Research Article |
Corresponding author: Avelinah Julius ( plagiovel80@yahoo.com ) Academic editor: Yasen Mutafchiev
© 2020 Avelinah Julius, Tadashi Kajita, Timothy M. A. Utteridge.
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:
Julius A, Kajita T, Utteridge TMA (2020) Two new species of Ardisia subgenus Tetrardisia (Primulaceae-Myrsinoideae) from Borneo. PhytoKeys 145: 139-148. https://doi.org/10.3897/phytokeys.145.48573
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Ardisia argentiana and A. nagaensis from subgenus Tetrardisia are herein described and illustrated as new species. They are endemic to Borneo and the Indonesian province of Central Kalimantan and to the Malaysian state of Sarawak, respectively. Ardisia argentiana is unique in its linear-oblong leaves, with a long, acuminate-caudate apex, and finely serrulate margins, while A. nagaensis can be easily recognized by its elliptic-lanceolate leaves.
Conservation, endemic, key, Malesia, Myrsinaceae, South-East Asia, taxonomy
Ardisia Sw. with ca. 450 to over 1000 species (Stevens 2001 onwards;
Ardisia is classified into subgenera on characters of habit, leaf morphology, disposition of flowers at inflorescence branch apices (racemes, umbels, corymbs), inflorescence position and floral morphology (see
The subgenus is defined by the combination of the small, woody shrub habit, leaves lacking bacterial nodules with a finely serrulate-denticulate leaf margin, and the usually tetramerous flowers with few ovules. During study of Ardisia specimens from Borneo deposited at KEP and E, two unidentified specimens with tetramerous flowers were encountered and suspected to be undescribed taxa. After close scrutiny of the relevant literature (e.g.
There are very few species of Ardisia with tetramerous flowers, and with few occurring in Malesia and surrounding regions: only a single taxon is recognized each from Java and Thailand, and three are known from Peninsular Malaysia, with none currently described from Borneo. With the publication of the two new species here, subgenus Tetrardisia now has six species, the previously described species, A. denticulata Blume (
This study was based on examination of herbarium specimens at E, K and KEP and the relevant taxonomic literature (e.g.
Differs from other members of the subgenus Tetrardisia in having linear-oblong leaves, with a long, acuminate-caudate apex, and finely serrulate margins.
INDONESIA. Borneo: Central Kalimantan, Kotawaringan [Kotawaringin] Timur, S. Mentaya, km 92 from Sangai, Plot 8, [1°18’S, 112°32’E], 100 m elevation, 18 May 1993, Argent et al. 93187 (holotype E!; iso: BO).
Shrub ca. 1 m high; stems sparsely scaly when young, soon glabrous, slightly flexuous, winged between the nodes with raised lines running along the internodes between the petiole bases. Indumentum of sessile, circular, peltate scales up to 0.05 mm in diameter, on young parts, leaves and inflorescence. Leaves alternate; petioles 3–5 mm long, winged by the decurrent leaf base, glabrous; lamina chartaceous, linear-oblong, (7–)10–11.5 × 1–2 cm, with dense gland-dots throughout apex acuminate-caudate, acumen 1–2.5 cm long; base cuneate, between the higher order venation, lacking hairs, sparsely scaly on the lower midrib, glabrous above, midrib sunken above, raised beneath; lateral veins ca. 48 pairs, semicraspedodromus, and with 1–2 intersecondary veins within each pair; intercostal veins obscure. Inflorescences terminal on lateral branches, proximally laxly paniculate with subumbellate branches with 2-flowers distally racemose; peduncle and rachis (1.5–)2.5–4 cm long, slightly flexuose, sparsely scaly; bracts linear-lanceolate, (0.4–)1.5–1.7 × 0.4 mm, very sparsely ciliate with few scattered hairs. Flowers ca. 8; pedicels slender, ca. 1.5 cm long, sparsely scaly; calyx-lobes 4, green, gland-dotted, lacking hairs, sparsely scaly outside, ovate, ca. 1 × 0.8 mm, apex acute, margin very sparsely ciliate with few scattered short hairs; corolla-lobes 4, reflexed, twisted apically, white with orange-brown elongated-dots, lanceolate, 6.4–7 × 1.7–2.3 mm, glabrous on both surfaces; stamens 4, spreading upright (in open flower), filament ±sessile, anthers twisted apically, lanceolate, ca. 5–6.2 × 0.8 mm, apex elongated into a hyaline tip, thecae open by longitudinal-slits, with scattered, lineate, orange brown dots behind, glabrous; ovary ovoid, ca. 7 × 6 mm, style and stigma filiform, ca. 6.8 mm long, ovules ca. 6 in 1-series. Fruits n.v.
Endemic to Borneo; known only from Sungai Mentaya, Kotawaringin Timur, Central Kalimantan.
The species epithet commemorates Graham Charles George Argent (1941–2019), a prominent botanist on tropical botany in South-East Asia and a leading expert on Ericaceae, especially the tropical ‘Vireya’ rhododendrons, and collector of the type material.
