Research Article |
Corresponding author: Alexandra N. Muellner-Riehl ( muellner-riehl@uni-leipzig.de ) Academic editor: Marc Appelhans
© 2020 Caroline M. Pannell, Jan Schnitzler, Alexandra N. Muellner-Riehl.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Pannell CM, Schnitzler J, Muellner-Riehl AN (2020) Two new species and a new species record of Aglaia (Meliaceae) from Indonesia. PhytoKeys 155: 33-51. https://doi.org/10.3897/phytokeys.155.53833
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Two new species of Aglaia from Indonesia are described, Aglaia monocaula restricted to West Papua, and Aglaia nyaruensis occurring on Borneo (Kalimantan, Brunei, Sabah and Sarawak). A phylogenetic analysis using nuclear ITS and ETS, and plastid rps15-ycf1 sequence data indicates that the two new species of Aglaia are also genetically distinct. Aglaia monocaula belongs to sectionAmoora, while A. nyaruensis is included in section Aglaia. A dichotomous key, drawings and three-locus DNA barcodes are provided as aids for the identification of the two new species of Aglaia. In addition, the geographic range of Aglaia mackiana (section Amoora) is expanded from a single previously known site in Papua New Guinea to West Papua, Indonesia.
Aglaia, conservation status, Indonesia, Meliaceae, phylogeny, taxonomy
The classification of the family Meliaceae continues to be refined (
From the 1990s, the genus received increasing scientific focus due to its bioactivity potential (
The two new species are described based on field observations and examination of herbarium specimens at BO, FHO, K, L, and A, using morphological characters that distinguish them from all other species in the genus Aglaia. Descriptions were written from herbarium specimens. Measurements were made with a tape-measure and calipers. The structure of the indumentum and its distribution was observed and described under a dissecting microscope at magnifications of more than 20×. Flowers were rehydrated by boiling in tap water. They were placed on a glass slide covered with 1 mm graph paper for scale and dissected, measured and described under a dissecting microscope. Additional information on locality, habitat, ecology, plant form, bark and wood characters and fruits was collected in the field and taken from herbarium labels. Conservation threat assessment followed IUCN Categories and Criteria (
Total genomic DNA was extracted for representative samples of each species of Aglaia described herein (Table
For all amplifications, we used the Phusion High-Fidelity DNA Polymerase (New England BioLabs, Ipswich, MA, United States) according to the manufacturer’s protocol. Annealing temperature for ITS (whole region or in two parts) and ETS was 51.5 °C, and for rps15-ycf1 51 °C. PCR products were cleaned using the NucleoSpin Extract II Kit (Macherey-Nagel, Düren, Germany). Sequencing reactions and analyses were run by LGC Genomics (Berlin, Germany).
All sequences were assembled and edited using Geneious (v7.06,
Newly generated sequences of ITS were combined with the data from
Section Amoora
Aglaia monocaula resembles Aglaia flavida, from which it differs through being a smaller, unbranched tree with reticulation subprominent and no indumentum on the lamina of the lower surface of the leaflets. It is unique in the genus in having a dark blackish-brown, slightly swollen, region at the base of the petiolules.
Aglaia monocaula Pannell A habit B detail of lower surface of the leaflet C apical shoot subtending infructescence in a leaf axil D part of the infructescence with fruits cut transversely to show the three seeds and longitudinally to show the junction between the two peltate cotyledons typical of the genus Aglaia (Drawn by Rosemary Wise, edited by Alexandra Muellner-Riehl).
Indonesia. West Papua: Kecamatan Ayfat, neighborhood of Ayawasi, fr. 12 Feb. 1995, K. Yumte 126 (L)
Tree, 3–10 m high, unbranched, with a terminal tuft of spirally inserted leaves; bole 4 cm in diameter; latex white. Twigs greyish-brown with large orange-brown pustules, densely covered with orange-brown and dark brown compact stellate hairs at the apex, glabrescent on older wood.
Leaves 47–70 cm long, 28–32 cm wide; petiole 11–30 cm long; the petiole, rachis and petiolules with few to numerous hairs like those on the twigs, glabrescent. Leaflets 15, the laterals opposite or subopposite, coriaceous, lamina 7–16 cm long 2–5.5 cm wide, elliptical, slightly up-curved at the margins, cuneate at the slightly asymmetrical base, tapering to an acuminate apex, the acumen obtuse and 10–12 mm long; lateral veins 5–14, ascending and curved upwards near the margin, not anastomosing, lateral veins and reticulation subprominent; midrib prominent below with sparse stellate scales, absent from lower leaflet surface, upper and lower leaflet surfaces minutely rugulose; petiolules 10–15 mm on lateral leaflets, slender, 20–40 mm long on terminal leaflet, all with a dark blackish-brown, slightly swollen, region at the base of the petiolules.
