PALM S Gardiner et al.: barfodii Vol. 56(2) 2012

LAUREN M. G ARDINER 1, Heterospathe JOHN DRANSFIELD 1, JEFF MARCUS 2 AND barfodii , a WILLIAM J. B AKER 1 1Royal Botanic Gardens, Kew, Richmond, Surrey, New TW9 3AB, United Kingdom from Papua [email protected] 2Floribunda Palms, P.O. Box 635, Mountain View, HI 96771, USA

1. Heterospathe barfodii in type locality, showing habit (Photo: Anders Barfod).

An extraordinary new species of Heterospathe from Papua New Guinea, which is already established in cultivation under an incorrect name, is described here for the first time.

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We first became aware of this puzzling, slender herbarium specimens are known from Bacan palm with a striking silvery-white crownshaft and Ternate islands in Maluku, , and when it featured on the cover of the Field Guide from cultivation in Bogor and Singapore to Palms in Papua New Guinea (Barfod et al. Botanic Gardens. The most recent specimen 2001) identified as “ sp.” Anders was collected by John Dransfield in Bogor Barfod from Aarhus University had found and Botanic Garden from the last cultivated , collected the palm with Roy Banka (PNG Forest which was struck by lightning during a storm Research Institute) and John Dowe (James in 1971. Its re-discovery in cultivation would Cook University) in 2000 in a relatively remote certainly have been significant. area of Milne Bay where it grew along a As part of a revision of Heterospathe for the roadside on the edge of disturbed forest (Fig. Palms of New Guinea project (Baker 2002, 1). Through research for a revision of Trudgen & Baker 2008), we compared Rhopaloblaste (Banka & Baker 2004), we specimens and photographs of the realized that this palm did not fit well within cultivated by Jeff Marcus with an extensive this , for example in its range of material from relevant herbaria and with elongate, rather than short peduncle and concluded that the cultivated palm is the same persistent, rather than caducous prophyll (Figs. as the palm collected by Barfod and colleagues 2 & 3). At that time, we decided that the in Milne Bay, Papua New Guinea. We also species most closely conformed to Heterospathe , found that the species does not match the except for one confounding fact – it had a specimens, type description and illustration of conspicuous crownshaft, a feature unknown in H. glauca (Scheffer 1876). The staminate any other species of that genus (Baker & of H. glauca are more robust and have Dransfield 2006, Dransfield et al. 2008). many more stamens than those of the In 2008 John Dransfield visited the nursery of unidentified palm (Fig. 6), 24 or more in the Jeff Marcus in Hawaii and was shown a palm case of the former and just 6–9 in the latter. cultivated under the name Heterospathe glauca Similarly, H. glauca bears large, globose (Scheff.) H.E. Moore (Figs. 4 & 5). Heterospathe “the size of a rifle ball” (Scheffer 1876) up to glauca is an enigmatic species described in the 17 mm in diameter, whereas the unidentified 19 th century as Ptychandra glauca Scheff. and palms bears similar but much smaller spherical not collected in recent years. Only a few fruits which are 7–9 mm in diameter (Fig. 7).

2. Heterospathe barfodii in type locality, showing maroon infrafoliar (Photo: Anders Barfod).

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3. Roy Banka with inflorescence of Heterospathe barfodii (Photo: John Dowe).

From the specimens and images available to us, is indeed a member of the genus Heterospathe , it remains uncertain whether or not H. glauca despite its unusual morphology, which means has a crownshaft, like that of the unidentified that the concept of the genus has to be palm. broadened to account for the presence of a crownshaft in rare instances. It is clear that the unidentified palm is in fact an undescribed species, which we publish here Quite how the species became established in as Heterospathe barfodii , in honor of the cultivation is unclear. Jeff Marcus obtained collector of the first specimen, Anders Barfod, from former IPS President Donn and his contribution to New Guinea palm Carlsmith in 1990. It has been distributed and exploration. Using DNA widely since this time throughout the USA and methods, we have confirmed that the species North Queensland, , but from whom

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4. Heterospathe barfodii growing at Floribunda Palms and Exotics, Hawaii (Photo: John Dransfield).

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5. Heterospathe barfodii growing at Floribunda Palms and Exotics, Hawaii, showing crownshaft, inflorescence and developing infructescence (Photo: Jeff Marcus).

