The islands of Fiji are home to about 25 palmspecies, at least 84% of which are endemic tothe island nation. At the generic level, Fiji’spalm flora shares its closest relationship withVanuatu (60%), which is well above its shared

distributions with the Solomon Islands (27%),the Bismarck Archipelago (25%), and Samoa(15%; Watling 2005). For eight (75%) of Fiji’sindigenous palm genera, Fiji is the southern oreastern limit of their ranges. Fiji has a native

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The Palmsof Fiji andTonga

CHRISTINE D. BACON

Department of BiologyColorado State UniversityFort Collins, CO 80523 USAcbacon@rams.colostate.edu

Although not the richest in the region, the palm floras of both Fiji and Tonga

are very exciting and were the focus of a recent collecting expedition in November

of 2009.

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1. Pritchardiapacifica incultivation atUniversity ofthe SouthPacific, Fiji.

palm flora comprising 11 genera – Alsmithia,Balaka, Calamus, Clinostigma, Cyphosperma,Heterospathe, Hydriastele, Metroxylon, Neo-veitchia, Physokentia, Pritchardia and Veitchia.

The islands of Tonga are mostly coralline inorigin, rather than volcanic (Whistler 1992).Tonga has a very similar flora as its northernneighbor, Samoa, because of their geographicproximity and both are also part of the “FijianRegion” that extends from the Santa CruzIslands and Vanuatu to Niue (Takhtajan 1969).Veitchia joannis, Pritchardia pacifica and P.thurstonii are the only native palms in Tonga.Areca catechu, Cocos nucifera, Livistona chinensis,Metroxylon warburgii, Pinanga coronata andPtychosperma macarthurii are commonlycultivated in Tonga, as they are in Fiji.

Pritchardia

The primary goal of our expedition was tosample Pritchardia pacifica (Fig. 1) in both Fijiand Tonga. Pritchardia pacifica is an interestingspecies because it is known only fromcultivation and appears to grow strictly aroundhuman activity. It is found from the Marshall

Islands in the northwest Pacific, south to theSolomon Islands, Vanuatu, Fiji and Tonga, andfurther east to Niue, Samoa and the TuamotuIslands of French Polynesia. It is a spectacularpalm with a rich history. Many legendssurround the palm, but dispute about its originremains, with Tonga as the most commonlycited origin (Watling 2005). We sampledindividuals of P. pacifica from across the mainisland of Tonga, Tongatapu, as well as on themain island of Fiji, Viti Levu, to test thehypothesis of a Tongan origin. We also plan togather samples from across the Pacific and usemolecular phylogenetics to test the historicalexpansion of the species. One potential issuewith testing hypotheses about P. pacifica is thatis has most probably been transported fromisland to island by sea-faring Polynesians,therefore the molecular results would trackhuman movement rather than rangeexpansion from seed dispersal and islandcolonization. One way to address this wouldbe to create alternative hypotheses of humanmovement through the Pacific and testwhether the palms follow human movementand colonization.

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2. Pritchardia thurstonii population at bottom center of coastal cliff in the ‘Eua National Park, ‘Eua Island,Tonga.

The second goal of our expedition was tocollect P. thurstonii (Fig. 2) in its native habitat.Hodel (2007) recently recognized populationsgrowing on ‘Eua (Tonga), on the Sovu islets inthe northern Lau island group (Fiji) and onOgea and Vulaga of the Southern Lau group(Fiji) as belonging to this species. Pritchardiathurstonii is an interesting species because in atribal-level analysis (Bacon et al., in prep.) itwas resolved as sister to the Hawaiian radiationof approximately 26 currently recognizedspecies (Hodel 2007). Study of P. thurstonii andother South Pacific Pritchardia species helps usunderstand what the historical distributionsof the genus were and potentially what theHawaiian colonizers looked like with respectto seed size and other morphologicalattributes.

Moreover, P. thurstonii was sampled in the fieldat the population level. Sampling at this levelis important because populations on bothTonga and Fiji are not as severely affected byrat predation on seeds in comparison toHawaii, where the Hawaiian species are barelyable to self-replace due to the devastation ofseed banks by rats. Our collections may helpestablish a baseline estimate of genetic

diversity held within and between populationsand can further be compared to Hawaiianestimates to quantify endangerment andpossibly identify populations of specialconservation concern.

Morphological characters used to distinguishP. pacifica and P. thurstonii include the lengthof the inflorescences in comparison to the leafpetiole, the number of inflorescences that areborn on the tree and their insertion in thecrown, and presence of lepidia on the abaxialsurface of the leaf (Watling 2005, Hodel 2007).Sampling both Pritchardia species is essential totest species boundaries using molecularevidence, which is one goal of my dissertationwork.

