but dioecious species generally possess dish-shaped flowers (61%).

C. COMPARISONS OF CURRENT POLLINATION WITH HABIT AND FLORAL TRAITS

We have good data on pollination systems for about 55% of the flora, and it is with these species that we make the following comparisons. As indicated previously, most of the species for which we have good information are either wind (47%) or bird (9%) pollinated. Only Libertia may be reliably insect pollinated.

1. Habit

All of the bird-pollinated species are perennial, ordinarily woody (shrubs and trees). Wind-pollinated species are perennial; the majority are herbs (60%, p = 0.0002, Table II).

2. Flower Size

The flowers of the bird-pollinated species are comparatively larger (large 46%, medium-sized 38%, very large 15%, Fig. 3A); when combined, large and medium-sized flowers proved to be associated with bird pollination (p <0.0001, Table II). On the contrary, most wind-pollinated species have very small (70%, p <0.0001, Table II) or small flowers (14%, Fig. 3A); both categories pooled were associated with wind pollination (p <0.0001, Table II).

3. Flower Shape

The floral features of most species show adaptations coincident with their pollination system (Fig. 3B). Thus, most bird-pollinated species have tubular flowers (57%). A substantial percentage (28%) are also dish-shaped flowers; however, these are all Asteraceae and, as required in Faegri and van der Pijl's classification (1979), are considered "dish" shaped, although each individual flower is obviously a diminutive tube. As would be predicted, wind-pollinated species have chiefly inconspicuous flowers (7l%,p <0.0001, Table II). However, there are also species with flower shapes that are not typically associated with wind pollination: bell (16%), tube (7%), or dish (5%).

4. Flower Color

Bird-pollinated species regularly have flowers with bright colors (red, orange, yellow, and mainly blue [36%]; Fig. 4A, colors combined: p <0.0001, Table II). On the other hand, flowers of wind-pollinated species are usually not showy (Fig. 4A), green (50%, p = 0.01, Table II) or brown (32%); both colors combined were associated with wind pollination (p <0.0001, Table II).

5. Sex

Some 80% of the bird-pollinated and 68% of the wind-pollinated species bear hermaphroditic flowers (Fig. 4B). The other sex forms are largely wind pollinated (Fig. 4B).

D. COMPARISONS OF THE HYPOTHESIZED POLLINATION OF COLONIZERS WITH HABIT AND FLORAL FEATURES

The following are obviously speculative, but the generalizations are informative in that they establish the likely evolutionary challenges faced by the colonizing species.

1. Habit

No typical habit seems to have been typical of insect-pollinated colonizers, but bird-pollinated colonizers likely had a perennial habit (from perennial herbs to trees). Within the species that are perennial herbs and shrubs, all pollination types were represented. Perennial herbs seem to have been primarily wind pollinated (53%) or insect pollinated (40%). Shrubs, however, were mostly (55%) insect pollinated, with no other class particularly strongly represented. Trees were mostly wind pollinated (60%, p = 0.03, Table II) or insect pollinated (37%).

2. Flower Size

The full range of flower sizes is found in insect-pollinated colonizers, although most would have been medium-sized or very small flowers (30% and 24%, respectively, Fig. 5A). As expected, very small flowers were typical of wind-pollinated (73%, p <0.0001, Table II) colonizers; large (50%) or medium-sized (33%) flowers were common in bird-pollinated colonizers (Fig. 5A).

3. Flower Shape

As expected, tubular flowers characterized all bird-pollinated colonizers, and inconspicuous flowers were common in almost all wind-pollinated species (85%, p <0.0001, Table II, Fig. 5B). The most frequent flower shapes among insect-pollinated colonizers were dish (50%, p <0.0001, Table II) and bell (20%, Fig. 5B).

4. Flower Color

The full range of floral colors was represented in insect-pollinated colonizers, with a predominance of white (56%, p <0.0001, Table II) or green (22%) colors (Fig. 6). As one might predict, showier flowers (blue 60%, red 20%, p <0.0001 for both colors combined, Table II) were typical of bird-pollinated colonizers (Fig. 6). The species with less showy green (82%, p <0.0001, Table II) or brown (9%) flowers were common in wind-pollinated colonizers (Fig. 6).

5. Sex

Given the sex distribution of the primary source flora, it is not surprising that hermaphroditic flowered species were the most common in the different pollination systems (78% of the insect-, 74% of the wind-, and 75% of the bird-pollinated species). However, dioecious colonizers seem to have been primarily insect pollinated (80%) or wind pollinated (20%). Monoecious colonizers are proposed as insect or wind pollinated (50% each), as are the few gynomonoecious species (40% insect and 40% wind).

E. CORRELATION BETWEEN CURRENT POLLINATION AND HYPOTHESIZED POLLINATION OF COLONIZERS

We assessed feasible changes in the pollination system for 39 genera (Table I), based on pollination system determination from our current studies together with our hypotheses of possible colonizers (see our discussion in the section above). About 70% of the genera seem to have retained the pollination system of the colonizers. Those that retained wind pollination include members of several monocot (Cyperaceae, Juncaceae, and Poaceae) and dicot families (Chenopodiaceae, Empetraceae, Euphorbiaceae [Fig. 7C], Gunneraceae, Haloragaceae, Urticaceae [Fig. 7D], Piperaceae, Plantaginaceae [Fig. 7A], Rubiaceae [Fig. 7B]). We propose the retention of hummingbird pollination for the Bromeliaceae (Greigia. Ochagavia), Campanulaceae (Lobelia), Fabaceae (Sophora), Lamiaceae (Cuminia [Fig. 8C]), Loranthaceae (Nothanthera), and Saxifragaceae (Escallonia [Fig. 8A]). The only species that seems to have retained insect pollination is Libertia.

At least some of the species of the other 30% of the genera bear a pollination system different from that of their presumed colonizers. For four genera, we suggest that the ancestral insect-pollination system switched to hummingbird pollination (Rhaphithamnus venustus, some Eryngium, Centaurodendron, and some Dendroseris [Fig. 8B]). Nicotiana cordifolia is now hummingbird pollinated, but the first colonizers could have been insect and/or hummingbird, so it is not included on the list of the species with changes. In addition, for seven other genera we suggest that the colonizers were likely insect pollinated and that on the islands they have become, of necessity, wind pollinated (Azara, Drimys [Fig. 9B], Juania, Pernettya, Robinsonia, Ugni [Fig. 9A]). Solanum species (Fig. 10B) may have changed from insect pollinated to pure autogamy, but autogamy is not uncommon in this genus, so this failsafe system may even have facilitated establishment of colonizers. In Wahlenbergia (Fig. 10A), we suggest that the ancesto rs were mainly insect pollinated, with a secondary system of autogamy, but that on the islands, and in the absence of insects, the selfing mechanism became the rule. The wind-aided mechanism of autogamy of W. berteroi must have arisen on the archipelago.

V. Discussion

A. HABIT

Like island systems as a whole (e.g., Darwin, 1859; Carlquist, 1965, 1974; Bramwell, 1972; Wagner et al., 1990), the Juan Fernandez Archipelago includes an inordinate number of perennial forms. There are shrubs/simple trees, for instance, in several Asteraceae endemic genera (e.g., Centaurodendron, Dendroseris, Robinsonia, Yunquea) and in individual species of Erigeron and Plantago whose presumed ancestors and relatives are smaller, herbaceous forms (Carlquist, 1965, 1974; Sanders et al., 1987). Molecular studies have provided confirmation of the evolution of arborescence in several plant groups from Hawaii (Wagner et al., 1990; Givnish et al., 1995; Baldwin, 1997; Sakai et al., 1997; Givnish, 1998) and the Canary Islands (Bohle et al., 1996; Kim et al., 1996; Francisco-Ortega et al., 1997; Panero et al., 1999).

Many hypotheses have been proposed to explain the preponderance of the woody habit, some with implications for reproductive biology. Wallace (1878) suggested that longevity (perennial habit) increased the opportunity for successful sexual reproduction when pollinators were scarce--as they are on some islands. Pollinators, other than hummingbirds, are certainly uncommon at best on the Juan Fernandez Islands. Bohle et al. (1996) went farther, claiming that Echium species from islands off the coast of Africa evolved the woody habit (from herbaceous perennial continental ancestors) to promote not just sexual reproduction but outcrossing--to reduce the inbreeding depression associated with the selfing of the initial colonizers. They also suggested that a secondary manifestation of outcrossing was rapid and extensive speciation on the islands. This latter pattern is perhaps foretold in the chromosomal races of the house mouse established on Madeira as well (Britton-Davidian et al., 2000).

B. FLORAL FEATURES

Angiosperms from oceanic islands are generally characterized by small, inconspicuous (recall = shape, "with no optical attraction") flowers versus showy, bright flowers with various shapes (e.g., Wallace, 1895; Carlquist, 1965, 1974), a trend particularly remarkable in the New Zealand flora (Godley, 1979; Lloyd, 1985; Webb & Kelly, 1993). The Juan Fernandez flora provides strong support for these generalizations: 60% of the flowers are very small/ small (and thus without "shape") and the most frequent flower colors are comparatively dull (green and then white, in decreasing order of frequency). Furthermore, this study, done species by species for the entire archipelago flora, is the first to statistically confirm the expected strong association among very small, inconspicuous, and green flowers. Finally, comparisons with the hypothesized colonizer flora imply that there has not been selection for change in flower color or size (selection neutral). This latter hypothesis implies that the small, inconspicuous f lowers characteristic of islands may better reflect selection for dispersal and establishment than adaptation in situ.

The distribution of flower colors on other islands (e.g., the Galapagos; McMullen, 1989, 1999) is comparable. This common pattern could be explained in several ways. For whatever reason, species with small green and white flowers may possess greater dispersability, or higher levels of SC. At least for the Juan Fernandez Archipelago, these species do not show a higher percentage of SC than do other elements of the flora. Another explanation is that these species with simple, nonshowy flowers may succeed by being serviced by a broader spectrum of pollinators. This hypothesis fails on the Juan Fernandez Islands because of the virtual lack of insect pollinators. Finally, species with these less distinctive flowers may be abiotically pollinated, a hypothesis supported by the high percentage of wind-pollinated species in the flora.

C. SEX

The bulk of the flora has hermaphroditic flowers (ca. 70%). Thus, as would be expected, of this majority condition among angiosperms, the island flora exhibits the range of habits and flower colors, sizes, and shapes. There are about twice as many monoecious (9%) and dioecious (9%) species in the Juan Fernandez Archipelago as there are in angiosperms as a whole (5% and 4% for monoecy and dioecy, respectively; Richards, 1997).

The Fernandezian monoecious species--included in the Euphorbiaceae, Urticaceae, and Cyperaceae--are largely herbaceous and have mostly very small, inconspicuous, green flowers. Because these families in general include many monoecious species (Cronquist, 1981), it is likely that the colonists in those groups were also monoecious. Most of these monoecious species are, and presumably were, wind pollinated--again a general feature of monoecious species (Richards, 1997).

