Number of tribes, genera and species of Poaceae in Easter Island
Rapa Nui (Easter Island, Isla de Pascua), also known as Te Pito O Te Henua, is a small oceanic island of volcanic origin discovered by the Dutch explorer Jakob Roggeveen in April 1722. It has belonged to Chile since 1888 and is administratively part of the Region of Valparaíso, Province Isla de Pascua. At around 163.6 km2, it is the largest island of the Chilean insular territory, situated in Polynesia,
The climate is warm and sub-tropical. The flora of Rapa Nui is extremely poor compared to other oceanic tropical islands . Approximately 40 % of the flora is indigenous. Nearly 23 % of the vascular flora is represented by endemic species, and some 20 species of the native flora, 10 of which are endemic, have disappeared or are endangered, principally due to invasive plants, fire, overgrazing and agriculture, among other factors . Nearly 90 % of the territory corresponds to herbaceous vegetation, with species of Poaceae (grasses) as the principal component, most of them alien . There are, however, very dense little forests composed of species that have come with human beings since the island was colonized. Wetlands are located chiefly in the craters of volcanoes, the largest in Rano Kau (Fig. 1) and others in Ranu Raraku and Ranu Aroi. The flora of these wetlands consists of
Grasslands present on the island can be divided into two types: 1. Very low grasslands, with high species diversity, overgrazed by livestock, especially horses. 2. Higher grasslands composed almost exclusively of
The number of species reported for the island is not consistent in the scientific literature. Most of the differences in number of species occur, probably, because the authors include or exclude cultivated plants, and due to synonyms and nomenclatural changes.
The aim of this chapter is to provide a synopsis of the diversity of the family Poaceae (Gramineae) in Rapa Nui, to provide a catalogue of all species of Poaceae and to analyse the completeness of the inventory, to analyse the taxonomic distribution, life cycle, photosynthetic pathway, and phytogeographical origin of Poaceae in Rapa Nui and to compare the diversity of Poaceae in Rapa Nui with those of other oceanic islands. To date, the most complete list of grass species published on the flora of Rapa Nui comprises 46 species . Our data indicate that in Rapa Nui, the family Poaceae comprises 50 species and one infraspecific taxon, from 37 genera and seven subfamilies. Recent taxonomic treatments have followed to update the nomenclature and the classification of the species.
2. History of the botanical expeditions to Rapa Nui
The first botanical collections made in Rapa Nui were those of Johann Reinhold Forster and his son Georg, during Cook’s second voyage aboard the “Resolution”, who sighted Rapa Nui on 13th March, 1774, and the next day landed at Hanga Roa . Both Cook and Forster made similar and interesting comments, mainly on crop species such as sugarcane, potatoes and bananas. They also mentioned
During a voyage of the Russian ship “Rurik”, commanded by Captain Kotzebue, and with Adelbert von Chamisso, a naturalist, on board, it passed by Rapa Nui on a short visit; apparently, a small botanical collection was made . Between 1825 and 1828, Captain Beechey’s journey of exploration in the “Blossom” visited several locations in Chile, of which an important record is retained in the publications of Hooker and Arnott (1830 and 1832). The ship arrived at Rapa Nui on 16th November, 1825, but there are no records of plants .
Endlicher  in
Savatier, during the campaign of the “Magicienne”, reached Rapa Nui in August, 1877. Plants collected on the island are kept in the herbarium of the Museum of Paris, but no list was published. In 1885, Hemsley, in his
In 1911, Francisco Fuentes was commissioned by the Chilean government to conduct a study on Rapa Nui. As a result of this work, Fuentes published his
Between 5th October, 1916, and 26th, September, 1917, Skottsberg and his wife made a major Swedish expedition to explore and study the Juan Fernández Archipelago and Rapa Nui, which culminated in the publication of
The Franco-Belgian mission exploring Rapa Nui from 29th July, 1934, to 3rd January, 1935, collected 61 species; the few that were not reported previously are obviously introduced, and known angiosperms number 142 . The list mentions nine species of grasses, some new with respect to the Skottsberg list, and others that were missing, and notes that there are numerous plants without flowers to identify.
