First record of the Gnetales in Australia: Gnetum gnemon L. (Gnetaceae) on Badu and Mua islands, Torres Strait, Queensland David G. Fell¹, David J. Stanton, Dick Williams, Frank Loban, Frank Nona, Troy Stow, John Wigness, Erimiah Manas & Guyai Uiduldam Summary Fell, D.G., Stanton, D.J., Williams, D., Loban, F., Nona, F., Stow, T., Wigness, J., Manas E., and Uiduldam G. (2015). First record of the Gnetales in Australia: Gnetum gnemon L. (Gnetaceae) on Badu and Mua Islands, Torres Strait, Queensland. Austrobaileya 9(3): 421–430. The gymnosperm order Gnetales is reported for the first time in Australia, from specimens of Gnetum gnemon collected on two continental islands in Torres Strait, Queensland. The distribution, habitat, ecology and local conservation status of the species are discussed, and its origins in Torres Strait are considered with reference to biogeographic and anthropogenic factors. Key Words: Gnetaceae, Gnetum gnemon, Australia flora, Queensland flora, Badu Island, Mua Island, Torres Strait ¹David G. Fell, P.O. Box 337, Alstonville, NSW 2477, Australia. Email: [email protected] Introduction Gnetum L. is the sole genus within the Gnetum gnemon L. has a wide distribution family Gnetaceae and order Gnetales of the in Malesia and the Western Pacific and is gymnosperms. The genus consists of about 40 native to Indonesia, Papua New Guinea, the species with 10 species in South America, one Solomons, and Vanuatu (Verheij & Sukendar in tropical West Africa, and 20–25 species in 1991; Manner & Eleivich 2006; Wonn & tropical and subtropical Asia (Markgraf 1929, Renner 2006; Bourke 2010). In Papua New 1977; Maheshwari & Vasil 1961; Verheij & Guinea it occurs in the Morobe, Eastern Sukendar, 1991; Carlquist 1994, 1996; Price Highlands, Western Highlands, Central, 1996; Wonn & Renner 2006, Hou et al. 2015). East Sepik, Sandaun (West Sepik), Manus The origins and evolutionary relationships Island, New Ireland, Western, Bougainville, of the gnetophytes are considered as central New Britain, Madang, Milne Bay, and Gulf toward understanding the origin of flowers Provinces (Conn & Damas 2006). It is and seed plant evolution (Markgraf 1929; previously known in Australia only from Chamberlain 1935; Price 1996; Hansen et al. cultivated trees in the vicinity of Darwin 1999; Winter et al. 1999; Manner & Eleivich (Atlas of Living Australia, 2012; I. Cowie 2006). Recent genetic studies now recognise pers. comm., November 2012). that gnetophytes are more closely related to Gnetum gnemon is a dioecious evergreen conifers than to angiosperms (Hansen et tree 15–30 m in height in the canopy or al. 1999; Winter et al. 1999; Becker et al. subcanopy with a trunk diameter at breast 2000), and that the major lineages of Gnetum height to 40 cm (Peekel 1984; Conn & Damas diverged in the Late Cretaceous (Hou et al. 2006; Manner & Eleivich 2006). The trunk is 2015). smooth and cylindrical with prominent raised swollen bands (100–300 mm apart) caused by the growth and fall of the opposite and symmetrical branches (Conn & Damas 2006). Leaves are simple and opposite, elliptic, lanceolate or oblong-oval, 15–25 cm long and 5–9 cm wide (Manner & Eleivich 2006). As Accepted for publication 21 August 2015 a gymnosperm, Gnetum does not have true 422 Austrobaileya 9(3): 421–430 (2015) flowers; the cones or strobili are presented at fruit are identified as a source of protein across the tip of a slender stem or axis (Chamberlain the nation (Cordon 1970). Cultivation of trees 1935; Manner & Eleivich 2006). It has yellow is achieved by propagating seed and/or from single-seeded fruits which turn purple red or cuttings (French 1986). In the Kiunga area of orange red at maturity (Maheshwari & Vasil Western Province the bark is used to make the 1961; Manner & Eleivich 2006), ripening fibre for string bags and other products and between September and December (Bourke it is also cultivated as a food plant (leaves) et al. 2004). in Daru (Western Province) (B. Waterhouse pers. comm., April 2014). Across its Asian, Malesian and Western Pacific range the genus Gnetum is known We report on the occurrence of from tropical rainforest up to 1,700 m with Gnetum gnemon on two islands in Torres rainfall of 750–5,000 mm/year (Manner & Strait, Queensland, thus recognizing the Eleivich 2006). It occurs in primary and gymnosperm order Gnetales for the first time secondary vegetation, with cultivated trees from Australia. The distribution, habitat and commonly found in home orchards and ecology of the species at these localities are subsistence gardens. discussed together with an assessment of its local conservation status. Its origins in Wild and cultivated Gnetum gnemon is an Torres Strait are considered with reference to important natural resource in many parts of biogeographic and anthropogenic factors. Asia, Malesia and the Pacific where it occurs. Documented