Inflorescences not seen.
Infructescence 11 cm long, 7 cm wide; peduncle 6 cm long, the peduncle and fruit stalks with few to numerous hairs like those on the twigs, glabrescent. Fruits 2.8 cm long, 1.8 cm wide, ovoid, pericarp bright scarlet or pinkish-red, inner pericarp white, dehiscent with three locules each containing 1 seed; seed white where attached to the central axis of the fruit by a large hilum, aril orange.
Known only from the area around Ayawasi village in West Papua.
Primary open forest on limestone ridge to 600 m, with an abundant growth of moss. Fruits eaten by kuskus.
Wood used for house beams.
sapa sai (K.Yumte)
The specific epithet of Aglaia monocaula refers to the unbranched habit of this small tree.
This species is known from only two fruiting specimens collected near Ayawasi village and is therefore assessed to be Data Deficient (provisional). Further collecting and monitoring is necessary to allow more conclusive estimations about the rareness and vulnerability of the species. However, the collections seen were made 24 and 25 years ago, so the likelihood of obtaining further material from this species is not great.
Indonesia. West Papua: top ridge of limestone hills south of Ayawasi village, fr., 1 May 1996, Polak 1221 (FHO)
This new species is represented by two fruiting specimens of monocaul trees that have leaves with a long petiolule on the terminal leaflet.
Section Aglaia
Aglaia nyaruensis resembles A. foveolata, from which it differs in its smooth leaflet lower surface, with the lateral veins and reticulation not prominent. These characters, combined with numerous pits on the leaflet upper and lower surfaces, make this species unique in the genus.
Aglaia nyaruensis Pannell A habit with male inflorescence B detail of upper leaflet surface C detail of lower leaflet showing distribution of indumentum D stellate hair E immature infructescence F male flower buds, densely covered with stellate hairs G male flower H half male flower (Drawn by Rosemary Wise, edited by Alexandra Muellner-Riehl).
Indonesia. Kalimantan: Central, Nyaru Menteng Arboretum, off km 28 road to Sampit, alt. 50m, fl., 28 Jan 1995, K. Sidyasa with Ambriansyah, Arifin & Priyono 1422 (holotype BO; isotypes K, L).
Tree, 10–22 m high, bole 8 m, diameter 20 cm, outer bark smooth, greyish or greyish-brown, shallowly fissured and lenticellate, inner bark pink, brownish-green or brown and fibrous, sapwood pale yellow, heartwood white; latex white or absent. Young twigs densely covered with reddish-brown stellate hairs and scales.
Leaves 25–31(–60) cm long, 14–21(–34) cm wide; petiole 7–9 cm long, the petiole, rhachis and petiolules densely covered with reddish-brown stellate hairs. Leaflets 11–15, the laterals subopposite, lamina 5.5–11.0 cm long 2.5–4.0 cm wide, narrowly oblong or elliptical, pale brownish-green when dry, rounded to cordate at the asymmetrical base, acuminate at apex with the acute acumen narrow and to 15 mm long; lateral veins 10–15, ascending and curved upwards near the margin, anastomosing some distance from the margin and with further reticulation between this and the margin of the leaflet, with shorter lateral veins in between. Midrib prominent below, lateral veins and reticulation not raised and barely visible in dried leaflets, the midrib on the upper surface of leaflet with numerous pale brown stellate hairs and scales, the midrib on the lower surface densely covered with reddish brown stellate hairs and scales, numerous on the lower leaflet surface when young, glabrescent, becoming sparse on the mature lamina near the midrib and absent from the rest of both surfaces of the lamina, with numerous pits on both surfaces; petiolules to 3 mm long on lateral leaflets to 10 mm long on terminal leaflet.