The highest levels of dioecy among flowering plants are found in oceanic islands (Carlquist, 1974; Richards, 1997). Although dioecy in the Juan Fernandez flora is twice the world average, it is not as high as in some other islands, such as New Zealand or Hawaii (Godley, 1979; Sakai et al., 1995a, 1995b, respectively). Furthermore, Arroyo and Uslar (1993) determined that about 9% of the sclerophyllous montane flora of central Chile is dioecious. Thus, the frequency of dioecy in the Juan Fernandez Archipelago is similar to that in a latitudinally comparable flora--and the presumed main continental source. This association characterizes other island systems as well (cf. Baker & Cox, 1984). All of the Fernandezian dioecious species are woody, and most are shrubs. Thus, dioecy is related to woodiness, as it has been in other floras (e.g., Baker, 1959; Bawa, 1980; Sakai et al., 1995b). This association is attributed to strong selection for outcrossing in large, long-lived plants that might otherwise self-cross (cf. Bawa, 1980). Dioecy also is not equally distributed among the various habits in the continental source flora in Chile (Arroyo & Uslar, 1993). Some 62% of the dioecious species are long-lived, perennial forms, and 17% of the dioecious perennials are shrubs.

In addition, dioecy is overrepresented on the Juan Fernandez Islands in species with small green flowers, a pattern also reported in Hawaii (Sakai et al., 1995a, 1 995b). Such flowers are often abiotically pollinated, as is true for all of the Fernandezian dioecious species, and for many other floras as well (e.g., Conn et al., 1980; Freeman et al., 1980; Muenchow, 1987; Steiner, 1988), including that of the Hawaiian Islands (Sakai et al., 1995b).

As indicated above, most of the dioecious species likely had dioecious ancestors. Almost all of the species of Coprosma known worldwide are dioecious (Oliver, 1935). Thus, we presume that the original colonists in this genus were dioecious as well. The island progenitors of Juania and Fagara may have been, at least, incipiently dioecious or have a clear tendency to it, because there are dioecious Fagara species in Hawaii (Wagner et al., 1990), and there are many dioecious palms (Cronquist, 1981). In two genera, it seems likely that dioecy arose in situ. The dioecy in Robinsonia may have had SI ancestors (Crawford et al., 1998; Anderson et al., 200 la). Pernettya rigida is the only cryptically dioecious species known so far in the islands. Because of this, careful anatomical, morphological, and experimental studies were required to understand that the flowers of P. rigida, though complete, are functionally dioecious (Anderson et al., 2000a). Most continental species of Pernettya bear hermaphroditic flowers, bu t some have unisexual flowers (Arroyo & Squeo, 1987; Anderson et al., 2000a). Thus, the first colonists of Pernettya may have been hermaphroditic, but perhaps with incipient dioecy expressed (Anderson et al., 2000a).

About 7% of the flora is gynomonoecious, with representatives from the Asteraceae, Chenopodiaceae, Cuminia (Lamiaceae), and Lactoris (Lactoridaceae). A review of the sex distribution in the first two families indicates that gynomonoecy seems likely to have been present in the colonizers. On the other hand, in Cuminia, the variability of sex expressions both we and Skottsberg (1928) observed makes it more likely that the gynomonoecy arose in situ. Lactoris is a monotypic endemic with debatable relationships (Stuessy et al., 1998a), so it is impossible to confidently project its colonist relatives.

There are few examples of gynodioecy, polygamy, and andromonoecy in this archipelago. The only species studied carefully is Rhaphithamnus venustus (Sun et al., 1996; Anderson et al., 200 la). The gynodioecy in this species seems to have arisen in situ and to be incipient, given the variability in sex expression (e.g., fertile and infertile anthers are similar).

D. BREEDING SYSTEM

"Baker's law" (Baker, 1955, 1967; Stebbins, 1957) proposes that the species most likely to become established after long-distance dispersal will be SC. In addition, the subsequent acquisition of self-pollination would favor establishment (Baker, 1955, 1967; Carlquist, 1974;

Ehrendorfer, 1979; Barrett, 1998). Even though comprehensive surveys of the compatibility status of island plants are surprisingly few (Barrett, 1998), the data that do exist support this hypothesis (e.g., Stephens, 1964; Rick, 1966; Gillett & Lim, 1970; Strid, 1970; Carlquist, 1974; Kores, 1979; Pandey, 1979; Roelofs, 1979; Rabakonandrianina, 1980; Cory, 1984; Lowrey & Crawford, 1985; McMullen, 1987, 1990; Webb & Kelly, 1993). More than 85% of the Fernandezian angiosperms studied carefully are SC (Bernardello et al., 1999; Anderson et al., 2001a). Thus, although SC is likely frequent in the Juan Fernandez Archipelago, the level of autogamy (automatic selfing within the same flower) seems to be quite low, at least in comparison with the Galapagos flora, where 26 of the 29 native and endemic species examined experimentally showed some degree of automatic self-pollination (McMullen, 1987, 1990). The lower rate of autogamy on the Juan Fernandez Islands may be attributable to the significant percentage of species that have temporal separation of the sexes through dichogamy (mostly protandry). A significant level of dichogamy also characterizes the New Zealand flora (Godley, 1979). Nonetheless, self-crossing is common, but in the form of geitonogamy (pollen transfer among flowers of the same individual). Geitonogamy is the most frequent mechanism of pollen transfer among the cosexual Fernandezian species we studied experimentally. And, in general, geitonogamy is probably the most widespread mode of self-pollination, virtually inevitable in SC plants that produce a number of flowers open at the same time (Lloyd & Schoen, 1992). In the Faroes Islands, Hagerup (1951) indicated that species with large inflorescences tended to be geitonogamous. However, Lloyd (1992) also pointed out that most cosexual SC species may be unable to avoid mixed mating--as would most likely occur in all of the SC hummingbird-pollinated species on the Juan Fernandez Islands.

In some Wahlenbergia taxa (W. fernandeziana and a natural hybrid between this species and W. grahamiae; Anderson et al., 2000b), autogamy perhaps provides an alternative to the presumed entomophilous pollination system of colonizers. Features of the Fernandezian Wahlenbergia, like protandry, nectar, and secondary pollen presentation (by pollen-collecting hairs on the style), reduce self-pollination and suggest ancestral allogamy and biotic pollination. The pollination biology of Wahlenbergia elsewhere (Lloyd & Yates, 1982; Petterson, 1997) also supports these suppositions. Autogamy is implicated by a very low P/O ratio and is promoted by SC and partial overlapping of male and female phases at the end of each flower's lifetime and by recurvature of stigmatic branches all the way either back to the style or to the inner corolla surface.

Three SI species are confirmed so far for the Juan Fernandez Islands (Anderson et al., 2001a). In other island systems, a few SI species were found as well (e.g., Carpenter, 1976; Corn, 1979; Grant & Grant, 1981; Carr et al., 1986). At least for two of the SI Fernandezian taxa, the Dendroseris spp., the ancestors are proposed to have been SI, based on the ubiquity of this condition among its extant closest relatives (Crawford et al., 1998; Anderson et al., 200la).

E. REWARD

Few species seem to offer pollen as a reward, but a significant number produce nectar (Skottsberg, 1928; Bernardello et al., 2000; Anderson et al., 2001a). However, with the exception of the two species of hummingbirds, there are virtually no floral visitors to collect this reward.

In general, nectary morphology of individual species follows the general pattern of the respective families, suggesting that nectaries came with the colonizers (Bernardello et al., 2000). In the small number of species in the archipelago that are hummingbird visited, nectar chemistry implies specialization for birds, again suggesting that the colonists were also ornithophilous. Likewise, the presence of nectaries and nectar in most non-hummingbird-pollinated species is most likely an indication of the pollination system of the first colonizers. Given the virtual lack of dedicated insect pollinators, the retention (or evolution of) nectar would not seem to reflect the extant biota. Particularly good examples of retention of elements of an ancestral system are the wind-pollinated, nectar-producing species, such as Pernettya rigida, Wahlenbergia berteroi, and the species of Robinsonia. The ancestors of P. rigida seem to have been insect pollinated, but today this cryptically dioecious species is wind pollinated, and visitors are extremely rare and ineffective (Anderson et al., 2000a). Wahlenbergia colonizers seem to have been entomophilous as well, but current pollination is largely autogamous (Anderson et al., 2000b). The situation in Robinsonia is not well understood. Their capitula are yellow/green and their flowers likely bear nectar, but visitors are particularly rare. In 1896, Johow reported flies visiting the genus, without citing the species. Skottsberg (1928) recorded no visitors, and in many hours of field observation in several expeditions, we never recorded any floral visitors either. Thus, given reasonable fruit set in the field, we postulated wind pollination (Anderson et al., 2001a), considering that shifts between insect and wind pollination have been reported to occur several times in this large family (Stebbins, 1970). Of course, fruits may be set apomictically as well, but at present we have no evidence of this.

F. VISITORS

In general, islands possess comparatively fewer animal species than do their source continents (MacArthur & Wilson, 1967). In consequence, their pollinator faunas are often smaller, with many groups completely absent (Carlquist, 1974; Woodell, 1979; McMullen, 1990, 1999; Inoue, 1993). The Juan Fernandez Islands can now become the exemplar of this pattern: Other than two species of hummingbird, floral visitors are absent or rare (Johow, 1896; Skottsberg, 1928; Bernardello et al., 1999; Anderson et al., 2001a). This small archipelago is notable because of the existence of the two hummingbird species, highlighted because one of them is the only endemic known on oceanic islands (Brooke, 1987; Meza, 1988; Colwell, 1989; Roy et al., 1998). The diet of the hummingbirds includes nectar from a total of 14 autochthonous plant species (Brooke, 1987; Meza, 1988; Colwell, 1989; Bernardello et al., 2000; Anderson et al., 2001a). In general, our analyses show that the ornithophilous species are woody and have tubular flower s with significant amounts of nectar. The chemical nectar composition of five of these species (Bernardello et al., 2000) indicates that they have a low concentration and high sucrose proportion, features common to the nectar of hummingbird-pollinated species (Pyke & Waser, 1981; Cruden et al., 1983; Baker & Baker, 1983a, 1983b, 1990). Rhaphithamnus venustus is the "most visited" ornithophilous endemic species (Brooke, 1987; Meza, 1988; Colwell, 1989; Sun et al., 1996; Anderson et al., 2001a). This species has a long flowering period and produces a large number of flowers with abundant, mostly sucrose-dominant nectar (Bernardello et al., 2000). In addition to the indigenous flora, these hummingbirds also visit flowers and feed on nectar from cultivated and adventitious species (Brooke, 1987; Meza, 1988; Colwell, 1989; Bernardello et al., 2000; Anderson et al., 2001a).

The Fernandezian insect fauna is small in general (cf. Johow, 1896; Kuschel, 1952; Wilson, 1973) and notably lacks species usually dedicated to floral visits (Skottsberg, 1928; Bernardello et al., 1999; Anderson et al., 2000a, 2000b). Consequently, it is not surprising that the records of insect visits to flowers are unusual. For instance, a few insects (beetles and moths) were observed on Berberis, Colletia, and Robinsonia species, either at the end of the nineteenth century (Johow, 1896) or early in the twentieth (Skottsberg, 1928); since then, there are no other records. It is worth mentioning that Skottsberg (1928) spent a total of at least six months in the field and recorded insects on only seven plant species (Table I). Although he gave no indication of the number of individual insects observed for each, based on his lack of citation of large numbers in an extensive text and the current dearth of flower visitors, we presume floral visitors have been rare. In more than 300 hours of observation in three field expeditions, we recorded only 23 floral visits by native insects. Furthermore, and with one exception (see below) the insects recorded are not specialists and, in general, should not be considered pollinators. Evidence of this comes from their behavior and from the absence or small amount of pollen on their bodies (Anderson et al., 2001a). They visit stamens, nectaries, corollas, leaves, other plant parts, and inanimate objects with equal interest and have very low or no fidelity to flowers in general (Anderson et al., 2000a, 2000b, 2001a). A possible exception is a newly described halictid bee species (Lasioglossum subgenus Dialictus; Engel, 2000). For the moment, this bee is a new endemic to the Robinson Crusoe Island. However, it is more than likely that it is a relatively new adventive to the island and that it represents an as yet uncollected and perhaps uncommon new continental species, rather than a continental waif that arrived on the island and evolved in situ (Anderson et al., 2001b).