Subsequently, several researchers have conducted the collection of plants, which have been deposited in various herbaria, especially in Chile in the herbarium of the National Museum of Natural History (SGO) and the herbarium of the University of Concepción (CONC) . In this work, 46 grass species have been reported.
3. Material and methods
Specimens were collected and preserved in the herbarium of the National Museum of Natural History at Santiago (SGO). Specimens were identified and photographs were taken using a Zeiss Stemi 2000 C stereomicroscope equipped with an Axiocam ERc5s camera. Images were processed with the software Zen 2011.
A database of the species of grasses of Rapa Nui was constructed, based on the databases of two important Chilean herbaria: CONC (Herbarium of the University of Concepción) and SGO (Herbarium of the National Museum of Natural History, Santiago). Specimens deposited in these herbaria and those collected for this project were included. The database contains the following fields: 1. Genus; 2. Species; 3. Common names in Rapa Nui; 4. Origin (native/introduced/endemic); 4. Geographical origin; 5. Photosynthetic pathway; 6. Life cycle; 7. Subfamily; 8. Tribe; 9. Collector’s name; 10. Collector’s number; 11. Latitude; 12. Longitude; 13. Altitude; 14. Locality; 15. Date of collection (year); 16. Date (year) of first registration; 17. Herbarium; 18. Herbarium number; 19. Bibliographic citations. A total of 369 specimens were included.
A checklist is provided, including Latin name, origin (endemic, native, introduced), homeland, life cycle (annual, perennial, annual or perennial), photosynthetic pathway (C3/C4) and classification (subfamily, tribe); however, the biogeographic status is sometimes difficult or impossible to establish. Meyer’s secondarization index was calculated as the number of native species/number of naturalized species .
The diversity of grasses of Rapa Nui was compared with the diversity of grasses of other oceanic islands (Galápagos, Pitcairn, Marquesas, Juan Fernández and Hawaii), using the regional diversity index (D). This index was calculated as D=S/log A, where S is the number of species in the region and A is the area in square kilometres . The floristic affinity between these islands was compared by cluster analysis of presence-absence data for 349 species, using Jaccard coefficient as the similarity measure, UPGMA algorithm and the statistical software Infostat . Species composition was taken from the literature [24-28]. Species accumulation curve and estimated richness was calculated using the software Estimates 8.0 .
Our database for the island included a total of 369 specimens collected over 12 decades (1900-2013), representing 51 species, 37 genera, 11 tribes and seven subfamilies (Table 1 and 2), that is, approximately 10 % of the total Chilean (continental and insular) grass flora (523 species and 57 infraspecific taxa) . The proportion of species relative to the number of genera is 1.36, similar to the proportion determined by Fuentes  for the entire flora of the island (135 species / 104 genera=1.29). Most of the genera are represented only by one or two species,
Only two species (3.9 %) of the family Poaceae are endemic to Rapa Nui (
Although there are relatively few botanical specimens of Poaceae collected in Rapa Nui, the species cited in the botanical literature are well represented in Chilean herbaria. Moreover, only one species was collected for the first time in 2013, suggesting that the inventory of species is fairly comprehensive. The first herbarium specimens entered into our database correspond to those made by Alexander Agassiz, who in 1904 collected 16 specimens representing 12 different species, about 20 % of the currently known diversity of Poaceae in Rapa Nui. By the middle of the 20th century, with the botanical expeditions made by Fuentes, Skottsberg, Balfour, Williamson & Co., Drapkin and the Mission Franco-Belge, the number of known species reached nearly 50 % of the currently known species number (Fig. 5). An important increase in the number of known species occurred after the botanical trips made by Michel Etienne, who published 24 wild and two cultivated species of Poaceae in Rapa Nui  and by Georg Zizka, who reported 46 species of grasses, the most comprehensive list until today [13, 14]. In the decade 1981-1990, a total of 149 specimens of Poaceae were collected, most of them by Zizka. In general, the herbarium collections of Poaceae from Rapa Nui are limited. Our database for the island included a total of 369 specimens over 12 decades (1900-2013), including a total of 51 taxa, most of which were known previously , and in Zizka’s [13, 14] papers.
