Three new species and a new name in Southeast Asian

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RESEARCH ARTICLE InTRoduCTIon
Gomphandra Wall.ex Lindl. is a genus of approximately 65 species of small trees and shrubs distributed from Sri Lanka and India to China, Indo-China, and throughout Southeast Asia to the Solomon Islands.The genus was revised by Sleumer in 1969 prior to its inclusion in Flora Malesiana (Sleumer 1971).Although it was traditionally placed in Icacinaceae, Kårehed (2001) provided evidence that the family is polyphyletic and placed Gomphandra in the segregate family Stemonuraceae.Gomphandra is the largest genus in the family, with 33 species previously recognized (Sleumer 1969) and approximately 30 new taxa.The genus is characterized by functionally dioecious flowers, a large sessile stigma, and fruit with longitudinal inner mesocarp ridges.Other genera in the family that share these characteristics also have laterally compressed fruit, usually with a fleshy lateral appendage, both of which are absent in Gomphandra.The site of appendage attachment may correspond to a distinctive pair of ridges and grooves flanking a central ridge on one side of the inner mesocarp in Gomphandra, a feature we refer to as a sulcus because the immediate impression is of a groove on the inner mesocarp (Fig. 1).The term 'pseudohilum' was considered for this feature, but it has already been used to refer to a feature of the seeds of Sapindaceae (Weckerle & Rutishauser 2005).The sulcus is not visible in fresh fruit but can be pronounced on dried mature fruits, though the ridges vary in prominence depending upon the species.Sleumer (1971) and others referred to the endocarp being ridged, but the ridges represent sclerified bundles of the inner mesocarp.
Many species of Gomphandra have a poor collection record, and a few are known only from the type collection.Ideally, a new species publication would include descriptions of pistillate flowers, staminate flowers, and mature fruit.However, for the poorly known species of Gomphandra, complete reproductive material is usually lacking.Nonetheless, new species can be described if their morphologies are distinctive enough to preclude confusion with previously described taxa.Two of the species described below were recognized as new by Sleumer (1969), but he did not choose to describe them at the time.Very few additional specimens have been collected since, and it is unlikely that more material will be collected in the future, as all three species described below are probably either extinct or critically endangered by habitat loss.Schori,; Map 1 Arbor ad 25 m alta.Folia coriacea, ovata-ellipsoidea ad obovata vel fere orbiculata, 5.6 -13.0 cm longa, 2.6 -6.7 cm lata, nervis secundariis 4 -5(-6) utrinque, desuper impressis, superioribus ante marginem confluentibus.Inflorescentiae ignotae.Infructescentiae cymosae, ramis primariis usque quatuor, axillares aut terminales.Fructus 1.9 -2.1 cm longi, 0.9 cm lati, ellipsoidei sed latissimi in partem superiorem; putamen protrusum ultra b a excentricum stigma, valde porcatum.-Typus: Haji Bujang S.30259 (holo A; iso K, L), Malaysia, Sarawak, 3rd Division, Batang Igan, Sungai Tutus Logging Camp, 6 Sept. 1970.Small tree to 25 m high and 40 cm diam; trunk fluted (to 2 m) or not.Twigs pubescent with minute, somewhat appressed hairs when young, eventually becoming glabrescent, 1.5 -3.0 mm diam, terminal buds falcate, densely covered with light brown pubescence.Leaves coriaceous, ovate-elliptic to obovate or almost orbicular, frequently conduplicate, apex acute to shortly and abruptly acuminate, base acute, lamina shortly decurrent onto the petiole, margin flat to slightly revolute, glabrous above when mature, sparsely pubescent along the veins below, shining above when dried, matte below, 5.6 -13.0 by 2.6 -6.7 cm; midrib sunken above, raised below, secondary veins 4-5(-6) pairs, impressed above, weakly brochidodromous with the upper pairs joining 1-2 mm from the margin, concolorous above, darker than the leaf surface below and raised, tertiary veins generally not visible above, ± raised below, percurrent and perpendicular to the midrib; petioles grooved, initially tomentellous but glabrescent with age, 1.0 -2.5 cm by 1.0 -1.6 mm.Inflorescences and flowers not known.Infructescences with up to 5 fruits developing (out of perhaps 12 or more flowers), either terminal on sympodial branches, or axillary, when axillary, most often on the older, naked portion of the twigs, cymose with 2 -3 degrees of branching, up to 4 branches borne on a common peduncle to 1.5(-2.0)cm long, branches to 1.2 cm long, fruit sessile or on short pedicels when more than one fruit develops per branch, all axes and calyx with minutely appressed pubescence.Fruit obovate-ellipsoid, widest in upper half, apex and base rounded, 1.9-2.1 by 0.9 cm, glabrous; subtended by persistent calyx 3 mm diam; inner mesocarp with c. 7 strong longitudinal ridges, stigma distinctly eccentric in dried fruit, 3 mm diam, facing away from sulcus.
Habitat & Ecology -Only collected from (peat) swamp forests at low elevations up to 3 m.