Much more abundant were the ants we collected in the flowers on some endemics. Ant specialist B. O. Wilson identified the ants as a recently introduced, highly invasive species from South America (Linepithema humile, "Argentine ants"). Because of their presumed relatively recent introduction, their hard and small bodies, and the lack of directed movement among the flowers, we consider that they are not yet significant in transferring pollen and, more importantly, played no role in the evolution of the flora or its pollination.

G. CURRENT POLLINATION

Carl Skottsberg was the first to carefully study pollination biology in the context of his three-year study of the floristics of the Femandezian plants (1928). Nevertheless, many of his conclusions were inferred from flower morphology and pollination of closest continental relatives rather than from actual observations. For instance, of the 49 species he reported as entomophilous (1928), he actually observed insects on flowers for only 7. The species he proposed as anemophilous were based largely on knowledge of families from the continent. For the ornithophilous species, he did observe hummingbirds visiting flowers for 9 species, including, for 3 species, the observation of pollen on the faces of collected birds. Since Skottsberg's work (1928), there was virtually no study of the reproductive biology until very recently (brief comments as part of some broader study include Crawford et al., 1990; Ricci & Eaton, 1994; focused studies include Sun et al., 1996; Bemardello et al., 1999, 2000; Anderson et al., 200 0a, 2000b, 2001a).

1. Hummingbird Pollination

The Juan Fernandez Islands are notable for having hummingbird pollination. The species that hummingbirds visit have abundant nectar and possess a floral structure suited to bird pollination (Bernardello et al., 2000; Anderson et al., 2001a). Wallace (1878) suggested that the large and showy flowers on Robinson Crusoe Island are an evolutionary consequence of dependence on hummingbirds. He was probably correct in his observation, but not in his explanation. Given that the percentage of bird pollination is about the same in today's flora as we estimated for the first colonists, we suggest that very few species switched from some other pollination system to ornithophily. Rather, the proportion of bird-pollinated species represents the successful establishment of ornithophilous colonists. Certainly, the association of hummingbirds with flowers is long-standing; in their study of the origin of plant-animal mutualisms in the woody flora of the temperate forest of southern South America, Aizen and Ezeurra (1998) con cluded that the evolution of ornithophily appears to have taken place well before the climatic cooling and biogeographical isolation of that region in the Tertiary. Thus, as might be expected, most of the ornithophilous families or genera include hummingbird-pollinated species elsewhere. There is good information that some members of Bromeliaceae (Bernardello et al., 1991), Campanulaceae (Galetto et al., 1993), Loranthaceae (Smith Ramirez, 1993; Armesto et al., 1996; Rivera et al., 1996), and Saxifragaceae (Armesto et al., 1996), as well as of the genera Nicotiana (Hernandez, 1981), Rhaphithamnus (Smith Ramirez, 1993; Sun et al., 1996), and Sophora (Arroyo, 1981), are visited by hummingbirds or other kinds of birds in southern South America.

We do not know the timing of the arrival of either plants or hummingbirds, nor whether both may have arrived several times. This is, approximately, a kind of "chicken-and-egg" conundrum. Colwell (1989) considers these plant species unlikely to have been successful colonists of these islands without the presence of hummingbirds. But, without the bird flowers and their large supply of nectar, what provided nutrition for the hummingbirds? The hummingbird-pollinated species are perennial, mostly woody, and SC: Thus, at least some of them could have reproduced to some limited extent before the hummingbirds arrived.

However, even SC does not ensure reproduction. Some of the SC ornithophilous species (the gynomonoecious, gynodioecious, and dichogamous species) require at least interflower pollen transfer. In many of these species, the abundance of simultaneous flowers, the amount of nectar exuded, and the hummingbird behavior would favor pollen transfer among flowers of the same plant (geitonogamy) over interplant pollen transfer (xenogamy). There are also some ornithophilous species that seem to have involved shifts from insect pollination of the first colonizers to bird pollination. Some hummingbird-pollinated members of the genera Centaurodendron and Dendroseris (Asteraceae), Eryngium (Apiaceae), and Rhaphithamnus (Verbenaceae) have comparatively larger flowers (Skottsberg, 1957; Sanders et al., 1987; Skottsberg, 1953; Sun et al., 1996, respectively) than do island congeners of those genera pollinated otherwise (or whose pollination is doubtful). Thus, in these instances, flower length may have increased due to a speci alization to hummingbird pollination. Sun et al. (1996) suggested that in Rhaphithamnus venustus it is likely that ornithophily arose as a result of flower elongation from the only entomophilous continental species of the genus, R. spinosus. However, given that hummingbirds occasionally visit the Argentine and Chilean R. spinosus (Sun et al., 1996), the colonizing ancestors of R. venustus may have been preadapted to ornithophily.

2. Wind Pollination

Wind is considered a significant pollination agent for remote island floras (Carlquist, 1966; Whitehead, 1969, 1983; Regal, 1982; Barrett, 1998), with anemophily characterizing significant percentages of species in island floras (Thornton, 1971; Carlquist, 1974; Ehrendorfer, 1979). Indeed, wind pollination is important in the Juan Fernandez Archipelago, as reported here and in our previous studies (Bernardello et al., 1999; Anderson et al., 2000a, 2000b, 2001a). In addition to some experimental studies, we propose anemophily as important, based on extensive and intensive studies demonstrating the absence of significant native-insect-pollinating groups and the consequent nearly complete lack of flower visiting insects.

For a portion of the flora, anemophily is not a surprise, given the condition among presumed ancestors. For a significant proportion of other species, we propose that the lack of any other alternative means of pollen transfer has led to a dependence on dispersal of pollen by wind. Similar arguments have been made for elements of the continental flora isolated in habitats with little insect-pollinator service (e.g., Berry & Calvo, 1989). Hagerup (1932, 1951) has reported instances in which the rarity or inactivity of insects seemed to result in selfing, anemophily, or even rain pollination. A number of the Fernandezian anemophilous species lack the full suite of characters (cf., Regal, 1982; Whitehead, 1983) typically associated with wind pollination. Thus, we conclude that floral morphology does not always give a precise indication of the pollination mode in this evolutionary young flora. Instead, the floral features many times seem to reflect retention of the traits of the reproductive systems of the origina l colonizers. These features may be expressed because there has been relatively little time for evolution to modify the ancestral characters or because there has been little or no selection against the ancestral traits (Anderson et al., 2001a). The latter argument may be the stronger, given the observations by Britton-Davidian et al. (2000) that selection (at least for some features) can act quickly: At least six different chromosomal races of the house mouse were established on Madeira in 400 years.

For another portion of the flora, anemophily is not so clearly expected. Few species colonize islands, so that successful colonizers may move into niches new to them, both ecologically and evolutionarily. In the new environment and with fewer competitors (vis-a-vis continents), unusual forms (e.g., the habit of Dendroseris on the Juan Fernandez Islands) may be expressed (e.g., Carlquist, 1974), or ancestral conditions may persist because of the lack of selection against them. As a consequence, wind pollination of certain species may be obscured by retention of ancestral entomophilous features. Thus, detailed study of individual species may be required to reveal the actual pollination system. More extensive studies of the pollination biology of island floras may reveal anemophily for species in which the morphology would not have predicted it (Anderson et al., 2001 a). In the Galapagos, for instance, McMullen (1987, 1990) indicated that floral morphology conductive to wind pollination is rarely encountered (al so see Barrett, 1998). A subsequent study (McMullen & Close, 1993), which analyzed the amount of airborne pollen produced by six species, suggested that it was not of a magnitude sufficient for wind pollination. However, detailed study of the reproductive biology of the Galapagos has only included a small fraction (10-15%) of the native flora, and subsequent analysis may reveal anemophily, a pollination system operating perhaps by default, to be more prominent than flower morphology would imply.

An interesting and extreme example that shows the impact of wind transferring pollen was detected in Wahlenbergia berteroi (Anderson et al., 2000b), a derived species (Lammers, 1996) that produces nectar but is not visited by hummingbirds or pollinating insects. In the other Wahlenbergia species of the archipelago, autogamy occurs at the end of the flower lifetime, when the stigmata branches recurve almost 360[degrees] and contact pollen held on the style. However, in W. berteroi the degree of stigmatic recurvature is insufficient to reach the style and facilitate autogamy. Nevertheless, in the field most of the flowers observed did have pollen on the stigmatic surfaces. To explain how pollen transfer occurs, we suggested an unusual method of autogamy, based on our field studies (Anderson et al., 2000b). As anthers dehisce in bud stage, the corolla throat closely surrounds the anther cylinder and thus becomes coated with pollen. As in other Wahlenbergia, the pollen is actually presented by a stylar brush that pushes the pollen up near the open mouth of the now open corolla. The ring of pollen in the throat is obvious because its white color stands out against the pink corolla. Given that experimental studies imply that pollen stays viable throughout the receptive stage of the stigma, we hypothesized that transfer from corolla to stigmata occurs when flowers are shaken by the ever-present wind, promoting contact in the narrow throat between the stigmata and corolla. A similar situation was reported in Iris versicolor from Kent Island, New Brunswick, where facultative self-pollination was promoted by wind that forced reflexed stigmas to brush against petals of the same flowers where pollen had settled (Zink & Wheelwright, 1997).

H. HYPOTHESIZED POLLINATION OF COLONIZERS

The majority of the colonizers seem to have been pollinated either by insects or by wind. However, this association is likely related not to the successful colonization of species with these pollination systems but to enhanced chances of reproduction of established plants through SC or, possibly, to some other feature, such as greater dispersal ability. Obviously, anemophilous species are preadapted to successfully reproduce on islands, because they do not need biotic pollinators (Ehrendorfer, 1979). Bird-pollinated species that dispersed successfully would have done well, given that these islands eventually also included two hummingbirds. However, entomophilous species would have encountered serious reproductive problems on islands with no dedicated insect pollinators.

According to Carlquist (1974), long-distance dispersal is often not achieved by a single individual or disseminule. For instance, there is evidence for multiple disseminules per species in the colonization of Krakatau islets (Docters van Leeuwen, 1936) and on a newly emerged island near Iceland (Einarsson, 1967). Thus, it is possible to envision even the establishment of SI or dioecious colonizers when two or more compatible plants arrive. Furthermore, perennials have an advantage over short-lived species: Longevity increases the chances of securing sufficient pollination for effective sexual reproduction that establishes and maintains a species (Wallace, 1895). Recall that most Fernandezian species are woody. The longer life cycles also increase the chances of encountering, eventually, effective pollinators: for example, the arrival of hummingbirds on these islands.