Most of the species of grasses of the island are introduced, some of them cited very early in the botanical literature, such as
In 1911, Francisco Fuentes collected, among other plants, a specimen published as the holotype of
5. Taxonomic distribution of Poaceae in Rapa Nui
The taxonomic distribution of the species of Poaceae in Rapa Nui is shown in Table 1. Seven subfamilies are represented. In continental Chile, eight subfamilies are present , of which only species of the subfamily Aristidoideae are absent in Rapa Nui.
The subfamily Arundinoideae comprises only one species:
The subfamily Bambusoideae is represented only by cultivated bamboos of the genus
The subfamily Chloridoideae comprises six genera and six species in Rapa Nui (Table 1), all belonging to the subtribe Cynodonteae, most introduced from Tropical Africa, such as
The subfamily Danthonioideae comprises only two species in Rapa Nui. One of the species was mentioned only once in the literature dealing with Rapa Nui flora , under the name
The only species of the subfamily Ehrhartoideae known in Rapa Nui is
The subfamily Panicoideae comprises 12 genera and 23 species, representing about 44 % of the grass flora of Rapa Nui (Table 1). This clearly contrasts with the total grass flora of Chile, where Panicoideae represents only
Panicoideae includes the second endemism of this family from the island,
The subfamily Pooideae comprises 16 species from 15 genera, being the second most diverse subfamily. Whereas Pooideae include most of the grasses of the Chilean flora (74.65 %) , in Rapa Nui it represents only
6. Phytogeographical origin of Poaceae in Rapa Nui
As in other oceanic islands, grasses are the most common alien plants occurring in Rapa Nui [3, 33, 35]. The number of endemic, native and alien species of the grass flora in the island is shown in Fig. 6, where we can see that alien plants represent the vast majority of the grass flora (Meyer’s secondarization index=0.24). Introduced species are mainly of African, European and Asian origin (Fig. 7, Table 2).
The proportion of alien species in six Poaceae subfamilies (Bambusoideae was not included as it contains only one cultivated species) is shown in Figure 8. All Chlorodoideae seem to be alien, most of African origin. However, the identity of some species, mainly those of
Most of the species of subfamily Panicoideae and subfamily Pooideae, which constitute the bulk of the grass flora of the island, are alien. The only species of subfamily Ehrhartoideae,
Rapa Nui belongs to the Polynesian Biogeographic Province  or Polynesian Floristic Region , included among the 25 biodiversity hotspots of the world [38, 39]. Specifically, it belongs to the Eastern Polynesia Subregion of Polynesia and represents the driest island of the subregion, with 1,325 mm of annual precipitation. In effect, there is no native vegetation on the island today and introduced plants outnumber native species due to the deforestation that occurred soon after the arrival of the first Polynesian inhabitants (38).
Introduced species that became naturalized or invasive represent one of the major threats to native species. It has been proposed that aboriginal people significantly modified the vegetation of the island  and the original vegetation communities were replaced by grasslands. In these plant communities, introduced species became increasingly abundant . According to Aldén , the composition of the flora underwent a rapid change from the 18th century, when European people begin to visit the island.
On the other hand, around 70,000 tourists visit the island each year, causing environmental deterioration . As shown in Figure 9, a sharp increase in the amount of alien species occurs in the 1990s; by this time, there are three times the number of alien species present compared to when Francisco Fuentes visited the island in 1911; nevertheless, in the 1910s, aliens already exceed the number of native species. Zizka  collected six species of grasses for the first time, all of them introduced. In some cases, nevertheless, we cannot be absolutely sure if certain species are indigenous or were accidentally introduced by man.