Male inflorescence 20 cm long, 10 cm wide; peduncle 8–9 cm long, the peduncle, rachis and first branches densely covered with reddish-brown stellate hairs and scales; higher orders of inflorescence branches with numerous reddish-brown stellate hairs and scales. Male flower 2 mm long, 2 mm wide; pedicel 1 mm long, the calyx and pedicel densely covered with reddish-brown stellate hairs and scales; calyx cup-shaped, deeply divided into five rounded lobes, petals 5, 1.75 mm long, 1 mm wide, yellow, obovate; staminal tube 1.5 mm long, 1.5 mm wide, obovoid with a wide mouth 1.5 mm across, anthers 6, 0.75 mm long, 0.25 mm wide, inserted half way down the tube inside and protruding through the aperture; ovary 0.5 mm long, 0.5 mm wide, ovoid, densely covered with brown stellate hairs and scales on the outside, with two locules each containing one vestigial ovule. Female flowers not seen.
Infructescence 24 cm long, 26 cm wide, peduncle 8 cm; peduncle rachis and branches densely covered with reddish-brown stellate hairs and scales. Young fruits 2 cm long, 1.5 cm wide, ellipsoid, reddish-orange, densely covered with reddish-brown stellate hairs and scales.
One record each from Kalimantan, Brunei, Sabah and Sarawak.
Peat swamp forest, swampy forest on white sand, on ultrabasic soil or on yellow-brown sandy soil over Tertiary clays, with deep litter and abundant humus and living roots. Altitude to 400 m.
Jalongan sasak (Bejang b. Sitam).
The specific epithet of Aglaia nyaruensis refers to the type locality, Nyaru Menteng in Kalimantan.
This species is known from one locality each in Kalimantan, Brunei, Sabah and Sarawak and is therefore considered to be Vulnerable.
Malaysia. Sarawak: Sibu, Haman Forest Reserve, c. 3 m alt, fr 18 June 1958, Bejang b. Sitam 9169 (K); Sabah: Sandakan, Bt Tawai Forest Reserve, 400 m alt., young flowers 26 June 1996, S. Diwol & L. Madani SAN 135187 (K). Brunei, Belait: Sungai Liang, Andalau Forest Reserve Compartment 5, 4°38'41"N, 114°30'20"E, 30 m alt., sterile, 8 March 2004, A.N. Muellner, C.M. Pannell, G. Challen, Jangurun, Muhd Yussof, Ibrahim ANM2039 (K).
New record for West Papua, Indonesia
Previously known only from the type locality in Papua New Guinea, this tall tree species in sectionAmoora, has the largest fruits recorded for the genus Aglaia. Collections from West Papua are of immature fruits and flower buds.
Aglaia mackiana A leaf with attachment to twig B apex of shoot C detail of upper leaflet surface D detail of lower leaflet surface E immature inflorescence F flower buds G peltate scales H transverse section of immature fruit with three seeds I seed, with large hilum and intact aril (Drawn by Rosemary Wise, edited by Alexandra Muellner-Riehl).
Indonesia, two records from West Papua. In Papua New Guinea, known only from the type locality in Chimbu Province.
Primary lowland forest on the coastal plain and to 450 m altitude; canopy 25–45 m high; associate species include Celtis, Sterculia, Pometia, Ficus, Oncospermum, and sundry Rubiaceae. Canopy tree to 45 m tall, branching above; bole c. 1 m diameter, buttressed below; bark tan, smooth, somewhat round flaky; fruits 12–16 cm diameter, light brown, lactiferous, 3-lobed. In Papua New Guinea, the fruit either dehisces on the tree and the seeds fall to the ground or the whole fruit falls from the tree and dehisces on impact with the ground. The seeds are swallowed whole by the Dwarf Cassowary and defaecated at up to 1000 m from the parent trees (
‘sapa peka’ (Wanda Ave 4394)
Named after Andrew Mack, who discovered this species in the course of his field work on the Dwarf Cassowary.
This species is known from only three localities, two in Papua and one in Papua New Guinea. It is therefore assessed to be Data Deficient (provisional). Further collecting and monitoring is necessary to allow more conclusive estimations about the rareness and vulnerability of the species. However, the collections seen were made 24, 25, 27, and 28 years ago, so the likelihood of obtaining further material from this species is not great.