Carlsmith obtained his material remains a Cruces Biological Station, in southern Costa mystery. Several individuals of the new palm Rica, where they were unnamed and lacked were observed by JD in 2008 in the Robert and provenance (Fig. 8). We speculate that they Catherine Wilson Botanical Garden at the Las could have originated from the same source as

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Donn Carlsmith’s plants – perhaps an early leaflet pair sometimes multifolded and distribution of seed from Papua New Guinea partially joined, tapering to bifid apices, bright via the Palm Society Seed Bank. It appears to green, concolorous, red-brown or whitish be a faster grower in cultivation than other medifixed ramenta to 5 mm long scattered on Heterospathe species. abaxial surface of proximal portion of midrib and secondary veins, transverse veinlets Unfortunately, the name Heterospathe glauca , conspicuous, proximal leaflets 39.5–49 cm which seemed so appropriate for this species long, 1.2–2.1 cm wide, middle leaflets 67–78 with its startling glaucous crownshaft, cannot cm long, 4–5.5 cm wide, distal leaflets 22–23.5 be used, but perhaps this will inspire palm cm long, 1.5–2.4 cm wide. Inflorescence experts to track down the true H. glauca , which 68–99 cm long, infrafoliar, branched to 3 is so poorly known and perhaps even orders, branches spreading, deep maroon at threatened in its native Maluku. anthesis; prophyll 25–34 cm long, 5–7 cm Heterospathe barfodii L.M. Gardiner & W.J. wide, opening apically, persistent, covered with Baker, sp. nov. Type: Papua New Guinea, Milne white farinaceous indumentum; peduncular Bay Province, along the Kabawawa highway, bract similar to prophyll, caducous or March 2000, Barfod et al. 4539 (holotype K!; persistent, exserted from prophyll, insertion isotypes AAU!, LAE, BRI, CANB). Fig. 9. ca. 14 cm from base of the peduncle, enclosed within prophyll; peduncle 32–33 cm long, Medium, solitary palm. Stem erect to (2–)8 m 20–22 mm wide; rachillae 14–36 cm long, tall, 7 –10 cm in diam.; scars 1–2 cm wide; 2.5–3 mm in diam., with ca. 6 triads per 1 cm, internodes 3–10 cm (up to 17 cm in juvenile rachilla bracts minute to 1 mm, triangular; all stem); green to dark brown but upper few inflorescence axes sparsely covered with internodes covered with white farinaceous floccose to lanate brown to pale brown indumentum. 7–9 in crown; sheaths indumentum, diminishing along branches. 55–67 cm long, forming a well-defined Staminate 2.9–3.7 mm long, 2.7–3.3 crownshaft, pale green, covered with dense, mm in diam. in bud, ovoid; 7.5–8.4 mm in thin farinaceous white indumentum with diam. and deep maroon at anthesis; sepals 3, scattered, minute, pale brown, white-edged imbricate, thick, triangular, 1.1–1.5 mm long, scales; 20–40 cm long, concave 1.0–1.7 mm wide; petals 3, 3.0–4.0 mm long, adaxially, bright green, indumentum as for 1.7–2.4 mm wide, valvate, triangular; stamens sheath; rachis 205–240 cm long, 10–12 mm 6–9, 4.2–5 mm long; filaments 3–3.5 mm long, wide at midpoint; leaflets 38–56 per side, borne 0.4–0.5 mm in diam., fused at base for 0.4 mm 4 cm apart (along mid-section of leaf), regularly to form a ring around pistillode, white, arranged, in a single plane, held horizontally inflexed; anthers 1.2–1.9 mm long, 0.6–0.9 with tips drooping slightly, narrowly to mm wide, medifixed; pistillode 1 mm long, broadly linear, leathery, singlefold but apical 0.8 mm in diam. at base, conical, minutely

6. Staminate flowers of Heterospathe barfodii on rachilla at anthesis (Photo: John Dransfield).

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7. The fruits of Heterospathe barfodii (Photo: John Dowe). trifid. Pistillate flower immature, bud ca. 4.9 Distribution: Known only from one wild- mm long, ca. 2.7 mm in diam., globose-ovoid, collected specimen, from the lowlands of deep maroon; gynoecium ca. 2.1 mm long, ca. mainland Milne Bay Province, Papua New 0.9 mm in diam., perianth imbricate. 1 Guinea. cm in diam., spherical, surface striate, red Habitat: Lowland tropical when mature; perianth cupule clasping; vegetation at about sea level. The type locality stigmatic remains subapical; endocarp thin is an open grassy area at the side of a dirt road bony, dark brown; occasionally bilobed with through the forest two developed carpels or one developed and one partially developed carpel. Seed 7 mm in Vernacular name and uses : There are no diam., spherical, pale brown; endosperm known records of local names or uses for this ruminate; embryo basal. palm.

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8. Row of mature Heterospathe barfodii trees at Las Cruces Biological Station, Costa Rica (Photo: John Dransfield). Conservation status: This palm meets the therefore its extent of occurrence is estimated criteria for threat category Critically to be less than 100 km 2. Similarly, the area of Endangered (CR B1ab(i,ii, iii, iv, v), C2(ai, aii), occupancy is estimated to be less than 10 km 2, D; IUCN 2001) as it is only known from a the palm is only known from one location, small population at a single location, and and the population size is suspected to number

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9. Heterospathe barfodii . A. crownshaft, inflorescences and leaf; B. apical and middle portion of leaf; C. detail of leaflet underside showing ramenta; D. inflorescence showing prophyll; E. buds on portion of rachilla; F. open staminate flower; G. staminate bud in section; H. fruit on rachilla; I. fruit in section showing ruminate seed. Scale bar: A = 50 cm; B, C, D = 8 cm; E, H = 1 cm; F = 5 mm; G = 3 mm; I = 7mm. A-I from Marcus 1 . Drawn by Lucy T. Smith. fewer than 50 mature individuals as no other location are vulnerable, and the area is under individuals were seen (A.S. Barfod, personal threat from logging, mining and oil palm communication). The type locality is a plantations. Therefore the population of this disturbed forest edge, along the side of a dirt species is likely to decline unless conservation road – by definition individual plants in this action is taken.