I. CONSERVATION

Proportionally, there are many more recorded extinctions of vascular plants from islands than from continental areas (Reid & Miller, 1989; Smith et al., 1993; Frankham, 1997). Thus, it is even more important to invest extra effort in protecting the remaining endangered island species (Cronk, 1997; Carlquist, 1998; Raven, 1998). Conservation or restoration programs cannot be effective without an understanding of the reproductive biology (Hamrick et al., 1991; Karron, 1991; Weller, 1994). In addition, data on reproductive biology are an integral part of understanding and, especially, interpreting problems of genetic diversity generated via molecular studies (Karron, 1987; Crawford, 1990; Barrett & Kohn, 1991; Holsinger, 1991).

For the Juan Fernandez flora, even though insect pollinators are virtually absent, wind pollination serves a large percentage of the species, and pollen transfer--at some level--seems not be a strongly limiting factor. However, a large number of the anemophilous species lack some features that would make wind pollination most effective (cf., Regal, 1982; Whitehead, 1983). Among them, one is very important for conservation purposes: close proximity of con-specifics; that is, relatively close spacing of compatible plants. The rapidly spreading populations of invasives and the continued foraging by rabbits and feral goats on native species (Stuessy et al., 1997, 1998b) are significantly reducing and separating the already small populations of many species. As Weller (1994) pointed out, wholesale destruction of natural areas may have contributed far more to rarity than have evolutionary inadequacies of plant breeding systems or pollination biology. The intrusion of invasives exacerbates this situation. The pollin ation systems of island plants are fragile, and habitat changes may reduce rates of already insufficient pollination--obviously with negative consequences for maintenance of populations and species.

There is particular concern for the SI and dioecious species, especially those with few individuals left. For instance, in the SI Dendroseris neriifolia (with only three individuals in the wild in Quebrada Lapiz, plus a few individuals cultivated at GONAF gardens in San Juan Bautista; Stuessy et al., 1998b), examination of open-pollinated flowers on cultivated individuals showed no pollen grains on stigmata or pollen tubes in styles; that is, no natural pollination (Anderson et al., 2000b). Furthermore, the open-pollinated seed set of the small cultivated population of the SI D. pruinata was extremely low. Thus, for both of those examples, at least in part because of the reproductive system (coupled with rarity), the species are at severe risk of extinction in the near future if the existing very few plants are not protected, and if their pollination/reproduction is not enhanced.

Among the dioecious species, two of the seven species of Robinsonia are already rare: R. macrocephala has not been collected since 1917, and R. berteroi is known from one male specimen (Quebrada Villagra; Stuessy et al., 1998b). Thus, care must be taken to continue programs of propagation for these very rare species. For Coprosma and Fagara, although more field data are needed, the relatively low density of compatible plants seems to be a concern for their effective sexual reproduction. Some other species are more stable at present. For instance, Juania australis was saved from extinction because of its ability to grow on the high and inaccessible ridges (Stuessy et al., 1998b) beyond easy access--at least of humans. However, there are still no studies of the reproductive biology of this endemic genus, so its success in seed production is unknown. Pernettya rigida has vegetative reproduction through stolons, sexual seed production mediated by wind, and dispersal of berries by birds. All of these features comb ine to promote this species as a colonizer of new habitats. For the same reasons it thrives on both main islands of the archipelago and seems not to be endangered (Anderson et al., 2000a).

The ultimate fate of some species may depend on preserving the plant-hummingbird relationship (Buchmann & Nabban, 1996; Kearns et al., 1998; Nabhan et al., 1998; Roy et al., 1999), including the web of organisms that affect both plant and pollinator. The density of conspecifics is fundamental to favor higher levels of interplant pollen transfer, Reproductive success depends on the quantity and quality of effective pollination visits, and both of these are likely to depend on local abundance (Kunin, 1997). Here, again, invasives pose a serious problem. They outcompete endemic species for habitat, and if they also produce abundant nectar and are visited by hummingbirds (as is true in general), their prosperity may also lead both to a reduced visitation rate and to reduced effective pollination of the endemics (Brooke, 1987; Colwell, 1989; Bemardello et al., 2000). The highly invasive bramble (Rubus ulmifolius) is heavily visited as a nectar source by the native hummingbird (Brooke, 1987), and it seems to be bet ter suited than the endemic hummingbird to feed on this plant species (Colwell, 1989). This fact, among others, is postulated to have differentially favored the native hummingbird at the expense of the endemic species, whose populations are declining (Colwell, 1989).

The hummingbird-pollinated Nicotiana cordiflora is endemic to Alejandro Selkirk Island, where the endemic hummingbird (S. fernandensis) is now considered extinct (Brooke, 1987; Colwell, 1989). Fortunately for the ornithophilous plant species, since 1981 the native hummingbird (S. sephaniodes) has become established on Alejandro Selkirk Island (although it is not clear if they are a breeding population or only adventives from Robinson Crusoe Island; Brooke, 1987; Colwell, 1989). Given the number of ornithophilous species and their importance to the flora, the complete loss of the hummingbirds could have immense conservation implications. Regrettably, the goat-maintained grassland of the island's lower slopes, largely composed of introduced species, and the native fern forest of the uplands, combine to render major portions of this island poor hummingbird habitat (Brooke, 1987). Thus, to maintain this plant--animal interaction, the introduced goats should be eliminated and the native habitats restored.

It is also crucial to maintain the interaction of hummingbirds with Dendroseris litoralis. This species is known presently only from several plants on Morro Spartan and adjacent Santa Clara Island; hence it is nearly extinct in the wild. Fortunately, it is extensively cultivated on Robinson Crusoe Island (Stuessy et al., 1998b).

A recently discovered bee (Engel, 2000) is considered irrelevant for the evolution of the reproductive systems on the islands (Anderson et al., 2001b). Although currently a rare endemic, we propose that the bee is likely also to be discovered in continental South America. However, this bee may be important in future conservation or restoration programs on the island. The bees--scarce at present--were observed collecting pollen on the Wahlenbergia hybrid (Anderson et al., 2000b; Engel, 2000) in San Juan Bautista and its environs. Although likely recent introductions, they could become beneficial, promoting allogamy for this and other taxa. Given the lack of significant native insect fauna, these bees would not seem to pose a competitive danger to other insects; they will displace no native pollinator fauna. However, just as the more common continental species of hummingbird may be responsible for the decline of the endemic hummingbird (Colwell, 1989), the introduced bees may outcompete the hummingbirds for flo ral nectar and have a negative effect on the vigor of these island "banner species."

Finally, a new pressure has been added: Introduced locusts are widespread in the most arid and dry part of Robinson Crusoe Island, near the landing strip. They are voracious, much affecting the scarce native and endemic species adapted to live in the arid low-island conditions. We observed them virtually destroying a population of W. berteroi, eating stems, leaves, even fruits, and seeds.

In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would do much to enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species. The depauperate soil of these islands is mostly eroded, exposing bedrock to degradation and retaining less rainwater. Erosion is, in fact, a considerable problem in these islands, particularly on the steep volcanic terrain (Sanders et al., 1982).

VI. Conclusions

The flora typically comprises perennials (ca. 90%). The majority of the species (60%) have comparatively insignificant flowers, either very small (46%) or small (14%). Inconspicuous flowers (a shape category for flowers with "no optical attraction") are widespread (41% of the taxa), as are dish-shaped flowers (35%). Green is the most frequent flower color (41% of the species), followed by white and yellow (26% and 12%, respectively). Most species (70%) are hermaphroditic, 9% are dioecious, and 9% are monoecious. Around 32% of the species are protandrous; 7%, protogynous. The few studies of compatibility of the flora (for 14% of the species) indicate that 85% of the species are self-compatible (SC) and that only 17% are self-incompatible (SI). Although SC is frequent, the level of autogamy is low. Nevertheless, geitonogamy, and thus selfing, would be the most frequent mechanism of pollen transfer. Outcrossing is mainly achieved through dioecy and SI, promoted by dichogamy, and facilitated by wind pollination.

About 55% of the species offer nectar as a reward; about 2%, pollen. Floral visitors are rare to uncommon. The activity of the two native hummingbird species (the endemic Sephanoides fernandensis and the native S. sephaniodes), recorded for 9% of the flora, is an exception to this generalization. Native insects (flies, moths, and beetles) were observed or recorded for only 11 species, but even on these, recorded floral visits were rare, In addition, the insects showed no marked preference for flowers over other plant parts and exhibited no observable species fidelity. Two species of introduced ants and a new endemic bee were recorded as well. It seems likely that the latter is a recent introduction.

We conclude that 9% of the flora is bird pollinated and 47% wind pollinated, including a special case of wind as an indirect force of pollen transfer in Wahlenbergia berteroi. Given the rare and imprecise nature of native-insect visitation, there is no certainty that any of the species are truly insect pollinated. This is a tribute to the paucity of the continental fauna, because we hypothesize that most of the original colonizers were either insect or wind pollinated.

Pairwise comparisons of many features were tested statistically. We confirmed the association between the following features known to be common for oceanic islands: green and very small flowers, green and inconspicuous flowers, and very small and inconspicuous flowers. Monoecious species are largely herbaceous and have mostly very small, inconspicuous, green flowers. Dioecious species are woody, chiefly with green and yellow, dish-shaped, medium-sized, wind-pollinated flowers. We propose that the progenitors of Coprosma, Juania, Fagara, and Pernettya already exhibited at least incipient dioecy; in Robinsonia dioecy seems to have arisen in situ. Both gynomonoecy in Cuminia and gynodioecy in Rhaphithamnus venustus likely arose in situ.

Hermaphroditic flowers and SC likely could have favored the reproduction and establishment of colonizers. There is a correlation between a number of floral features and the hypothesized pollination of colonizers. Therefore, the flower color, shape, and size of the extant flora may, to some large extent, express the pollination syndromes of colonizers rather than representing extant pollination. In addition, the presence of nectar in the extant flora does not necessarily indicate biotic pollination; for example, some clearly wind-pollinated species also bear nectar. Thus, studies of the reproductive biology of oceanic island plants need to be conducted species by species before broad, effective generalizations can be made, because the observed features can be misleading.

We assessed possible changes in the pollination system by comparison of species for which we have reliable data (for 39 genera) with the hypothesized pollination of their colonist progenitors. Most (70%) retained the pollination system of the colonizers, either by wind or hummingbirds. In 30% of genera, at least some species bear a different pollination system: In four genera the ancestral insect-pollination system seems to have switched to hummingbird pollination, and in seven other genera from insect to wind pollination. For the insect-to-bird pollination examples, flower length seems to have increased due to a specialization to hummingbird pollination. In Wahlenbergia, the ancestors were likely insect pollinated, with a secondary system of autogamy; but in the islands, with paucity of insect pollination, this selfing mechanism became the rule. A wind-aided mechanism of autogamy in W. berteroi must have arisen in situ. We also suspect that much of the remaining 44% of the flora for which pollination is unkn own is likely to be wind pollinated or autogamous (or both). The bird-pollinated species are mostly obvious, and insect pollination is mostly insignificant. Thus, it is likely that a much higher percentage of the flora switched from insect pollination of colonizers to abiotic or autogamous endemics.

The species that arrived having preadaptations to bird pollination seem to have retained those features. The hummingbird-pollinated species that were perennial and SC could have reproduced and become established even before the hummingbirds arrived. Considering the many simultaneous flowers produced by these species, the amount of nectar exuded, the hummingbird behavior, and the existence of SC, hummingbird visits likely favor geitonogamous rather than xenogamous pollen transfer.