Indonesia. West Papua: surroundings of Ayawasi, 1°09'S, 132°12'E, c. 450m, fruit, 30 April 1996, Wanda Ave 4394 (L); Sarmi, coastal plain, 1–3 km N of Sewan on the Waske River. 2°4'S, 138°46'E, 10–20 m, fr., 3 June 1993 McDonald and Ismail 3786 (BO, L, K). Papua New Guinea. Chimbu Province: Crater Mountain Biological Research Station, 145043–45'S, 6'05–58'E, leaves only, 1992, Mack 699 (FHO! holotype); same locality, ?1995, fruit only, Ross Sinclair RS 105 (FHO!); same locality, seeds only, 18 Aug. 1995, Mack s.n. (FHO!); same locality, fallen male inflorescences only, no date, Mack 297 (A):
To accommodate the three species the following couplets (in bold, labelled (i) and (ii), can be inserted into the existing key to Malesian Aglaia by
1a | Leaf always a single blade | 2 |
1b | Leaves trifoliolate or imparipinnate | 7 |
7a (1) | Leaflets with few or no hairs or scales on the lower surface, the reticulation continuous and subprominent on one or both surfaces | 8 |
7b | Leaflets with at least some scales or hairs on the lower surface, although these may be few and difficult to see, reticulation not continuous and subprominent on either surface, or if subprominent, then with indumentum on lower surface of leaflet | 10 |
8a (7) | Leaflets with reticulation subprominent on the lower surface | 8a (i) & (ii) |
8a(i) | Tree branched, leaflets 2–7, fruits indehiscent | 66. A. cumingiana |
8a(ii) | Tree unbranched, leaflets 15, fruits dehiscent | A. monocaula |
8b | Leaflets with reticulation subprominent on both surfaces | 9 |
10a (7) | Leaflets linear-lanceolate or narrowly elliptical, most being at least 5 times longer than wide | 11 |
10b | Leaflets ovate, elliptical, oblong, obovate, lanceolate or oblanceolate, most less than 5 times longer than wide | 14 |
14a (10) | Indumentum dense, of white or pale brown hairs or scales which totally conceal the lower surface of leaflet | 15 |
14b | Indumentum reddish-brown or, if pale, not totally concealing the lower surface of leaflet | 20 |
20a (14) | Lower surface of leaflet so densely covered with reddish-brown or orange brown hairs or scales, that the surface is not or barely visible between them | 21 |
20b | Hairs or scales absent from the lower surface or, when present, the lower surface of leaflet readily visible between them | 24 |
24a (20) | Indumentum of peltate scales, sometimes with stellate scales interspersed | 25 |
24b | Indumentum of stellate hairs or scales; peltate scales absent | 69 |
25a (24) | Indumentum of peltate scales only | 26 |
25b | Indumentum of peltate and stellate scales (or with at least some of the scales with a long fimbriate margin) | 60 |
26a (25) | Scales densely covering lower surface of leaflet | 27 |
26b | Scales ± absent to numerous on lower surface of leaflet | 31 |
31a (26) | Scales few to numerous on lower surface of leaflet | 32 |
31b | Scales ± absent from lower surface of leaflet but may densely cover the midrib below and immediately adjacent to it and occasionally on the lateral veins | 46 |
46a (31) | Scales densely covering the midrib on lower surface of leaflet and immediately adjacent to the midrib, occasionally also on the lateral veins | 47 |
46b | Scales ± absent from lower surface of leaflet | 54 |
47a (46) | Scales large (many 0.2 mm across), orange-brown, reddish-brown or almost white, with a tendency to flake off | 48 |
47b | Scales less than 0.2 mm across or if larger, then dark reddish-brown or purplish-brown and adhering closely to the leaflet | 50 |
50a (47) | Anthers and/or staminal tube with simple white hairs | 51 |
50b | Anthers and staminal tube without hairs | 52 |
52a (50) | Leaflets with purplish-brown fimbriate peltate scales densely covering the midrib below and ± absent from the rest of the lower surface of the leaflet | 34. A. glabrata |
52b (50) | Leaflets with dark reddish-brown peltate scales numerous on the midrib below | 53 |
53a (52) | Leaflets 7–23, stellate scales absent | 53a (i) & (ii) |
53a(i) | Leaflets 11–23, 50–88 cm long, 36–54 cm wide, fruits dehiscent, at least 12.5 cm long and 10 cm wide | A. mackiana |
53a (ii) | Leaflets (7-)9–13(-15), 5–18.5 cm long, 1.5–4.5 cm wide, fruits indehiscent, 1.5–3 cm long, 2–3.5 cm wide | 33. A. scortechinii |
53b | Leaflets 3–5(-7), some stellate scales interspersed among the peltate scales | 50. A. odoratissima |