In Tonga, P. thurstonii is found in the ‘EuaNational Park on the island of ‘Eua, which isan easy excursion from the main island ofTongatapu, where the capital city Nuku’alofais located. Either a three-hour boat ferry or aneight-minute flight gets one to ‘Eua, where afour-wheel drive and National Park guide canbe hired to access the populations. ‘Eua isgeologically different from the rest of theTongan islands and it appears to besignificantly older (Whistler 1992). Thenational park is located on the southeasternescarpment of the island. There are

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3. Balaka microcarpa from Colo-i-Suva National ForestPark, outside of Suva, Fiji.

4. Veitchia vitiensis from Colo-i-Suva National ForestPark, outside of Suva, Fiji.

approximately 250 P. thurstonii individuals,including juveniles, in two populationsseparated by steep cliffs and ocean. We wereable to sample the largest population, but theother was largely inaccessible, and our guideflatly refused to help navigate to thepopulation. The smaller population was seenfrom the southernmost end of the mainpopulation (Fig. 2) and was estimated toconsist of approximately 85 individuals.

We found two individuals of P. thurstoniioutside of the ‘Eua National Park, one in thenorthern extreme of the island and the otherin the southwest. Upon visiting both of thesesingle trees, we observed that they weregrowing at the mouths of caves. In the north,the cave was small and near the top of the seacliffs, whereas the tree in the southwest wasgrowing near a large cave system that forms atthe base of the sea cliffs. Human remains havebeen found in both the above-mentionedTongan caves. In Hawaiian traditional culture,people placed the remains of their ancestors incaves on sea cliffs, an important detail becausecustoms between islands in the Pacific arehypothesized to be related because of theircommon Polynesian ancestors. So why arethese Pritchardia growing at the mouths ofthese caves that hold human remains? Wespeculated that Pritchardia was deliberatelyplanted at the mouths of caves. But why? Weasked a ‘Eua shaman elder whether anytraditional uses for P. thurstonii were known,

and he explained that the stem of P. thurstoniiwas crushed and the liquid derived from it wasdrank to keep “death spirits” away. Of course,much of the traditional meaning of local plantuse can be lost in translation and throughmodernization of island peoples. We do notknow if the traditional use of the palmaccounted for its presence at the mouth of thecaves. We can only report that the associationbetween palms and sea caves seems to be morethan just coincidental.

Due to economic misfortune, the nationalFijian airline recently went out of business, sowe were unable to visit the populations of P.thurstonii in the Sovu Islets of Fiji. The onlycurrent means of traveling there is a ferry thatleaves once a month, although it has not beenrunning reliably since September 2009. Insteadof collecting P. thurstonii as we had originallyplanned, we turned our attention to collectingas many palm species on the main Fijian islandof Viti Levu as possible.

Fijian palms

Our Fijian expedition focused on higherelevation palms found in the mountain rangewhere the highest peak, Mt. Victoria, is located.Viti Levu, the largest island in Fiji, has thegreatest number of species among all theislands of Fiji because larger islands harborlarger numbers of species and because thetopographic variability provides for greaterhabitat diversity. Viti Levu has experienced arecent boom in tourism, and many foresttracks along the popular southern Coral Coast

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5. Flowers of Physokentia petiolata from the TomaniiviForest Reserve, near Navai village, on Viti Levu, Fiji.

6. Balaka longirostris flowers from the TomaniiviForest Reserve, near Navai village, on Viti Levu, Fiji.

area have been highly affected by developmentand deforestation. This human activity hascaused populations of Balaka longirostris,Cyphosperma sp. nov. ‘naboutini,’ Heterospathephillipsii, H. vitiensis, Metroxylon vitiensis,Physokentia petiolata, Veitchia joannis and V.vitiensis to decline and some to near extinction(Johnson 1996, Watling 2005). Forest usage isprevalent across the whole island, with localvillages using the understory and forest edgesfor plantations of taro, sweet potato, manioc,banana and guava, amongst many others. Inthe northern areas, development is less severebecause the King’s Road is not paved in someareas and the coast is less desirable for resortspeculators.

We spent a day northeast of the Fijian capitalof Suva in the Colo-i-Suva National Forest Park.Established in 1872, the Colo-i-Suva isrenowned for its easy access from Suva, itsverdant rainforests and its bird diversity. TheColo-i-Suva has about 4.5 km of trails and aseries of natural waterfalls formed by theWaisila Creek (upper Waimanu River). Thepark was planted with African mahogany inthe 1940s and 1950s, destroying some areas ofthe park, but some areas are still very nicely

preserved. We were rewarded with two palmspecies there, Balaka microcarpa (Fig. 3) andVeitchia vitiensis (Fig. 4).

After two days of driving, we conducted fieldwork from the Navai Village at the foot of Fiji’shighest peak, Mt. Victoria. Near Navai, we sawthe endemic Cyphosperma tanga, a criticallyendangered palm of which the Navai villageis very proud (see Back Cover). ScatteredVeitchia vitiensis individuals were found closeto the C. tanga population. In assessing the C.tanga population, we recorded many seedlingsand juveniles, so even though there is onlyone population remaining, it has apparentlybeen able to maintain its numbers.