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Additional specimens examined: Cultivated: New Guinea taxa is incomplete, resolves it in United States of America, Hawaii, Floribunda a clade with two other New Guinea taxa, H. Palms and Exotics, June 2009, Marcus 1 (K!). delicatula H.E. Moore and H. elegans subsp. humilis (Becc.) M.S. Trudgen & W.J. Baker. In Notes: The genus Heterospathe Scheff. is defined this revised analysis, remains quite by a combination of characters that occur in distinct from Heterospathe . other genera of tribe , rather than by any unique synapomorphies: “ leaf sheath Acknowledgments splitting to the base, interfoliar inflorescences The authors thank Anders Barfod and John (at least at anthesis), peduncle longer than the Dowe for their valuable contributions to this rachis, peduncular bract longer than the manuscript, and Lucy Smith for her analytical prophyll, prophyll persistent, and peduncular illustration of the new species. bracts persistent” (Norup et al. 2006). Heterospathe barfodii shares most of these LITERATURE CITED characters. Although H. barfodii has a distinct BAKER , W.J. 2002. The Palms of New Guinea crownshaft with the outermost leaf sheaths Project. Flora Malesiana Bulletin 13: 35–37. remaining largely tubular, it appears that the oldest sheath is at least partially open with BAKER , W.J. AND J. D RANSFIELD . 2006. Field Guide fibrous margins, a condition that can be to the Palms of New Guinea. Royal Botanic observed on inner leaf sheaths of other species Gardens, Kew. of Heterospathe . It would appear that the BAKER , W. J., M.V. N ORUP , J.J. C LARKSON , T.L.P. possession of a crownshaft is a matter of degree COUVREUR , J.L. D OWE , C.E. L EWIS , J.C. P INTAUD , in Heterospathe . The fact that the inflorescences V. S AVOLAINEN , T. W ILMOT AND M.W. C HASE . are infrafoliar in H. barfodii is most likely 2011. Phylogenetic relationships among related to the physical constraints imposed by arecoid palms (: Arecoideae). a well-defined crownshaft. Annals of Botany 108: 1417–1432. As noted by Norup et al. (2006), the BANKA , R. AND W.J. B AKER . 2004. A monograph combination of a crownshaft and persistent of the genus Rhopaloblaste (Arecaceae). Kew prophyll, as is found in Heterospathe barfodii , Bulletin 59: 47–60. occurs elsewhere in tribe Areceae only in Dransfieldia , , and some BARFOD , A.S., R. B ANKA AND J.L. D OWE . 2001. species of . Of these, H. barfodii most Field Guide to Palms in New Guinea. AAU closely resembles a robust Dransfieldia , on Reports 40. Department of Systematic account of its crownshaft, infrafoliar Botany, University of Aarhus. inflorescence, elongate peduncle and persistent DRANSFIELD , J., N.W. U HL , C.B. A SMUSSEN -L ANGE , prophyll. However, Dransfieldia is a much more W.J. B AKER , M.M. H ARLEY AND C.E. L EWIS . slender palm, with slim, bullet-shaped 2008. Genera Palmarum: Evolution and staminate buds well-spaced along the rachillae Classification of the Palms. Royal Botanic (unlike the relatively crowded arrangement of Gardens, Kew. ovoid buds along the rachillae in H. barfodii ), more stamens (up to 19) and an inner whorl NORUP , M.V., J. D RANSFIELD , M.W. C HASE , A.S. of erect stamen filaments (the filaments all BARFOD , E.S. F ERNANDO AND W.J. B AKER . 2006. being inflexed in H. barfodii ). Dransfieldia is Homoplasious character combinations and also restricted in geographic distribution to generic delimitation: a case study from the the far-western Papua province in Indonesian Indo-Pacific arecoid palms (Arecaceae). New Guinea. We have generated DNA American Journal of Botany 93: 1065–1080. sequence data for the two low-copy nuclear SCHEFFER , R.H.C.C. 1876. Sur quelques palmiers genes PRK and RPB2 from the cultivated du groupe des Arecinées. Annales du Jardin material of H. barfodii , integrating these into Botanique de Buitenzorg 1: 103–164. the recent arecoid dataset of Baker et al. (2011) and re-analysing the data following their TRUDGEN , M.S. AND W.J. B AKER . 2008. A revision methods. The resulting molecular phylogeny of the Heterospathe elegans (Arecaceae) places the species firmly within the genus complex in New Guinea. Kew Bulletin 63: Heterospathe , and although the sampling of 639–647.

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