Data deficient (DD). The only specimen available was collected in 1993 and the species is known only from a single collection location south of Bukit Raya in Central Kalimantan, and thus meets the B1a criterion for Critically Endangered (CR) status. The species was found to the south of the Bukit Baka – Bukit Raya National Park, and, to date, there are no further collections of the species from inside this protected area. Satellite imagery in Google Earth from 2015 shows that the collection locality still has some forest coverage and was not penetrated with roads or logging tracks, and it also appears to not have been converted for agricultural use such as oil palm plantations. However, lack of collections and field observations of the species do not allow inference of decline or fluctuation in population size or EOO and AOO, and we are unable to fulfil the criteria to preliminary assess this species as Critically Endangered.
Ardisia argentiana is a distinct species on account of the combination of branches with wing-like raised lines running between the petiole bases, linear-oblong leaves less than 2 cm wide with finely serrulate margins, the terminal, laxly paniculate inflorescence with a hairy scaly rachis and few, tetramerous flowers. This new species is morphologically similar to Ardisia nagaensis, but differs from that species in leaf morphology, especially the shorter pedicels (A. argentiana 3–5 mm long; A. nagaensis 1.5– 3 cm long), linear-oblong leaves less than 2 cm wide (A. nagaensis elliptic-lanceolate to 5.5 cm wide) and the fewer flowered inflorescence (A. argentiana with 8 flowers vs. A. nagaensis with 24).
The flower is described as ‘white’ in the specimen label which probably refers to the corolla and anthers. The leaf resembles Ardisia mystica. B.C. Stone (
Similar to Ardisia tetrasepala in its simple and compact inflorescence with corymbose flowers but differs mainly vegetatively in having leaves laxly arranged, longer petioles that are covered with glandular hairs, a chartaceous lamina without marginal secondary veins, surface with scattered stellate hairs near the margin beneath and leaf base obtuse or ±cuneate. The corolla lobes are smooth without lepidote scales (vs. lepidote scales present in A. tetrasepala).
MALAYSIA. Borneo: Sarawak, Tatau District, Ulu Merirai, Gua Naga, [02°39’12”N, 113°03’05”E], 11 July 2005, Julia et al. S95726 (holotype KEP!; iso: SAR, SING)
Ardisia nagaensis Julius, T. Kajita & Utteridge, sp. nov. A habit B glandular hairs on stem C erect, simple hairs on petiole D the venation close-up E stellate hairs close-up F enlarged inflorescence G flower bud H mature flower I open calyx J open corolla with stamen K corolla lobe L stamen, front and back view M placenta and ovules. (Drawn by Mohamad Aidil Noordin from Julia et al. S95726).
A small shrub, less than 1 m high; stems glabrescent, slightly flexuous, winged between the nodes with obscure raised lines running along the internodes between the petiole bases. Indumentum of simple, short (stalked), glandular, or stellate (star-liked), pale brown to rusty hairs (visible under microscope). Leaves alternate; petioles 1.5–3 cm long, sparsely to densely hairy towards midrib, simple, short, glandular hairs; lamina chartaceous, with dense gland-dots throughout the leaf, glabrous on both surfaces except for margin with scattered stellate hairs beneath, elliptic-lanceolate, 17–19 × 5–5.5 cm, apex long acuminate and slightly caudate with acumen 1.5–2.0 cm long; base obtuse or ±cuneate, margin obscurely denticulate being entire in appearance, midrib flat or slightly sunken above, raised beneath, glabrous except hairy with glandular hairs beneath, denser near leaf base; lateral veins 13–15 pairs, distinct above, prominent beneath, inter-secondary veins present in between; intercostal veins percurrent, distinct beneath. Inflorescence terminal on lateral branches, simple with flowers arranged in corymbs; peduncle and rachis 4–5 cm long, densely hairy with glandular hairs; bracts ovate to elliptic, 0.5–1.5 mm long, hairy with glandular hairs. Flowers ca. 24; pedicels slender, 1.2–1.8 cm long, up to 2 cm long in fruiting, covered with dense, glandular hairs; calyx-lobes 4, purplish, gland-dotted, lacking hairs, broadly ovate, 1–1.5 × 1–1.5 mm, apex obtuse, margin incised, sparsely ciliate with short, thick hairs; corolla-lobes 4, reflexed, twisted apically, purplish with transparent margin, lanceolate, ca. 8 × 2.5 mm, glabrous on both surfaces; stamens 4, spreading upright (in open flower), filament ±sessile, anthers twisted apically, lanceolate, ca. 6 × 1 mm, apex without a prolonged hyaline tip, thecae opening by longitudinal slits, with lineate, black dots behind, glabrous; ovary globose, ca. 1 × 0.8 mm, style and stigma filiform, ca. 6 mm long, ovules 4–6 in 1-series. Fruits ripe bright red, globose, ca. 6 × 5 mm.
Endemic in Borneo, Sarawak; known only from Gua Naga, Ulu Merirai area, Tatau District.
Ardisia nagaensis is very local and was found at only one locality, Gua Naga, for which it is named.