Anemophily for a portion of the flora is expected, given the characteristics of the proposed progenitors; for other species, the lack of alternative means of pollination must have led to anemophily. For the latter, the features associated with wind pollination may not be so obvious. If there has not been sufficient evolutionary time or selection pressure, ancestral and misleading features of abiotic pollination may be retained. More extensive studies of the pollination biology of island floras may reveal anemophily for species in which features of the flowers would not have predicted it.

Given the many recorded extinctions of vascular plants from islands versus those from continental areas, it is imperative to invest additional effort in protecting the remaining island species. Conservation or restoration programs cannot be effective without a deep and broad understanding of the reproductive biology of the plants. In order to conserve these plants, programs must involve a combination of reproductive and environmental measures. The ultimate fate of some species may depend on preserving the plant--hummingbird relationship, including the web of organisms that affect both plant and pollinator. The populations of introduced animals and weeds must be controlled. Experimentally produced allogamous seeds would enhance diversity in restoration programs. In addition, the preservation of habitat seems to be the central challenge to indirectly protect the unique island species.

VII. Acknowledgments

We thank the National Science Foundation, the University of Connecticut Research Foundation, the Department of Ecology and Evolutionary Biology, CONICET, CONICOR, and SECYT--Universidad Nacional de Cordoba (Argentina) for financial support and a sabbatical leave. We are grateful to Edward O. Wilson, Jane O'Donnell, Michael S. Engel, Haroldo Toro, and Charles Michener for insect identification, to Nalini Ravishanker for statistical advice, to Maryke Schlehofer, Leonardo Galetto, and Dick Jensen for helping in many ways, and to Virge A. Kask for help with the illustrations and Jim Hill with the manuscript. The assistance of CONAF (Chile) is sincerely acknowledged, especially Ivan Leiva (chief of the Juan Fernandez National Park) and park guides Jorge Angulo, Guillermo Araya, Oscar Chamorro, Bernardo Lopez, Manuel Recabarren, and Ramon Schiller, without whose invaluable help this work would have been impossible. Further assistance was provided by the Meteorological Service of Robinson Crusoe Island (especially O svaldo Jara and Alex Meneses) and by Fernandezians Jose M. Gutierrez, Valeria Salzmann, Jorge Palomino, and Juanita Lopez. We dedicate this article to the memory of G. L. Stebbins, for his stimulating work on plant breeding systems and evolution.

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[Figure 1 omitted]

[Figure 2 omitted]

[Figure 3 omitted]

[Figure 4 omitted]

[Figure 5 omitted]

[Figure 6 omitted]

Table I

Habit, flower, and floral biology features, breeding system, floral
visitors observed, and pollination of the Juan Fernandez autochthonous
angiosperm flora

Dicots

                                          Species (b)


Family, genus (a)                    Native

Apiaceae
 Apium
                                     A. australe
 Centella
                                     C. asiatica
 Eryngium




Asteraceae (i)
 Abrotanella
                                     A. linearifolia
 Centaurodendron (k)



 Dendroseris (k)











 Erigeron




 Gamochaeta

 Lagenophora

                                     L. hariotti
 Robinsonia (k)







 Taraxacum


                                     T. fernandezianum
                                     T. subspathulatum
 Yunquea (k)

Berberidaceae
 Berberis


Boraginaceae
 Selkirkia (k)

Brassicaceae
 Cardamine

                                     C. chenopodioides
Campanulaceae
 Lobelia
                                     L. alata
 Wahlenbergia








Caryophyllaceae
 Spergularia



Chenopodiaceae
 Chenopodium



 Sarcocornia

                                     S. fruticosa
Convolvulaceae
 Calystegia
                                     C. tuguriorum
 Dichondra
                                     D. sericea
Empetraceae
 Empetrum
                                     E. rubrum
Ericaceae
 Pernettya


Euphorbiaceae
 Dysopsis

Fabaceae
 Sophora


Flacourtiaceae
 Azara


Gunneraceae
 Gunnera





Haloragaceae
 Haloragis


Lamiaceae
 Cuminia (k)

Lactoridaceae
 Lactoris (k)

Loranthaceae
 Notanthera
                                     N. heterophylla
Myrtaceae
 Myrceugenia


 Myteola
                                     M. nummularia
 Ugni

Piperaceae
 Peperomia




Plantaginaceae
 Plantago
                                     P. australis
                                     P. frima

Ranunculaceae
 Ranunculus

Rhamnaceae
 Colletia

Rosacese
 Acaena


 Margyracaena


 Margyricarpus


 Rubus

                                     R. geoides
Rubiacese
 Coprosma


 Galium


 Hedyotis

                                     H. salzmannii
 Nertera

                                     N. granadensis
Rutaceae
 Fagara


Santalaceae
 Santalum

Saxifragaceae
 Escallonia


Scrophulariaceae
 Euphrasia

 Mimulus
                                     M. glabratus
Solanaceae
 Nicotiana

 Solanum

                                     S. pentlandii
                                        ssp. interandinum

Urticaceae
 Boehmeria


 Parietaria
                                     P. debilis
 Urtica


                                     U. berteroana


Verbenaceae
 Rhaphithamnus

Winteraceae
 Drimys


                                           Species (b)


Family, genus (a)                    Endemic

Apiaceae
 Apium
                                     A. fernandezianum
 Centella

 Eryngium
                                     E. bupleuroides
                                     E. inaccessum
                                     E. sarcopphyllum
                                     E. x fernandezianum
Asteraceae (i)
 Abrotanella

 Centaurodendron (k)
                                     C. dracaenoides
                                     C. palmiforme

 Dendroseris (k)
                                     D. berteroana
                                     D. gigantea
                                     D. litoralis
                                     D. macrantha
                                     D. macrophylla
                                     D. marginata
                                     D. micrantha
                                     D. neriifolia
                                     D. pinnata
                                     D. pruinata
                                     D. regia
 Erigeron
                                     E. fernandezianus
                                     E. ingae
                                     E. luteoviridis
                                     E. rupicola
 Gamochaeta
                                     G. fernandeziana
 Lagenophora


 Robinsonia (k)
                                     R. berteroi
                                     R. evenia
                                     R. gayana
                                     R. gracilis
                                     R. macrocephala
                                     R. masafuerae
                                     R. thurifera
 Taraxacum




 Yunquea (k)
                                     Y. tenzii
Berberidaceae
 Berberis
                                     B. corymbosa
                                     B. masafuerana
Boraginaceae
 Selkirkia (k)
                                     S. berteroi
Brassicaceae
 Cardamine

                                     C. krusselii
Campanulaceae
 Lobelia

 Wahlenbergia
                                     W. berteroi
                                     W. fernandeziana
                                     W. grahamiae
                                     W. masafuerae
                                     W. tuberosa
                                     W. fernandeziana x
                                        W. grahamiae
                                        (= hybrid)
Caryophyllaceae
 Spergularia

                                     S. confertifolia
                                     S. masafuerana
Chenopodiaceae
 Chenopodium
                                     Ch. crusoeanum
                                     Ch. nesodendron
                                     Ch. sanctae-clarae
 Sarcocornia


Convolvulaceae
 Calystegia

 Dichondra

Empetraceae
 Empetrum

Ericaceae
 Pernettya

                                     P. rigida
Euphorbiaceae
 Dysopsis
                                     D. hirsuta
Fabaceae
 Sophora
                                     S. fernandeziana
                                     S. masafuerana
Flacourtiaceae
 Azara
                                     A. serrata var.
                                      fernandeziana
Gunneraceae
 Gunnera
                                     G. bracteata
                                     G. glabra
                                     G. masafuerae
                                     G. peltata
                                     G. bracteata x peltata
Haloragaceae
 Haloragis
                                     H. masafuerana
                                     H. masatierrana
Lamiaceae
 Cuminia (k)
                                     C. eriantha
Lactoridaceae
 Lactoris (k)
                                     L. fernandeziana
Loranthaceae
 Notanthera

Myrtaceae
 Myrceugenia
                                     M. fernandeziana
                                     M. schulzei
 Myteola

 Ugni
                                     U. selkirkii
Piperaceae
 Peperomia
                                     P. berteroana
                                     p. fernandeziana
                                     P. margaritifera
                                     P. skottsbergii
Plantaginaceae
 Plantago


                                     P. fernandezia
Ranunculaceae
 Ranunculus
                                     R. caprarum
Rhamnaceae
 Colletia
                                     C. spartioides
Rosacese
 Acaena

                                     A. masafuerana
 Margyracaena

                                     M. skottsbergii
 Margyricarpus

                                     M. digynus
 Rubus


Rubiacese
 Coprosma
                                     C. oliveri
                                     C. pyrifolia
 Galium

                                     G. masafueranum
 Hedyotis


 Nertera


Rutaceae
 Fagara
                                     F. externa
                                     F. mayu
Santalaceae
 Santalum
                                     S. fernandezianum
Saxifragaceae
 Escallonia

                                     E. callcottiae
Scrophulariaceae
 Euphrasia
                                     E. formosissima
 Mimulus

Solanaceae
 Nicotiana
                                     N. cordifolia
 Solanum



                                     S. fernandezianum
Urticaceae
 Boehmeria

                                     B. excelsa
 Parietaria

 Urtica



                                     U. glomeruliflora
                                     U. masafuerae
Verbenaceae
 Rhaphithamnus
                                     R. venustus
Winteraceae
 Drimys
                                     D. confertifolia




Family, genus (a)                     Habit

Apiaceae
 Apium                               Perennial herb

 Centella                            Perennial herb

 Eryngium                            Shrub/tree




Asteraceae (i)
 Abrotanella                         Perennial herb

 Centaurodendron (k)                 Tree



 Dendroseris (k)                     Shrub/tree











 Erigeron                            Shrub




 Gamochaeta                          Perennial herb

 Lagenophora                         Perennial herb


 Robinsonia (k)                      Shrub







 Taraxacum                           Perennial herb




 Yunquea (k)                         Tree

Berberidaceae
 Berberis                            Shrub


Boraginaceae
 Selkirkia (k)                       Shrub

Brassicaceae
 Cardamine                           Annual or perennial
                                      herb

Campanulaceae
 Lobelia                             Annual herb

 Wahlenbergia                        Shrub








Caryophyllaceae
 Spergularia                         Perennial herb



Chenopodiaceae
 Chenopodium                         Shrub



 Sarcocornia                         Perennial herb


Convolvulaceae
 Calystegia                          Shrub

 Dichondra                           Perennial herb

Empetraceae
 Empetrum                            Shrub

Ericaceae
 Pernettya                           Shrub


Euphorbiaceae
 Dysopsis                            Perennial herb

Fabaceae
 Sophora                             Tree


Flacourtiaceae
 Azara                               Tree


Gunneraceae
 Gunnera                             Perennial herb





Haloragaceae
 Haloragis                           Perennial herb


Lamiaceae
 Cuminia (k)                         Shrub

Lactoridaceae
 Lactoris (k)                        Shrub

Loranthaceae
 Notanthera                          Shrub

Myrtaceae
 Myrceugenia                         Tree


 Myteola                             Shrub

 Ugni                                Shrub

Piperaceae
 Peperomia                           Perennial herb




Plantaginaceae
 Plantago                            Shrub



Ranunculaceae
 Ranunculus                          Perennial herb

Rhamnaceae
 Colletia                            Shrub

Rosacese
 Acaena                              Perennial herb


 Margyracaena                        Perennial
                                      herb

 Margyricarpus                       Perennial
                                      herb

 Rubus                               Perennial
                                      herb