Field notes -Bark smooth to narrowly longitudinally fissured with a somewhat scabrous surface, grey or pale yellowish grey, inner bark orange.Fruit pinkish white, orangey-pink, or yellowish grey.
Notes -Gomphandra palustris is distinguished by its rounded, coriaceous leaves with impressed brochidodromous veins, and its strongly ridged, asymmetrical fruit.It is not similar to any other species in Borneo and is probably most closely related to the Philippine G. luzoniensis (Merr.)Merr.group, whose members have a similar infructescence structure, fruit size and colour.All collections, to date, are fruiting specimens collected from June through October.On specimens with immature fruit, the young ovaries are strongly cylindrical, with no evident asymmetry.

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This species was mentioned by Anderson (1963) as Gomphandra aff.comosa King, and he cited three additional specimens, which are presumably at SAR: Anderson 3116 and 14502 (2nd Division, Triso P. F.), and Anderson 4189 (4th Division, Sg.Dua); unfortunately, we have been unable to see and verify these collections.Using specimens we have seen to map the distribution, the species is known from peat swamp forests along the northern coast of Sarawak.Following IUCN (2001) guidelines to estimate threat, these collections have a small extent of occurrence (11 779 km 2 ) suggesting a conservation rating of Vulnerable (VU).However, using satellite imagery, most of the collections (apart from Burley & Lee 353) are now areas of plantation (probably oil palm), and, even with the additional collections we have not seen, a conservation of Endangered (EN B1ab(i,iii,iv)) is more appropriate for this species.
Distribution -Endemic to the Philippines, restricted to Mt Halcon on Mindoro.
Habitat & Ecology -On a ridge at 1 200 m, in a forest dominated by gymnosperms and Tristania, on gravel/sand, possibly from a quartz base, and from cloud forest at 1 350 m.Leaf galls are present on the A and K specimens.According to the Mangyan man who collected the specimen for Ridsdale et al., "bees" pollinated the flowers and birds ate the fruit.Fruit doves or hornbills might be able to swallow the fruit whole, but other birds would be limited to consuming the thin mesocarp.
Field notes -Outer bark smooth, brown and greenish, sometimes with short vertical lines of lenticels, inner layer of outer bark green, inner bark cream.Leaves dark green.Fruit yellow-cream to yellowish orange.
Notes -This species is most similar to G. oligantha Sleumer, but it differs in its coriaceous leaves with a revolute margin, obscure venation, longer petioles, and shorter acumen, and in its thick, shorter peduncle and globose fruit.ICBN Rec.7A we have not designated either of these duplicates as holotype.
Gomphandra halconensis is known from two fruiting collections from Mt Halcon.Mt Halcon is currently closed to all visitors and is now under ancestral lands jurisdiction.According to the Mangyan man who collected Ridsdale et al.'s specimens, only one tree was present in Ramayan village, and the tree has since been cut.Apparently much of the forest on Mt Halcon has been cut for charcoal production and slash-and-burn agriculture (kaingin), but perhaps this species can still be found on the mountain's slopes.Following IUCN (2001) guidelines, the limited number of localities and the small area of occupancy (extent of occurrence cannot be calculated for two points), together with known problems of habitat destruction on Mt Halcon, give a conservation rating of Critically Endangered (CR2ab(iii)).
Habitat & Ecology -On ridge at 1 705 m, on clay loam in secondary disturbed forest and slightly disturbed gully forest.
Field notes -Leaves glossy above and dull and paler below, midrib yellowish.Fruit yellowish or slightly pink (both specimens with submature fruit).
Etymology -The specific epithet honours Harold C. Conklin, an anthropologist at Yale University's Peabody Museum, who first collected this species as part of his research for what became the Ethnographic Atlas of Ifugao.
Notes -Gomphandra conklinii is distinguished by its relatively small, coriaceous leaves with irregular venation and by its fruits, which are the largest of any Gomphandra in the Philippines.PPI collectors (12920) did not see ripe fruit but reported that the fruit are purple when ripe.This anecdote is somewhat doubtful, given the fruit colours that are reported for other collections of this species, and considering that all observations of ripe fruit in other species of Gomphandra list the colour as white, ivory, yellow, pink, salmon, or red.Based on herbarium records, this species blooms in September and fruits from March to July.
Gomphandra conklinii was first collected in 1961 and has been found as recently as 1994.While Ifugao Province has retained its forested areas and in fact has more forest now than it did when Conklin collected this species, much of the forest is highly degraded.The 1990s collection sites have been extensively cut or converted to agriculture and then abandoned to grassland.The local forester in Banaue did not recognize photos of herbarium specimens of this species.There are also reports that previously common species have become very scarce or even extirpated in the past generation, presumably as a result of overharvesting.Applying IUCN (2001) standards, especially the geographic range which is extremely small (e.g., area of occupancy is only 4 km 2 ), and taking into account forest degradation, this species is Critically Endangered (CRB2ab(iii)).