Data deficient (DD). The only specimen available was collected in 2005 and the species is known only from a single collection location from Ulu Merirai in Central Sarawak, and thus meets the B1a criterion for Critically Endangered (CR) status. The Ulu Merirai region is an area of sandstone with limestone cliffs, and supports several point endemics, including several newly described species of Begonia L. (
Compared to the other members of subgenus Tetrardisia, this new species described here has an affinity with A. tetrasepala, endemic to Peninsular Malaysia [Johor, Gunung Pulai]. This is because both are characterized by a simple and compact inflorescence, whereas other taxa in xTetrardisia have compound and laxly flowers arranged. Ardisia nagaensis differs from A. tetrasepala by the leaf characters viz. the petiole length (A. nagaensis with 1.5–3 cm long vs. A. tetrasepala with 0.5–0.8 cm long), the leaf base (A. nagaensis with obtuse or ±cuneate leaf base vs. A. tetrasepala with cordate-rounded) and the venation (A. nagaensis without marginal secondary veins vs. A. tetrasepala with marginal secondary veins). Further morphological comparison between these two species is given in the Table
The calyx is described as ‘purplish’ in the specimen label which probably refers to both the calyx and corolla. The corolla margin on one side is transparent as observed in A. denticulata.
Ardisia nagaensis | A. tetrasepala | |
Leaf texture | Chartaceous | Subcoriaceous |
Leaf shape and size | Elliptic-lanceolate, 17–19 × 5–5.5 cm | Oblong-elliptic, 10–19.5 × 2.5–5 cm |
Leaf base | Obtuse or ±cuneate | Cordate-rounded |
Leaf margin | Obscurely denticulate being entire in appearance | Seemingly entire immature but finely crenulate when mature |
Leaf apex | Long acuminate and slightly caudate with acumen 1.5–2.0 cm long | Acuminate-caudate, acumen 1.5–2.5 cm long |
Leaf surface | Glabrous on both surfaces except for margin with scattered stellate hairs beneath | Glabrous above except sparsely hairy to glabrescent on midrib Beneath |
Lateral veins | Lateral veins 13–15 pairs, distinct above, prominent beneath, inter-secondary veins present in between | Lateral veins 20–24, distinct and slightly prominent on both surfaces, brochidodromus with secondary veins looping and joining 0.2–0.5 mm in from the margin with distinct secondary loops |
Intercostal veins | Percurrent, distinct beneath | Reticulate, distinct and slightly prominent on both surfaces |
Flowers | ca. 24 | 8–13 |
Pedicel | 12–18 mm long | 5–10 mm long |
Calyx lobes | Purplish, lacking hairs, broadly ovate, 1–1.5 × 1–1.5 mm | Green, glabrous inside, sparsely hairy outside, ovate, ca. 1 × 1 mm |
1 | Leaves entire or obscurely undulate or obscurely crenulate | 2 |
– | Leaves distinctly crenulate-denticulate | 5 |
2 | Inflorescences paniculate, 15–20 cm long. Thailand and Malay Peninsula | A. porosa |
– | Inflorescences subumbullate or racemose, 3–5 cm long | 3 |
3 | Leaf lamina linear-oblong, 1–2 cm wide, margin distinctly finely serrulate in the distal half. Inflorescence branched and flowers loosely arranged in racemes. Borneo | A. argentiana |
– | Leaf lamina elliptic, elliptic-lanceolate or elliptic-oblong, 3.5–7 cm wide | 4 |
4 | Leaves with scattered stellate hairs near the margin abaxially, margin obscurely serrulate being entire in appearance; petioles 1.5–3 cm long. Inflorescence with ca. 24 flowers. Borneo | A. nagaensis |
– | Leaves puberulent along the midrib abaxially, margins entire proximally becoming faintly crenulate distally; petioles 5–8 mm long. Inflorescences with 8–20 flowers. Malay Peninsula | A. tetrasepala |
5 | Leaves somewhat bullate, lateral nerves 12–20 pairs; pedicels puberulous. Malay Peninsula and Java | A. denticulata |
– | Leaves not bullate, lateral nerves numerous; pedicels glabrous. Thailand | A. tetramera |
This work is part of the first author’s PhD funded by the Ministry of Education, Culture, Sports, Science and Technology (MEXT) and the Japan Society for the Promotion of Science (JSPS) KAKENHI 16J12040. In addition, the study was also carried out as part of the Flora of Malaysia Project at the Forest Research Institute Malaysia, funded by The Ministry of Science, Technology and Innovation of Malaysia (Project No. 01-04-01-0000 Khas2), and the Ministry of Water, Land and Natural Resources (formerly Ministry of Natural Resources and Environment) of Malaysia under the 10th and 11th Malaysian Plans (SPPII No. P23085100010021 and P23085100018003, respectively). Useful comments from reviewers on how to improve this manuscript are greatly appreciated. The authors also would like to thank Hazel Wilks (K) and Mohamad Aidil Noordin (KEP) for preparing the botanical illustrations.