Rubiacese
 Coprosma                            Tree


 Galium                              Perennial
                                      herb

 Hedyotis                            Perennial
                                      herb

 Nertera                             Perennial
                                      herb

Rutaceae
 Fagara                              Tree


Santalaceae
 Santalum                            Tree

Saxifragaceae
 Escallonia                          Shrub/tree


Scrophulariaceae
 Euphrasia                           Shrub

 Mimulus                             Annual herb

Solanaceae
 Nicotiana                           Shrub

 Solanum                             Perennial
                                      herb



Urticaceae
 Boehmeria                           Tree


 Parietaria                          Annual herb

 Urtica                              Annual or
                                      perennial
                                      herb



Verbenaceae
 Rhaphithamnus                       Tree

Winteraceae
 Drimys                              Tree


                                              Flower

                                     Size, in mm
Family, genus (a)                    (category)

Apiaceae
 Apium                               2 x 2 (very small)

 Centella                            2 x 2 (very small)

 Eryngium                            5 x 4 (small)




Asteraceae (i)
 Abrotanella                         3 x 2 (very small)

 Centaurodendron (k)                 20 x 8 (large)



 Dendroseris (k)                     22 x 27 (very large)











 Erigeron                            8 x 8 (medium)




 Gamochaeta                          3 x 3 (very small)

 Lagenophora                         6 x 6 (medium)


 Robinsonia (k)                      9 x 6 (medium)







 Taraxacum                           30 x 8 (large)




 Yunquea (k)                         30 x 15 (very large)

Berberidaceae
 Berberis                            6 x 8 (medium)


Boraginaceae
 Selkirkia (k)                       13 x 7 (medium)

Brassicaceae
 Cardamine                           6 x 4 (small)


Campanulaceae
 Lobelia                             11 x 5 (medium)

 Wahlenbergia                        12 x 8 (medium)








Caryophyllaceae
 Spergularia                         2.5 x 2.5 (very small)



Chenopodiaceae
 Chenopodium                         1 x 1.5 (very small)



 Sarcocornia                         2 x 2 (very small)


Convolvulaceae
 Calystegia                          40 x 50 (very large)

 Dichondra                           2.5 x 2.5 (very small)

Empetraceae
 Empetrum                            3 x 2 (very small)

Ericaceae
 Pernettya                           8 x 4 (medium)


Euphorbiaceae
 Dysopsis                            2 x 2 (very small)

Fabaceae
 Sophora                             30 x 5 (large)


Flacourtiaceae
 Azara                               4 x 6 (small)


Gunneraceae
 Gunnera                             1.2 x 1.2 (very small)





Haloragaceae
 Haloragis                           4.3 x 7 (medium)


Lamiaceae
 Cuminia (k)                         15 x 5 (medium)

Lactoridaceae
 Lactoris (k)                        3.5 x 3 (small)

Loranthaceae
 Notanthera                          12 x 2 (small)

Myrtaceae
 Myrceugenia                         2.5 x 2.5 (very small)


 Myteola                             8 x 6 (medium)

 Ugni                                8 x 8 (medium)

Piperaceae
 Peperomia                           1 x 1 (very small)




Plantaginaceae
 Plantago                            5 x 2 (small)



Ranunculaceae
 Ranunculus                          8 x 2 (small)

Rhamnaceae
 Colletia                            5 x 3 (small)

Rosacese
 Acaena                              2 x 1.5 (very
                                      small)

 Margyracaena                        2 x 1.5 (very
                                      small)

 Margyricarpus                       1.5 x 1.5 (very
                                      small)

 Rubus                               20 x 15 (large)


Rubiacese
 Coprosma                            7 x 6 (medium)


 Galium                              2 x 3 (very small)


 Hedyotis                            5 x 2 (small)


 Nertera                             2 x 2 (very small)


Rutaceae
 Fagara                              4 x 4 (small)


Santalaceae
 Santalum                            4 x 4 (small)

Saxifragaceae
 Escallonia                          15 x 5 (medium)


Scrophulariaceae
 Euphrasia                           12 x 6 (medium)

 Mimulus                             25 x 10 (large)

Solanaceae
 Nicotiana                           25 x 4 (medium)

 Solanum                             10 x 8 (medium)




Urticaceae
 Boehmeria                           1.5 x 1.5 (very
                                      ssmall)

 Parietaria                          1 x 1 (very small)

 Urtica                              1.5 x 1 (very
                                       small)




Verbenaceae
 Rhaphithamnus                       29 x 6 (large)

Winteraceae
 Drimys                              4 x 23 (medium)


                                                    Flower


Family, genus (a)                    Shape (c)        Color

Apiaceae
 Apium                               Dish             White/green

 Centella                            Dish             White/purplish

 Eryngium                            Dish
                                                      Green
                                                      Violet
                                                      Green
                                                      Violet
Asteraceae (i)
 Abrotanella                         Dish (discoid)   Brown/red

 Centaurodendron (k)                 Dish (discoid)   Blue, dark purple



 Dendroseris (k)                     Dish (radiate)
                                                      White
                                                      White
                                                      Orange
                                                      Orange
                                                      Orange
                                                      Orange
                                                      White
                                                      White
                                                      White
                                                      White
                                                      White
 Erigeron                            Dish (radiate)   White/yellow




 Gamochaeta                          Dish (discoid)   White

 Lagenophora                         Dish (radiate)   White/pink


 Robinsonia (k)                      Dish (radiate)
                                                      Yellow
                                                      Yellow
                                                      Yellowish green
                                                      Yellow
                                                      Yellow
                                                      Green
                                                      Yellowish green
 Taraxacum                           Dish (ligulate)  Yellow




 Yunquea (k)                         Dish

Berberidaceae
 Berberis                            Dish             Yellow


Boraginaceae
 Selkirkia (k)                       Bell             White

Brassicaceae
 Cardamine                           Dish             White


Campanulaceae
 Lobelia                             Tube             Blue

 Wahlenbergia                        Dish, tube
                                                      Pink
                                                      White
                                                      White
                                                      White
                                                      White
                                                      White


Caryophyllaceae
 Spergularia                         Inconspicuous    White,
                                                       white/pink


Chenopodiaceae
 Chenopodium                         Inconspicuous    Green



 Sarcocornia                         Inconspicuous    Green


Convolvulaceae
 Calystegia                          Bell             White

 Dichondra                           Bell             Green

Empetraceae
 Empetrum                            Bell             Red

Ericaceae
 Pernettya                           Bell             White


Euphorbiaceae
 Dysopsis                            Dish             Green

Fabaceae
 Sophora                             Flag             Yellow


Flacourtiaceae
 Azara                               Brush            Yellow


Gunneraceae
 Gunnera                             Inconspicuous    Red





Haloragaceae
 Haloragis                           Dish             Green


Lamiaceae
 Cuminia (k)                         Tube             Violet

Lactoridaceae
 Lactoris (k)                        Inconspicuous    Green

Loranthaceae
 Notanthera                          Tube             White

Myrtaceae
 Myrceugenia                         Bell             White


 Myteola                             Bell             White

 Ugni                                Bell             White

Piperaceae
 Peperomia                           Inconspicuous    Green




Plantaginaceae
 Plantago                            Tube             Brown



Ranunculaceae
 Ranunculus                          Tube             White/pink

Rhamnaceae
 Colletia                            Tube             White/pink

Rosacese
 Acaena                              Inconspicuous    Green


 Margyracaena                        Inconspicuous    Green


 Margyricarpus                       Inconspicuous    Green


 Rubus                               Dish             White


Rubiacese
 Coprosma                            Tube             Green/brown


 Galium                              Tube             White/green


 Hedyotis                            Tube             White


 Nertera                             Tube             White/green


Rutaceae
 Fagara                              Inconspicuous    Brown


Santalaceae
 Santalum                            Dish             White

Saxifragaceae
 Escallonia                          Tube             Red


Scrophulariaceae
 Euphrasia                           Tube             White

 Mimulus                             Tube             Yellow

Solanaceae
 Nicotiana                           Tube             Dark red

 Solanum                             Dish             Violet,yellow
                                                      /white



Urticaceae
 Boehmeria                           Inconspicuous    Green


 Parietaria                          Inconspicuous    Green

 Urtica                              Inconspicuous    Green





Verbenaceae
 Rhaphithamnus                       Tube             Violet

Winteraceae
 Drimys                              Dish             White





Family, genus (a)                    Sex

Apiaceae
 Apium                               Hermaphroditic

 Centella                            Hermaphroditic

 Eryngium                            Hermaphroditic




Asteraceae (i)
 Abrotanella                         Gynomonecious

 Centaurodendron (k)                 Andromonecious



 Dendroseris (k)                     Hermaphroditic











 Erigeron                            Gynomonecious




 Gamochaeta                          Hermaphroditic

 Lagenophora                         Monoecious/
                                      gynomonecious

 Robinsonia (k)                      Dioecious







 Taraxacum                           Hermaphroditic/
                                      gynomonoe
                                      cious


 Yunquea (k)                         Hermaphroditic

Berberidaceae
 Berberis                            Hermaphroditic


Boraginaceae
 Selkirkia (k)                       Hermaphroditic

Brassicaceae
 Cardamine                           Hermaphroditic


Campanulaceae
 Lobelia                             Hermaphroditic

 Wahlenbergia                        Hermaphroditic








Caryophyllaceae
 Spergularia                         Hermaphroditic



Chenopodiaceae
 Chenopodium                         Gynomonoecious



 Sarcocornia                         Hermaphroditic/
                                      polygamous

Convolvulaceae
 Calystegia                          Hermaphroditic

 Dichondra                           Hermaphroditic

Empetraceae
 Empetrum                            Polygamous

Ericaceae
 Pernettya                           Dioecious


Euphorbiaceae
 Dysopsis                            Monoecious

Fabaceae
 Sophora                             Hermaphroditic


Flacourtiaceae
 Azara                               Hermaphroditic


Gunneraceae
 Gunnera                             Hermaphroditic





Haloragaceae
 Haloragis                           Hermaphroditic


Lamiaceae
 Cuminia (k)                         Gynomonoecious

Lactoridaceae
 Lactoris (k)                        Gynomonoecious

Loranthaceae
 Notanthera                          Hermaphroditic

Myrtaceae
 Myrceugenia                         Hermaphroditic


 Myteola                             Hermaphroditic

 Ugni                                Hermaphroditic

Piperaceae
 Peperomia                           Hermaphroditic




Plantaginaceae
 Plantago                            Hermaphroditic



Ranunculaceae
 Ranunculus                          Hermaphroditic

Rhamnaceae
 Colletia                            Hermaphroditic

Rosacese
 Acaena                              Hermaphroditic


 Margyracaena                        Hermaphroditic


 Margyricarpus                       Hermaphroditic


 Rubus                               Hermaphroditic


Rubiacese
 Coprosma                            Dioecious


 Galium                              Hermaphroditic


 Hedyotis                            Hermaphroditic


 Nertera                             Hermaphroditic


Rutaceae
 Fagara                              Hermaphroditic


Santalaceae
 Santalum                            Hermaphroditic

Saxifragaceae
 Escallonia                          Hermaphroditic


Scrophulariaceae
 Euphrasia                           Hermaphroditic

 Mimulus                             Hermaphroditic

Solanaceae
 Nicotiana                           Hermaphroditic

 Solanum                             Hermaphroditic




Urticaceae
 Boehmeria                           Monoecious


 Parietaria                          Polygamous

 Urtica                              Monoecious





Verbenaceae
 Rhaphithamnus                       Gynodioecious

Winteraceae
 Drimys                              Hermaphroditic





Family, genus (a)                    Dichogamy (d)