uses include: food (leaves Materials and methods for wrapping food items, young cones and A survey of the vegetation of the Torres Strait leaves cooked with meat for flavoring; seeds Islands, Queensland, Australia, was carried ground into a flour for fried flat cakes; flowers out in 2007 (Stanton et al. 2009). The survey’s (stroboli) eaten; young fruits eaten raw or primary objective was to map vegetation cooked; timber (poles for house construction, communities at a scale of 1:25,000 and tool handles, burned for fuel, and pulped for Regional Ecosystems at a scale of 1:50,000, papermaking); stem bark fibre (string bags, and was supplemented by floristic inventory ropes, bowstring on musical instruments, and collections of voucher specimens for construction of fishing lines and fishnets, Australian herbaria. Additional surveys on assembling arrowheads and arrowshafts); the Badu and Mua islands between 2009 agroforestry and rehabilitation (intercropping and 2015 have been carried out as part of a for rambutan and breadfruit, trellis for biodiversity management planning program Dioscorea yams, improving soil fertility, through the Land and Sea Management Unit windbreaks, boundary markers) (Henderson of the Torres Strait Regional Authority (3D & Hancock 1989; Verheij & Sukendar 1991; Environmental 2011a, 2011b; Gynther et al. Ohtsuksa 1983; Peekel 1984; Salim et al. 2002; 2014; Reis et al. 2015). Walter & Sam 2002; Manner & Eleivich 2006; Quartermain & Tomi 2010). Extracts derived Collections of Gnetum gnemon were from Gnetum are reported useful as health made at Badu Island in October 2007 (Fell supplements (Kato et al. 2009) and the nuts 10206 & Stanton) (Fig. 1) and on Mua Island form an important home industry throughout in April 2011 (Fell 10803 & Stanton) (Fig. Indonesia (Cadiz & Florido 2001). 2), with further confirmed observations on Mua in March 2014 and on Badu in May In Papua New Guinea it is an important 2015 (Fell pers. obs.). Voucher specimens are wild and cultivated food plant in many lodged with the Queensland Herbarium (BRI) locations (Powell 1976; Bourke 2004) and with duplicates to the Australian Tropical known in Tok pisin as ‘Tulip’ (two leaves) Herbarium (CNS). (French 1986). The Gidra people of the Oriomo River area of southwest Papua New Guinea adjacent to Torres Strait, eat the fruits and leaves (Ohtsuka 1983), and the leaves and Fell et all., Gnetum gnemon in Australia 423 Fig. 1. Leaves of Gnetum gnemon on Badu Island (Fell 10206 & Stanton). Photo: D.G. Fell. Results and discussion rock is overlain on its margins by younger unconsolidated deposits including alluvial Gnetum gnemon in Torres Strait deposits and extensive dune fields of varying Badu Island age and geomorphic expression (Willmott & Powell 1977; Garnett & Jackes 1983; 3D Badu Island is situated approximately midway Environmental 2011a). between the tip of Cape York Peninsula and mainland New Guinea, and belongs to the A total of 49 vegetation communities Near Western Group of Torres Strait Islands. within 20 broad vegetation groups and 32 It is a continental island of 10,467 ha, centred Regional Ecosystems are recognised across at 10° 07’ S, 142° 09’ E and located around the island, within which 592 plant species 92 km south of the Papua New Guinea (PNG) have been recorded. The flora comprises coastline and 70 km NW of Cape York (Map 560 native species with 17 ferns, one cycad, 1). The island is the homeland of the Badulgal one conifer and 572 flowering plants (3D people with a population of 784 people as Environmental 2011a). at the 2011 census (Australian Bureau of Mua Island Statistics 2012). Mua (also known as Moa) Island lies adjacent The coarse grained Badu granite and immediately to the east-south east of dominates the landscape, forming the island’s Badu, being separated by a narrow (2.5 km rugged interior of low rocky hillocks with wide) channel. Situated approximately 55 km massive granite boulders. Granite basement 424 Austrobaileya 9(3): 421–430 (2015) north of the tip of Cape York, Queensland and in the north-east, rising to 376 m. A rugged 94 km south of the New Guinea mainland east and south facing coastline features rocky (Map 1), it has a total area of 17,001 ha, coastal headlands, and an expansive coastal and is the second largest island in Torres plain forms a broad enclave behind the island’s Strait. There are two Torres Strait Islander north-eastern coastline. Vegetation is diverse communities on the island: at Kubin and St with a total of 62 vegetation communities Pauls, which had populations of 163 and 258 occurring within 23 broad vegetation groups respectively as at the 2011 census (Australian and 44 regional ecosystems. The flora is the Bureau of Statistics 2012). richest in the Torres Strait region with 609 native species, comprising 19 ferns, one Mua is formed on continental igneous