69a (24) | Leaflets with few to densely covered with stellate hairs or scales on the lower surface; when sparse, some hairs or scales occur evenly distributed between the veins and their presence visible with the naked eye | 70 |
69b | Leaflets without or with few hairs on the lower surface, with scales visible only with a lens or densely covered with hairs on the midrib only, few and unevenly scattered on the rest of the lower surface | 88 |
88a (69) | Lower surface of leaflet with numerous stellate or peltate scales | 89 |
88b | Leaflets with hairs or scales few on the lower surface between the veins when mature, but sometimes densely covering the midrib | 93 |
93a (88) | Stellate hairs or scales more than 0.15 mm in diam., numerous on or densely covering the midrib, sometimes also on the lateral veins, almost absent elsewhere | 94 |
93b | Stellate hairs or scales either very small, less than 0.15 mm in diameter, or almost totally absent from the midrib below and from the rest of the lower surface of leaflet | 109 |
94a (93) | Leaves ± sessile or with a short peduncle of not more than 1 cm; the basal part of leaflets much smaller than the rest and subrotund | 61. A. subsessilis |
94b | Leaves not sessile, the basal leaflets only slightly smaller than the rest and of similar shape | 95 |
95a (94) | Reticulation subprominent on lower surface and often on upper surface of leaflet | 96 |
95b | Reticulation may be visible, but not subprominent | 97 |
97a (96) | Petals 3, densely covered with stellate scales on the outside; fruits dehiscent | 9. A. lepidopetala |
97b | Petals 5, without scales on the outside, fruits indehiscent | 98 |
98a (97) | Tree unbranched; leaflets shiny | 69. A. coriacea |
98b | Tree branched; leaflets not shiny | 99 |
99a (98) | Fruit c. 0.5 cm in diameter, with few stellate scales | 64. A. aherniana |
99b | Ripe fruit 1 cm or more in diameter, with dense indumentum | 100 |
100a (99) | Leaflet apex with a parallel-sided acumen | 101 |
100b | Leaflet apex with a tapering acumen | 102 |
101a (100) | Leaflets coriaceous | 42. A. forbesii |
101b | Leaflets not coriaceous | 101b (i) & (ii) |
101b(i) | lateral veins subprominent | 40. A. leptantha |
101b(ii) | lateral veins not raised | 43a A. nyaruensis |
The final lengths of our alignments were 1006 bp (ITS), 515 bp (ETS), and 600 bp (rps15-ycf1). The results of our phylogenetic analyses of the combined nuclear data were largely congruent with the infrageneric relationships of
Majority-rule consensus tree of the combined nuclear dataset (ITS and ETS). Node values indicate Bayesian posterior probabilities. Clade numbers on the right refer to the clades identified by
DNA barcodes
Three-locus DNA barcodes (Table
Voucher information and GenBank accession numbers for Aglaia monocaula and A. nyaruensis.
Taxon | Locality | Voucher | ITS | ETS | rps15-ycf1 |
---|---|---|---|---|---|
A. monocaula | West Papua | Polak 1221 (FHO) | MT439806 | MT439713 | MT409504 |
A. nyaruensis | Kalimantan | Sidiyasa et al. 1422 (L) | MT439808 | MT439716 | MT409506 |
Kalimantan | G. Laman et al. 1397 (A) | MT439807 | MT439715 | MT409505 | |
Brunei | Muellner et al. 2039 (K,BRUN) | KF212126 | MT439714 | – |
We would like to thank the directors and curators of the following herbaria for permission to remove samples from herbarium specimens for DNA extraction and sequencing: Naturalis, Leiden (L), Royal Botanic Gardens, Kew (K), Harvard University Herbaria (A), Smithsonian Institution, Washington (US), Daubeny Herbarium, University of Oxford (FHO), living collections and Herbarium of the University of Vienna (WU) and L for a loan received and processed by the National Botanic Gardens of Ireland (DBN); Claudia Krüger for assistance in DNA laboratory work; Rosemary Wise for the detailed botanical illustration of the species; and David Kenfack as well as an additional anonymous reviewer and the editors for useful comments on the manuscript. Financial support for this study was provided by the project “Indonesian Plant Biodiversity and Human Health – BIOHEALTH” (BMBF grant no. 16GW0120K) to Alexandra N. Muellner-Riehl. Jan Schnitzler is currently supported by the German Centre for Integrative Biodiversity Research (iDiv) Halle‐Jena‐Leipzig funded by the German Research Foundation (DFG – FZT 118).
Table S1. Voucher information, origin, and GenBank accession numbers for all species included in this study
Data type: molecular data
Explanation note: New GenBank accessions (this study) are highlighted in bold.