Apiaceae
 Apium

 Centella

 Eryngium                            Protogyny




Asteraceae (i)
 Abrotanella                         Protandry

 Centaurodendron (k)                 Protandry



 Dendroseris (k)                     Protandry











 Erigeron                            Protandry




 Gamochaeta                          Protandry

 Lagenophora                         Protandry


 Robinsonia (k)                      Protandry







 Taraxacum                           Protandry




 Yunquea (k)                         Protandry

Berberidaceae
 Berberis


Boraginaceae
 Selkirkia (k)

Brassicaceae
 Cardamine


Campanulaceae
 Lobelia                             Protandry

 Wahlenbergia                        Protandry








Caryophyllaceae
 Spergularia



Chenopodiaceae
 Chenopodium



 Sarcocornia


Convolvulaceae
 Calystegia

 Dichondra

Empetraceae
 Empetrum

Ericaceae
 Pernettya


Euphorbiaceae
 Dysopsis

Fabaceae
 Sophora


Flacourtiaceae
 Azara


Gunneraceae
 Gunnera                             Protandry





Haloragaceae
 Haloragis                           Protandry


Lamiaceae
 Cuminia (k)

Lactoridaceae
 Lactoris (k)                        Protogyny

Loranthaceae
 Notanthera

Myrtaceae
 Myrceugenia                         Protogyny


 Myteola

 Ugni                                Protandry

Piperaceae
 Peperomia




Plantaginaceae
 Plantago                            Protogyny



Ranunculaceae
 Ranunculus

Rhamnaceae
 Colletia

Rosacese
 Acaena


 Margyracaena


 Margyricarpus


 Rubus


Rubiacese
 Coprosma


 Galium


 Hedyotis


 Nertera


Rutaceae
 Fagara


Santalaceae
 Santalum

Saxifragaceae
 Escallonia                          Protandry


Scrophulariaceae
 Euphrasia

 Mimulus

Solanaceae
 Nicotiana                           Protandry

 Solanum                             Protandry




Urticaceae
 Boehmeria


 Parietaria

 Urtica





Verbenaceae
 Rhaphithamnus                       Protandry

Winteraceae
 Drimys                              Protogyny



                                     Breeding
                                     system                     Floral
Family, genus (a)                    determined (d)             reward

Apiaceae
 Apium                                                          Nectar

 Centella                                                       Nectar

 Eryngium                                                       Nectar




Asteraceae (i)
 Abrotanella                                                    Nectar

 Centaurodendron (k)                                            Nectar



 Dendroseris (k)                                                Nectar


                                     Self-compatible




                                     Self-incompatible



 Erigeron                                                       Nectar




 Gamochaeta                                                     Nectar

 Lagenophora                                                    Nectar


 Robinsonia (k)                                                 Nectar







 Taraxacum                           Agamospermous              Nectar




 Yunquea (k)                                                    Nectar

Berberidaceae
 Berberis                                                       Nectar
                                     Self-incompatible

Boraginaceae
 Selkirkia (k)                                                  Nectar

Brassicaceae
 Cardamine                                                      Nectar


Campanulaceae
 Lobelia                                                        Nectar

 Wahlenbergia                                                   Nectar
                                     Self-compatible
                                     Self-compatible



                                     Self-compatible


Caryophyllaceae
 Spergularia                                                    Nectar



Chenopodiaceae
 Chenopodium



 Sarcocornia


Convolvulaceae
 Calystegia                                                     Nectar

 Dichondra                                                      Nectar

Empetraceae
 Empetrum

Ericaceae
 Pernettya                                                      Nectar


Euphorbiaceae
 Dysopsis

Fabaceae
 Sophora
                                                                Nectar

Flacourtiaceae
 Azara                               Self-compatible            Nectar


Gunneraceae
 Gunnera





Haloragaceae
 Haloragis

                                     Self-compatible
Lamiaceae
 Cuminia (k)                                                    Nectar

Lactoridaceae
 Lactoris (k)                        Self-compatible

Loranthaceae
 Notanthera                                                     Nectar

Myrtaceae
 Myrceugenia


 Myteola                             Self-compatible (n)        Nectar

 Ugni                                                           Nectar

Piperaceae
 Peperomia




Plantaginaceae
 Plantago


                                     Self-compatible
Ranunculaceae
 Ranunculus                                                     Nectar

Rhamnaceae
 Colletia                                                       Nectar

Rosacese
 Acaena                                                         Nectar


 Margyracaena                                                   Nectar


 Margyricarpus                                                  Nectar


 Rubus                               Self-compatible (a)        Nectar


Rubiacese
 Coprosma                                                       Nectar


 Galium                                                         Nectar


 Hedyotis                                                       Nectar


 Nertera                                                        Nectar


Rutaceae
 Fagara                                                         Nectar


Santalaceae
 Santalum

Saxifragaceae
 Escallonia                          Self-compatible, selfer    Nectar


Scrophulariaceae
 Euphrasia                                                      Nectar

 Mimulus                                                        Nectar

Solanaceae
 Nicotiana                           Self-compatible            Nectar

 Solanum                                                        Pollen



                                     Self-compatible, (o) self
Urticaceae
 Boehmeria                           Self-compatible


 Parietaria

 Urtica





Verbenaceae
 Rhaphithamnus                       Self-compatible            Nectar

Winteraceae
 Drimys                                                         Pollen




                                     Floral visitors
Family, genus (a)                    observed (d)

Apiaceae
 Apium

 Centella

 Eryngium
                                     Hummingbirds, (g, h) flies (g, i)



Asteraceae (i)
 Abrotanella

 Centaurodendron (k)
                                     Hummingbirds (m)


 Dendroseris (k)                     Flies (i)


                                     Hummiogbirds (g, h, l, m)




                                     Flies & moths (l)



 Erigeron
                                     None (l)



 Gamochaeta

 Lagenophora


 Robinsonia (k)                      Flies (i)

                                     None (g, l)
                                     None (g, l)
                                     None (g, l)


                                     None (g, l)
 Taraxacum




 Yunquea (k)

Berberidaceae
 Berberis                            None (l)
                                     Beetles (g)

Boraginaceae
 Selkirkia (k)

Brassicaceae
 Cardamine


Campanulaceae
 Lobelia

 Wahlenbergia
                                     Ants & flies (l)
                                     Months & anys (l)
                                     Moths (g)


                                     Moths, ants, and bee (l)


Caryophyllaceae
 Spergularia



Chenopodiaceae
 Chenopodium


                                     None (l)
 Sarcocornia


Convolvulaceae
 Calystegia

 Dichondra

Empetraceae
 Empetrum

Ericaceae
 Pernettya                           Flies & beetles,
                                     (l) moths (g)

Euphorbiaceae
 Dysopsis                            None (l)

Fabaceae
 Sophora
                                     Hummingbirds (g, h)

Flacourtiaceae
 Azara                               None (g, l)


Gunneraceae
 Gunnera                             None (l)


                                     Not observed (l)


Haloragaceae
 Haloragis

                                     None (l)
Lamiaceae
 Cuminia (k)                         Hummingbirds (g, l)

Lactoridaceae
 Lactoris (k)                        None (l)

Loranthaceae
 Notanthera                          Hummingbirds (g)

Myrtaceae
 Myrceugenia
                                     None (g)

 Myteola

 Ugni                                None (g, l)

Piperaceae
 Peperomia

                                     None (l)


Plantaginaceae
 Plantago


                                     None (l)
Ranunculaceae
 Ranunculus

Rhamnaceae
 Colletia                            Moths (g)

Rosacese
 Acaena


 Margyracaena                        None (l)


 Margyricarpus                       None (l)


 Rubus


Rubiacese
 Coprosma
                                     None (l)

 Galium


 Hedyotis


 Nertera


Rutaceae
 Fagara


Santalaceae
 Santalum

Saxifragaceae
 Escallonia                          Hummingbirds, (g, h, l)
                                     flies, (l) moths (g, l)

Scrophulariaceae
 Euphrasia

 Mimulus                             None (g)

Solanaceae
 Nicotiana                           Hummingbirds (g, l)

 Solanum




Urticaceae
 Boehmeria                           None (l)


 Parietaria

 Urtica





Verbenaceae
 Rhaphithamnus                       Hummingbirds (g, h, l, m)

Winteraceae
 Drimys                              None (l)





Family, genus (a)                    Pollination (d, e)

Apiaceae
 Apium

 Centella

 Eryngium
                                     Hummingbird



Asteraceae (i)
 Abrotanella

 Centaurodendron (k)
                                     Hummingbird


 Dendroseris (k)


                                     Hummingbird








 Erigeron




 Gamochaeta

 Lagenophora


 Robinsonia (k)







 Taraxacum




 Yunquea (k)

Berberidaceae
 Berberis


Boraginaceae
 Selkirkia (k)

Brassicaceae
 Cardamine


Campanulaceae
 Lobelia                             Hummingbird

 Wahlenbergia
                                     Selfer, wind







Caryophyllaceae
 Spergularia



Chenopodiaceae
 Chenopodium                         Wind



 Sarcocornia                         Wind


Convolvulaceae
 Calystegia

 Dichondra

Empetraceae
 Empetrum                            Wind

Ericaceae
 Pernettya                           Wind


Euphorbiaceae
 Dysopsis                            Wind

Fabaceae
 Sophora
                                     Hummingbird

Flacourtiaceae
 Azara                               Wind


Gunneraceae
 Gunnera                             Wind





Haloragaceae
 Haloragis                           Wind


Lamiaceae
 Cuminia (k)                         Hummingbird

Lactoridaceae
 Lactoris (k)                        Wind

Loranthaceae
 Notanthera                          Hummingbird

Myrtaceae
 Myrceugenia


 Myteola

 Ugni                                Wind

Piperaceae
 Peperomia                           Wind




Plantaginaceae
 Plantago                            Wind



Ranunculaceae
 Ranunculus

Rhamnaceae
 Colletia

Rosacese
 Acaena


 Margyracaena


 Margyricarpus


 Rubus


Rubiacese
 Coprosma                            Wind


 Galium


 Hedyotis


 Nertera                             Wind


Rutaceae
 Fagara


Santalaceae
 Santalum

Saxifragaceae
 Escallonia                          Hummingbird, selfer


Scrophulariaceae
 Euphrasia

 Mimulus

Solanaceae
 Nicotiana                           Hummingbird

 Solanum



                                     Selfer
Urticaceae
 Boehmeria                           Wind


 Parietaria                          Wind

 Urtica                              Wind





Verbenaceae
 Rhaphithamnus                       Hummingbird

Winteraceae
 Drimys                              Wind



                                     Pollination
                                     of
Family, genus (a)                    colonizers (f)