cycad, two conifers and 654 flowering plants basement rocks and is topographically diverse, (3D Environmental 2012a). dominated by the high point of Banks Peak Fig. 2. Upland habitat of Gnetum gnemon on Banks Peak, Mua Island. Photo: D.J. Stanton. Local Habitat & Ecology The Badu occurrence is within a closed (swamp) forest characterised by poorly In Torres Strait Gnetum occurs in poorly drained soils which may be seasonally drained lowland swamp forest (Badu), inundated. Typical canopy tree species are lowland riparian rainforest (Mua), and on Carallia brachiata (Lour.) Merr., Deplanchea steep granitic hillslopes in well-developed tetraphylla (R.Br.) F.Muell., Lophostemon evergreen vineforest (Mua) (Fig. 3). suaveolens (Sol. ex Gaertn.) Peter G.Wilson Fell et all., Gnetum gnemon in Australia 425 including Acmenosperma claviflorum (Roxb.) Kausel., Anthocarapa nitidula (Benth.) T.D.Penn. ex Mabb., Calophyllum sil Lauterb., Manilkara kanosiensis H.J.Lan & B.Meeuse, Sterculia shillinglawii F.Muell. subsp. shillinglawii and Syzygium beuttnerianum (K.Schum.) Nied. Gnetum habitat on the steep southern slopes of the peak features a wind sheared canopy of Acmenosperma claviflorum, Calophyllum sil, Podocarpus grayae De Laub. Licuala ramsayi var. tuckeri Barfod & Dowe and Pandanus zea H.St. John. Riparian rainforest on alluvial sands and silts constitute Gnetum lowland habitat on Mua. Characteristic canopy species are Buchanania arborescens (Blume) Blume, Horsfieldia australiana S.T.Blake, Maranthes corymbosa Blume, Syzygium angophoroides and S.bamagense B.Hyland. No uses are documented or known for Gnetum gnemon in the Torres Strait. Specimens were shown to Land and Sea Rangers during our visits to Badu in November 2007, November 2010 and May Fig. 3. Multi stemmed trunk of Gnetum gnemon in lowland riparian rainforest on Mua Island. Photo: D.G. 2015, and on Mua in March 2011 and March Fell. 2014. They did not recognise the plant, and stated that they had no name or use for it (D. Williams pers. comm., 2015; T. Stow pers. & J.T.Waterh., Maranthes corymbosa Blume, comm., 2007; J. Wigness pers. comm., 2011). Melaleuca dealbata S.T. Blake and Syzygium angophoroides (F.Muell.) B.Hyland with a Natural or translocated? sharp ecotone to the surrounding woodland Some plants and their populations in vegetation. Torres Strait are a reflection of natural On Mua, Gnetum occurs in upland and biogeographic distributions, whereas others lowland situations. Upland habitats are on are anthropogenic and may indicate the the steep granite hillslopes and crests of influence of people as dispersal agents Banks Peak, the highest topographic feature (Denham 2008; Denham et al. 2009; McNiven in Torres Strait. Gnetum was recorded 2008). between approximately 250–350 m altitude The accepted hypothesis that the in evergreen notophyll vineforest and in pantropical range of Gnetum reflects a wind sheared evergreen vinethicket (Fig. Gondwanan history was established by 2). The evergreen vineforests are developed Markgraf (1929). However, Wonn & Renner on upper sheltered slopes where weathering (2006) consider that its present range follows of the granite has produced a well-drained a more recent radiation in the Malesian and relatively fertile sandy loam soil (3D region, and therefore is not Gondwanan. Environmental 2011b). The rainforest type Using analysis of fossil-calibrated molecular- is a unique and newly described ecosystem clocks, these authors investigated Gnetum for Queensland, restricted to only a few lineages now found in Africa, South America mountain top locations in Torres Strait (3D and Southeast Asia with results suggesting the Environmental 2009). The canopy height influence of ancient long-distance dispersal of ranges from 23 m to 35 m with dominants 426 Austrobaileya 9(3): 421–430 (2015) seeds across seawater. Further, its distribution the authors in 2007, 2009, 2011, 2014 and 2015 throughout Asia is thought to have occurred assessed the condition of the habitat as good, during radiation through Malesia where being free of weeds although diggings by feral opportunities for overland seed dispersal pigs (Sus scrofa) were observed throughout coincided with times of low sea levels (Wonn the habitat. The even age of the canopy & Renner 2006). and the species composition together with evidence of old charred stumps indicate that Torres Strait was formed by rising sea the habitat is transitional and subject to past levels in the Holocene (c. 8000–6500 years burning. Invasive weeds such as Singapore before present) that inundated the Arafura daisy (Sphagneticola trilobata (L.) Pruski), Plain, a low-lying land bridge that connected praxelis (Praxelis clematidea R.M.King & Australia and New Guinea for much of H.Rob.), annual mission grass (Pennisetum recent geological history (Jennings 1972; pedicellatum Trin. subsp. pedicellatum) Woodroffe