Conklin's collector, Ballogan (Banlugan is a spelling variation), reported the Ifugao common name 'halipan' for this species and said that the branches were used as a supplement to betel nuts.However, in 2007, when the first author visited Banaue, 'hali-pon' referred exclusively to Ficus spp., and no one could recall using twigs of any plant along with betel nuts.Mr. Ballogan said that while collecting for Conklin, he would ask locally about common names, and a common name was reported on the herbarium label if three people gave the same identification.On his own collection, Conklin reported 'lungi' as a common name and said that the timber was used for house construction.This use is much more in line with other reports of economic uses of Gomphandra, but we have not had an opportunity to search for this species under this other common name.
Habitat & Ecology -Primary and secondary rain forest, lower montane Castanopsis forest zone, lowland rain forest dominated by Anisoptera, tall alluvial forest and flood plain forest, on steep to moderate slopes, along streams, in riverine swamp forest, at forest edges, in a limestone area and on ultrabasic alluvial soil, and in disturbed forest on flat, well-drained land, 5-500 m elevation.
Field notes -Bark grey, grey-green, greenish brown, smooth or with pustular lenticels.Leaves dark to medium green and matte or shiny above, paler or yellow green and matte below.Calyx (staminate flowers) green, petals light green.Fruit reportedly yellow or orange when ripe (no collections of ripe fruit).
Notes -Gomphandra rarinervis is one of the most easily recognized Gomphandra species in New Guinea, with large leaves, widely spaced veins, a long, prominently ribbed drupe, and spreading pubescence on both the twigs and leaves.
Specimens of this species have most often been identified as G. papuana (Becc.)Sleumer, or less frequently as G. montana (Schellenb.)Sleumer.Gomphandra rarinervis has been collected from the northern part of Papua New Guinea, and it does not appear to overlap in distribution with G. papuana, which occurs in Irian Jaya, Western, Gulf, Chimbu, Central, and Northern Provinces, with a disjunct population in West Sepik Province near Vanimo.Gomphandra papuana has similar leaves, but the pubescence on the leaves and twigs is very tightly appressed, and while the fruits are a similar size, the inner mesocarp seldom develops, so no ribs are evident, the stigma is much smoother, and the epicarp has a granular texture.Gomphandra montana was described from East Sepik Province, and unfortunately all the fruiting syntypes appear to have been destroyed at B, but the staminate specimens have much smaller leaves with close parallel veins, and a long-peduncled inflorescence that is multiply branched.Material from Central Province, originally described as G. carrii Sleumer, probably belongs to G. montana.The fruits are up to 3 cm long (matching Schellenberg's description), but they lack the beak of G. rarinervis, have more numerous and less prominent ridges, and are borne on a long-peduncled, multiply branched infructescence.
Gomphandra rarinervis matches type material of Stemonurus puberulus, but the epithet 'puberula' was used in Gomphandra by Ridley (1915), so a replacement name is needed.The epithet 'rarinervis' has been chosen in reference to the widely spaced veins on the leaves.The syntypes of Stemonurus puberulus were at B, and Sleumer inappropriately designated Lauterbach 2204 as a lectotype in 1969, after the specimen had been destroyed.The other specimen cited in the protologue of S. puberulus, Lauterbach 2483, evidently did not have any duplicates.The curator at WRSL did not respond to inquiries about their duplicate of the Lauterbach 2204 specimen, which Sleumer also cited, but a fragment at Leiden (ex WRSL), confirmed the identity of the basionym and must be chosen as the lectotype.
To date, G. rarinervis has been collected from 22 localities in Morobe and Madang Provinces and is given a conservation rating of Least Concern (LC) using IUCN (2001) guidelines.However, the extent of occurrence (39 241 km 2 ) is large, and above any of the IUCN threat thresholds, because of a couple of 'outlier' collections (especially Robbins 2033b, but also Kiapranis et al. LAE 87104).If either of these localities no longer held viable habitat for G. rarinervis then a rating of Near Threatened (NT) is more appropriate.

Fig. 1
Fig. 1 Sulcus.a.The stigma is displaced toward a pair of parallel grooves (emphasized here as thick black lines) divided by a ridge, all of which is here referred to as a sulcus.The longitudinal ribs of the inner mesocarp are visible to either side; b. sulcus in profile.The thick black bars correspond to the thick lines shown in the lateral view.The ridges and grooves of the sulcus are not more prominent than those of the longitudinal ridges in this species (G.luzoniensis, Reynoso et al.PPI 17155).-Scale bars = 1 cm.
The herbarium label forRidsdale et al. 1702  indicates that duplicate collections were deposited in the Philippines at PNH and LBC.Despite considerable effort, we have not been able to confirm the presence of the specimen at PNH in either the accessioned or the unaccessioned material, and we have great concerns about the long-term conservation of the specimen in LBC due to housing conditions.Accordingly, to comply with Map 1 Distribution of G. palustris in Sarawak, Malaysia, Borneo.Map 2 Distributions of G. halconensis (-) in Mindoro and G. conklinii (:) in Luzon, Philippines.