From a physiological point of view, introduced species mostly show C4 photosynthesis (Fig. 10), and most of the species, both alien and native, are perennial (Fig. 11). It has been demonstrated that alien species distantly related to the native flora are more likely to become harmful weeds for regional ecosystems, supporting Darwin’s naturalization hypothesis; thus, special attention should be paid to newly introduced species for which there are no close relatives in the regional flora . In Rapa Nui, our data show that species from 30 genera and two subfamilies that do not include native species have been introduced.
7. Comparing the diversity of Poaceae in Rapa Nui with other Pacific Islands
The diversity of grasses in Rapa Nui, calculated as the number of species per area expressed in square kilometres (regional diversity index) is slightly lower than that of the Juan Fernández Archipelago (Fig. 12). According to the literature, Poaceae in Juan Fernández Archipelago comprises 32 genera and 53 species [33, 43]. This number is relatively small compared with Hawaii (216 species) and Galápagos (94 species). From Desventuradas islands (San Félix, San Ambrosio), only two species have been recorded (
|16.||N||Australia, Asia & Pacific||C3||P||Pooideae||Poeae|
|17.||N||Australia, Asia & Pacific||C3||P||Pooideae||Poeae|
|19.||I||Australia, Asia & Pacific||C4||A||Panicoideae||Paniceae|
|26.||N||New Zealand, Australia, New Guinea, Rapa Nui||C3||A/P||Pooideae||Poeae|
|49.||I||Europe, Asia, Africa||C3||A||Pooideae||Poeae|
|50.||I||Europe, Asia, Africa||C3||A||Pooideae||Poeae|
8. Taxonomic sampling effort
The collection effort (sampling) is an important part of the taxonomic work on which the knowledge of species richness and, ultimately, the knowledge of biodiversity are based. Although collectors try to find all the species in a region, this goal is almost impossible, or at least very difficult to achieve; thus, the real number of species can only be estimated from the number of observed (collected) species (48). As shown in Table 3 and Figure 14, for a total of 50 observed species, the species richness estimated by different estimators ranges from 58.3 (Bootstrap) to 86.8 (Chao). An increased collection effort for Poaceae in Rapa Nui could yield between eight and 36 additional hitherto unsampled species. As shown in Fig. 15, collections are concentrated only in a few localities, chiefly in Rano Kau, near Hanga Roa, Anakena, and Rano Raraku.
|Sobs (Mao Tau)||50|
9. Concluding remarks
The floras of the oceanic islands are especially prone to serious threats from alien invaders , because they have a propensity to include highly adapted specialist rather than generalist species . This is of particular interest, as these ecosystems comprise high numbers of endemic plants, in contrast with continental regions of similar size . For these reasons, it is necessary to have on hand complete lists of the flora indicating alien plants that could be agricultural weeds and invasive species that can put pressure on native ecosystems.
Poaceae contains an important number of species that behave as weeds in natural environments (invasive), as well as in ruderal and agricultural habitats all over the world. In Rapa Nui, grasses are the most diverse family of vascular plants; nearly 90 % of the island is covered by grasslands and alien grasses represent nearly 80 % of the Poaceae of the island. A similar situation occurs in other oceanic islands . Several alien species introduced to Rapa Nui are noxious weeds (
On the other hand, only two Poaceae endemic to Rapa Nui are recognized. Another two Poaceae considered endemic in previous studies have been the object of taxonomic studies that demonstrate their non-endemic status:
Herbarium specimens provide valuable information to appreciate regional plant diversity, as well as to understand plant invasions and geography [46, 47]. A wide array of data (phenology, flowering periodicity, distribution, altitude, morphometry, minimum residence time, new distributional records, etc.), that are important to biodiversity monitoring can be obtained from herbarium specimens . However, herbarium collections in Rapa Nui are relatively scarce.