In the book, Palms of the Fiji Islands, Watling(2005) reported a legend about how C. tangagot to Navai, which we confirmed with manyof the village elders. The story goes that twochiefs were traveling in the Navai area with awhale’s tooth (tabua) wrapped in the leaves ofthe C. tanga, locally known as taqwa. Theystopped to rest and decided to bury thewrapped sacred tooth. From that exact spotgrew the taqwa palm, and today it is onlyfound in that one locality. Traditionally in Fijitabua was made from teeth from the upper

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7. Emergent Clinostigma exorrhizum population in the Tomaniivi Forest Reserve, near Navai village, on VitiLevu, Fiji.

jaw of a sperm whale and are regarded asperhaps the most important cultural items inFijian society. When the practice became morewidely known in the early 1800s thousandsof fake teeth made from ivory and walrus tuskscame on the market, and mass-production ledto the development of the European art ofscrimshaw.

On the next day collecting from Navai villagewe set out to find the majestic Clinostigmaexorrhizum populations. Scaling steep terrain inthe upper montane forest of the TomaniiviNature Reserve, we were amazed by the skill ofour guide and his dogs, which were able tohunt a wild piglet without any spoken wordsor weapons. Along the trail we collectedPhysokentia petiolata (Fig. 5), distinctive with itsround and dark black fruits, and Balakalongirostris (Fig. 6), which the local Navaivillagers use to make spears for hunting.Nearing the top of the ridge we came upon thebeautiful C. exorrhizum (Fig. 7), as it usuallyfound only above 700 m in elevation. This isthe largest palm I had ever collected and standsaround 20 m in height. None of theindividuals we were able to collect had maturefruits, but it has been reported that oneindividual infructescence can produce up to14,000 seeds (Watling, 2005).

Overall, the trip was a success and importantcollections were made for my dissertationresearch. My specific goals for the materialcollected from this field work are to discovernot only the origin of the enigmatic Pritchardiapacifica, but also test species boundaries in thegenus, as well as look at patterns of geneticdiversity and how it is organized on the

landscape. It is from field work such as this,that biologists like myself get lots of ideas andI look forward to report more results in futurereports. I also collected material for futureprojects on the biogeography of Pacific palms.The biogeography of this region is extremelyinteresting with respect to patterns ofdiversification and adaptive radiation, andthere is still so much to uncover.

Acknowledgments

Work in Fiji was done in collaboration withMarika Tuiwawa and Alifereti Naikatini of thePacific Regional Herbarium at the Universityof the South Pacific in Suva, Fiji. In Tonga,Liuaki Fusitu’a from the Ministry of Educationand Leody Vainikolo from the Ministry ofAgriculture facilitated the permitting process.Iliki from Navai village (Fiji), Jo from ‘Eua(Tonga), and Geovanni Romero assisted withthe field collections. This research wassupported by the Montgomery BotanicalCenter.

LITERATURE CITED

HODEL, D.R. 2007. A review of the genusPritchardia. Palms Supplement to 51: S1–S52.

TAKHTAJAN, A.L. 1969. Flowering Plants: Originand Dispersal. Oliver & Boyd, Edinburgh.

WATLING D. 2005. Palms of the Fiji Islands.Environmental Consultants Ltd., Suva, Fiji.

WHISTLER, W.A. 1989. The unique flowers ofPolynesia: Tonga. Bulletin of the NationalTropical Botanical Garden 19: 81–84.

WHISTLER, W.A. 1992. Vegetation of Samoa andTonga. Pacific Science 46: 159–178.

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PALM BOOK AND JOURNAL COLLECTION FOR SALE

An exceptionally complete private assemblage of books, journals and other materials isoffered for private sale as a collection. It is comprised of 1,300 books in English and 20 foreignlanguages. Notable titles are the Barbosa Rodrigues Sertum Palmarum Brasiliensium, two-volume folio of 1903, with 174 chromolithograph plates; Kerchove de Denterghem’s classicwork Les Palmiers (1878); an fine copy of Seemann’s Popular History of the Palms (1856) inits original binding; Martius’s Palmarum familias ejusque general (1824) and Steck’s 1757 studySagu. Also included is a nearly complete collection of the palm publications of OdoardoBeccari. All the standard works on palms of the past century are in the collection, includingpalm floras and palm conference proceedings. Economic species books are a particularcollection strength: coconut (254), oil (167), date (132), rattans (98) and sago (30). Amongthe 70 journals are full runs of Principes/Palms; Palms & Cycads (Australia) and The PalmJournal (California) and journal runs of economic palms. A third component consists of 800items made up of journal reprints on palms, pamphlets, maps, microfiches, CD-ROMs andvideos. The collection would be an ideal resource for any institution researching palms. APDF file of the items in the collection is available. Contact Dennis Johnson; telephone 513-631-8766 or email djohn37@aol.com.