Apiaceae
 Apium                               Insect

 Centella                            Insect

 Eryngium                            Insect




Asteraceae (i)
 Abrotanella                         Insect

 Centaurodendron (k)                 Insect



 Dendroseris (k)                     Insect











 Erigeron                            Insect




 Gamochaeta                          Insect

 Lagenophora                         Insect


 Robinsonia (k)                      Insect







 Taraxacum                           Insect




 Yunquea (k)                         Insect

Berberidaceae
 Berberis                            Insect


Boraginaceae
 Selkirkia (k)                       Insect

Brassicaceae
 Cardamine                           Insect


Campanulaceae
 Lobelia                             Hummingbird

 Wahlenbergia                        Insect








Caryophyllaceae
 Spergularia                         Insect



Chenopodiaceae
 Chenopodium                         Wind



 Sarcocornia                         Wind


Convolvulaceae
 Calystegia                          Insect

 Dichondra                           Insect

Empetraceae
 Empetrum                            Wind

Ericaceae
 Pernettya                           Insect


Euphorbiaceae
 Dysopsis                            Insect

Fabaceae
 Sophora
                                     Hummingbird

Flacourtiaceae
 Azara                               Insect


Gunneraceae
 Gunnera                             Wind





Haloragaceae
 Haloragis                           Wind


Lamiaceae
 Cuminia (k)                         Hummingbird

Lactoridaceae
 Lactoris (k)                        Wind

Loranthaceae
 Notanthera                          Hummingbird

Myrtaceae
 Myrceugenia                         Insect


 Myteola                             Insect

 Ugni                                Insect

Piperaceae
 Peperomia                           Insect




Plantaginaceae
 Plantago                            Wind



Ranunculaceae
 Ranunculus                          Insect

Rhamnaceae
 Colletia                            Insect

Rosacese
 Acaena                              Insect


 Margyracaena                        Wind


 Margyricarpus                       Wind


 Rubus                               Insect


Rubiacese
 Coprosma                            Wind


 Galium                              Insect


 Hedyotis                            Insect


 Nertera                             Wind


Rutaceae
 Fagara                              Insect


Santalaceae
 Santalum                            Insect

Saxifragaceae
 Escallonia                          Hummingbird, insect


Scrophulariaceae
 Euphrasia                           Insect

 Mimulus                             Insect

Solanaceae
 Nicotiana                           Hummingbird, insect

 Solanum                             Insect




Urticaceae
 Boehmeria                           Wind


 Parietaria                          Wind

 Urtica                              Wind





Verbenaceae
 Rhaphithamnus                       Insect

Winteraceae
 Drimys                              Insect


Monocots

                                          Species (b)


Family, genus (a)                    Native

Arecaceae
 Juania (k)

Bromeliaceae
 Greigia


 Ochagavia (k)

Cyperaceae
 Carex, Cyperus, Eleocharis,         9 spp.
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia
                                     L. chilensis
Juncaceae
 Juncus
                                     J. cappillaceus
                                     J. imbricatus
                                     J. pallescens
                                     J. planifolius
                                     J. procerus

 Luzula                              [_NOT REPRODUCIBLE IN ASCII]

Orchidaceae
 Gavilea

Poaceae
 Agrostis, Bromus, Chaetotropis,     10 spp.
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum

                                           Species (b)


Family, genus (a)                    Endemic

Arecaceae
 Juania (k)
                                     J. australis
Bromeliaceae
 Greigia
                                     G. berteroi

 Ochagavia (k)
                                     O. elegans
Cyperaceae
 Carex, Cyperus, Eleocharis,         4 spp.
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia

Juncaceae
 Juncus






 Luzula
                                     L. masafuerana
Orchidaceae
 Gavilea
                                     G. insularis
Poaceae
 Agrostis, Bromus, Chaetotropis,     5 spp.
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum




Family, genus (a)                     Habit

Arecaceae
 Juania (k)                          Tree

Bromeliaceae
 Greigia                             Shrub


 Ochagavia (k)                       Shrub

Cyperaceae
 Carex, Cyperus, Eleocharis,         Perennial herb
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            Perennial herb

Juncaceae
 Juncus                              Perennial herb






 Luzula                              Perennial herb

Orchidaceae
 Gavilea                             Perennial herb

Poaceae
 Agrostis, Bromus, Chaetotropis,     Annual or perennial herb
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum

                                              Flower

                                     Size, in mm
Family, genus (a)                    (category)

Arecaceae
 Juania (k)                          10 x 4 (medium)

Bromeliaceae
 Greigia                             30 x 10 (large)


 Ochagavia (k)                       35 x 10 (large)

Cyperaceae
 Carex, Cyperus, Eleocharis,         2.5 x 1.5 (very small)
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            15 x 20 (large)

Juncaceae
 Juncus                              4 x 2 (very small)






 Luzula                              2.5 x 2 (very small)

Orchidaceae
 Gavilea                             15 x 10 (large)

Poaceae
 Agrostis, Bromus, Chaetotropis,     2.5 x 1.5 (very small)
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum

                                                    Flower


Family, genus (a)                    Shape (c)        Color

Arecaceae
 Juania (k)                          Dish             Green

Bromeliaceae
 Greigia                             Tube


 Ochagavia (k)                       Tube             Red/violet

Cyperaceae
 Carex, Cyperus, Eleocharis,         Inconspicuous    Green/brown
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            Dish             White

Juncaceae
 Juncus                              Inconspicuous    Green/brown






 Luzula                              Inconspicuous    Green/brown

Orchidaceae
 Gavilea                             Flag             Yellow

Poaceae
 Agrostis, Bromus, Chaetotropis,     Inconspicuous    Green/brown
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum




Family, genus (a)                    Sex

Arecaceae
 Juania (k)                          Dioecious

Bromeliaceae
 Greigia                             Hermaphroditic


 Ochagavia (k)                       Hermaphroditic

Cyperaceae
 Carex, Cyperus, Eleocharis,         Hermaphroditic, 6 spp.;
  Macherina, Oreobolus, Scirpus,      monoecious, 7 spp.;
  Uncinia                             andromonecious, 1 sp.
Iridaceae
 Libertia                            Hermaphroditic

Juncaceae
 Juncus                              Hermaphroditic






 Luzula                              Hermaphroditic

Orchidaceae
 Gavilea                             Hermaphroditic

Poaceae
 Agrostis, Bromus, Chaetotropis,     Hermaphroditic, 10 spp.
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum




Family, genus (a)                    Dichogamy (d)

Arecaceae
 Juania (k)

Bromeliaceae
 Greigia


 Ochagavia (k)                       Protandry

Cyperaceae
 Carex, Cyperus, Eleocharis,
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia

Juncaceae
 Juncus






 Luzula

Orchidaceae
 Gavilea

Poaceae
 Agrostis, Bromus, Chaetotropis,
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum


                                     Breeding
                                     system                     Floral
Family, genus (a)                    determined (d)             reward

Arecaceae
 Juania (k)

Bromeliaceae
 Greigia                                                        Nectar


 Ochagavia (k)                                                  Nectar

Cyperaceae
 Carex, Cyperus, Eleocharis,
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                                                       Nectar

Juncaceae
 Juncus






 Luzula

Orchidaceae
 Gavilea                                                        Nectar

Poaceae
 Agrostis, Bromus, Chaetotropis,
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum



                                     Floral visitors
Family, genus (a)                    observed (d)

Arecaceae
 Juania (k)

Bromeliaceae
 Greigia                             Hummingbirds


 Ochagavia (k)                       Hummingbird (g)

Cyperaceae
 Carex, Cyperus, Eleocharis,
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            Files, (g, l) moths (g)

Juncaceae
 Juncus






 Luzula

Orchidaceae
 Gavilea

Poaceae
 Agrostis, Bromus, Chaetotropis,
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum




Family, genus (a)                    Pollination (d, e)

Arecaceae
 Juania (k)                          Wind

Bromeliaceae
 Greigia                             Hummingbird


 Ochagavia (k)                       Hummingbird

Cyperaceae
 Carex, Cyperus, Eleocharis,         Wind
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            Insect

Juncaceae
 Juncus                              Wind






 Luzula                              Wind

Orchidaceae
 Gavilea

Poaceae
 Agrostis, Bromus, Chaetotropis,     Wind
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum


                                     Pollination
                                     of
Family, genus (a)                    colonizers (f)

Arecaceae
 Juania (k)                          Insect

Bromeliaceae
 Greigia                             Hummingbird


 Ochagavia (k)                       Hummingbird

Cyperaceae
 Carex, Cyperus, Eleocharis,         Wind
  Macherina, Oreobolus, Scirpus,
  Uncinia
Iridaceae
 Libertia                            Insect

Juncaceae
 Juncus                              Wind






 Luzula                              Wind

Orchidaceae
 Gavilea                             Insect

Poaceae
 Agrostis, Bromus, Chaetotropis,     Wind
  Chusquea, Danthonia,
  Leptophyllachloa, Megalachne, (k)
  Nassella, Piptochaetium,
  Podophorus, (k) Trisetum

(a)Characteristics are recorded at the genus level unless noted. When
there is more than one species per genus, the general character state or
average is given; however, the distinctly different species are also
noted.

(b)In the species columns, empty cells indicate that no species of the
kind are known for the respective genus; varieties are only included
when they are the only representative of a taxon in the archipelago.

(c)Flower-shape categories are after Faegri and van der Pijl (1979). For
the Asteraceae, the type of capitulum is also shown, in parentheses.

(d)Empty cells indicate missing information.

(e)Includes conclusions for species for which data or literature offer
reasonable evidence.

(f)Hypothetical, based on what is known from the literature on the
pollination of close relatives.

(g)Observed by Skottsberg (1928).

(h)Observed by Meza (1988).

(i)Observed by Johow (1896).

(j)For the Asteraceae, data represent the size of the capitulum.

(k)Endemic genera.

(l)Observed by the authors.

(m)Observed by Brooke (1987).

(n)From Moore (1983).

(o)From Ramanna and Hermsen (1981).
Table II

Statistical tests (chi-square and general association
coefficient) of features of the Juan Fernandez flora that were
positively associated. In all cases degrees of freedom = 1. p =
probability for both chi-square ([x.sup.2]) and general association
coefficient.

                                                 General
                                               association
Feature                             [x.sup.2]  coefficient     p

Very small flowers and perennial      31.65       31.44     <.0001
 herbs
Hermaphroditic and white flowers      13.17       13.07     0.0003
Monoecious and green flowers          17.21       17.09     <.0001
Inconspicuous and green flowers       48.50       48.17     <.0001
Bell-shaped and white flowers         12.07       11.95     0.0005
Very small and green flowers          50.09       49.77     <.0001
Inconspicuous and very small         118.47      117.69     <.0001
 flowers
Perennial herbs and current wind      13.96       13.80     0.0002
 pollination
Large- or medium-sized flowers and    25.50       25.20     <.0001
 and current bird pollination
Very small flowers and current        24.49       24.21     <.0001
 wind pollination
Very small or small flowers and       40.27       39.81     <.0001
 current wind pollination
Inconspicuous flowers and current     27.41       27.10     <.0001
 wind pollination
Bright-colored flowers and current    59.16       58.44     <.0001
 bird pollination
Green flowers and current wind         6.22        6.14       0.01
 pollination
Green or brown flowers and current    47.03       46.53     <.0001
 wind pollination
Trees and wind pollination of          4.76        4.72       0.03
 colonizers
Very small flowers and wind           36.31       35.88     <.0001
 pollination of colonizers
Inconspicuous flowers and wind        48.43       47.86     <.0001
 pollination of colonizers
Dish-shaped flowers and insect        26.51       26.20     <.0001
 pollination of colonizers
White flowers and insect              24.77       24.46     <.0001
 pollination of colonizers
Bright-colored flowers and bird       44.82       44.27     <.0001
 pollination of colonizers
Green flowers and wind pollination    31.02       30.65     <.0001
 of colonizers