et al. 2000). Given the occurrence and leucaena (Leucaena leucocephala of G. gnemon in neighboring areas of central- (Lam.) de Wit) occur on the islands, but are southern New Guinea, it is likely the Torres mostly associated with habitation and along Strait records are refugial populations that roadsides. Singapore daisy has successfully reflect its biogeographic distribution across invaded degraded swamp forests within and the Malesian region. on the margins of the Badu community and However, the potential for the taxon to is considered a serious threat to these habitats have been introduced into Torres Strait by across the island. human vectors should not be discounted. The Resilience to fire of Gnetum gnemon is species is widely cultivated across its range, not documented; however, its mesic habitat is including in New Guinea, and hence it is likely to be sensitive to fire. Fire has been an possible that it was translocated to the islands. ongoing influence on the island’s vegetation, Movements of human populations and trade and fire management is an increasing focus between New Guinea and Torres Strait have of the Land and Sea Ranger program (3D greatly influenced the distribution patterns Environmental 2012a, 2012b). Given the of a number of important tropical crop plants small population size and highly localised and has influenced cultivation practices occurrence, there is a risk of stochastic in the region (Haddon 1935; Barrau 1963; extinction of the species on this island. Harris 1977; Barham 2000; Denham 2008; Denham et al. 2009; McNiven et al. 2006; The upland rainforest habitat of the McNiven 2008). Furthermore, the impact of Mua population is similarly remote and Torres Strait islanders upon their pre-contact inaccessible, and no known direct threats surroundings included the introduction of are evident. Population size is not known; new plant species from other islands (Shnukal however, the spatial extent of the forest 2004; McNiven 2008), a practice that is type is mapped as < 477 ha (Stanton et al. ongoing in contemporary culture. The fact 2009). While numerous weeds occur on the that local people do not have a local name or a island they are limited to the margins of the use for the plant does not mean that it was not island communities Kubin and St Pauls and used in the past. along some major tracks and roads. The undisturbed and remote nature of the upland Conservation Status habitat suggests that there would appear to Across the majority of its range Gnetum be a very low risk of stochastic extinction gnemon is a common tree species in the wild of the population. Its lowland occurrence on that is also widely cultivated throughout Mua occupies approximately 20 ha within Malesia. In Torres Strait, a conservative a linear band of riparian rainforest located estimate of its population size on Badu is 500– three kilometres from the coast. The ecotone 1000 individuals within an area of available to adjacent Corymbia and Melaleuca grassy habitat of < 200 ha. Surveys carried out by woodland vegetation is sharp and controlled Fell et all., Gnetum gnemon in Australia 427 by seasonal burning practices. Weeds such as Regional Authority. Field assistance on Badu Brazilian joyweed (Alternanthera brasiliana was provided by Badu Island Ranger Keijie (L.) Kuntze) and leucaena occur nearby. Weed Bowie as well as by Mua Ranger Jackson incursions, and inappropriate fire management Ware, Warraber Trainee Ranger Young Billy which erode fire sensitive riparian margins, and Mer Trainee Ranger Tauki Passi. On represent potentially threatening processes to Mua Island, Ranger Mentor Simon Conaty the integrity of the lowland Gnetum habitat. supported field work and his assistance is most appreciated. Thanks to David Gooding who Integration of further ecological and provided GIS support to detailed vegetation genetic studies are necessary to better mapping over Torres Strait islands. Botanical understand the conservation and management identifications were provided by the staff of requirements of Gnetum gnemon in Torres the Queensland Herbarium. Comments on Strait. Future assessments on the conservation an earlier version of this paper from Garrick values of Gnetum gnemon in Torres Strait Hitchcock, Rob Neal, Paul Forster and should acknowledge that conservation of Barbara Waterhouse are greatly appreciated. peripheral populations is dependent upon the genetic divergence from other conspecific References populations, and may be beneficial to the 3D eNviroNMeNtal (2011a). Biodiversity Profile for protection of the evolutionary process (Lesica Badu Island. Report to Torres Strait Regional & Allendorf 1995). Authority. 3D Environmental: Brisbane. 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Position of Badu and Mua islands in Torres Strait indicating position between Papua New Guinea and mainland Australia.