A new Haniffia species (Zingiberaceae) and a new generic record from Sarawak, Malaysian Borneo

Plant material

Fresh leaf material of Haniffia santubongensis was collected from the type locality, Mount Santubong. The type specimen with the spirit material was deposited to SAR.

Phylogenetic analyses

Phylogenetic analyses were performed with PAUP*4.0b10 (Swofford [2002]) for maximum parsimony (MP) reconstruction with all characters equally weighted. The most parsimonious trees were obtained with heuristic searches of 1,000 replicates with random stepwise sequences addition, tree bisection-reconnection (TBR) branch swapping, collapse of zero-length branches, with the multiple-tree option in effect, and saving up to 10,000 trees from each random sequence addition.

The most suitable nucleotide substitution model for each of the gene regions was selected in jModeltest ver. 0.1.1 (Posada [2008]) using Akaike information criterion (AIC). General time reversible (GTR + I + G) was the nucleotide substitution model selected. Maximum likelihood (ML) analyses were carried out using RAxML 7.2.6 (Stamatakis et al. [2008]). Maximum likelihood bootstrap values were obtained by running 10,000 replicates. Bayesian phylogenetic analyses were performed with MrBayes ver. 3.1.2 (Huelsenbeck and Ronquist [2001]). Markov chain Monte Carlo (MCMC) was repeated twice to assure parameter convergence. The MCMC algorithm was run for 2,000,000 generations with one cold and three heated chains, starting from random trees and sampling one out of every 100 generations. Convergence was assessed by using the standard deviation of split frequencies as convergence index with values < 0.005 interpreted as indicating good convergence. The first 10% of trees were discarded as burn-in. Remaining trees were used to construct 50% majority-rule consensus trees.

Morphology and biogeography

In overall appearance H. santubongensis is most similar to H. cyanescens, sharing with that species a wide bluish labellum and semi-glossy fruits. However, H. santubongensis is clearly distinct from H. cyanescens by the lateral staminodes with an oblique bifid tip, the labellum distally notched (not deeply divided), and differences in labellum colouration and patterning (most notably the presence of a median yellow callus in H. santubongensis).

Haniffia santubongensis represents a new generic record for Borneo where it is locally endemic to Gunung Santubong. Combined with the three locally endemic species in Peninsular Malaysia and P. Thailand, it provides further compelling evidence that these Haniffia species, along with numerous other examples in families as diverse as the aroids, the palms, Rubiaceae Juss., and the genus Hanguana Blume, represent relictual fragments of the Riau Pocket phytochore (Ashton [2005]; Corner [1960]).

Molecular analyses

The combined ITS-plastid dataset contained the new species and 19 species in the Kaempferia Clade recognized in Leong-Škorničková et al. ([2011]) together with two outgroup species. The dataset comprises 2,115 characters: 1,682 characters were constant, 251 variable, but parsimony uninformative, and 182 (8.6%) were parsimony informative. The data analyses produced 32 shortest trees of length 633 steps with a Consistency Index = 0.7615 and Retention Index = 0.7729. The tree topologies obtained from three analyses are similar and consistent with the results from Leong-Škorničková et al. ([2011]). Figure 1 shows the partial tree obtained from maximum likelihood analysis. The new species is shown to be a sister taxon to H. albiflora and H. flavescens. Haniffia cyanescens is supported basally to the three Haniffia species. The new species of Haniffia differs by nine nucleotide substitutions (two in trn K intron and seven in ITS), an addition of 17 bps (trn K intron), an eight bp substitution (ITS), and seven bp deletion (ITS).

Taxonomic treatment

Key to Haniffia species

1a. Corolla lobes and staminodes pale yellow; labellum pale yellow with golden yellow median band, apex emarginate. . . . . . . . H. flavescens

1b. Corolla lobes and staminodes white; labellum white, white with purple veins or dark blue-violet; apex bilobed. . . . . . . . . 2

2a. Leaf blade 10–14?×?2–3.5 cm; ligule with long white hairs; flowers 2–5 in each inflorescence. S Thailand. . . . . . . . . H. albiflora

2b. Leaf blade 17–21 × 3–4.9 cm; ligule glabrous or sparsely pubescent; flowers 5–7 in each inflorescence. Malaysia. . . . . . . . . 3

3a. Lateral staminodes bifid at apex; labellum distally notched, with a central yellow median band. NW Borneo. . . . . . . . H. santubongensis

3b. Lateral staminodes not bifid; labellum deeply spit, without a central yellow median band. Peninsular Malaysia. . . . . . . . 4

4a. Labellum dark blue-violet with white veins; lateral staminodes obovate to oblanceolate. . . . . . . . H. cyanescens var. cyanescens

4b. Labellum white with violet veins; lateral staminodes lanceolate. . . . . . . . . . H. cyanescens var. penangiana

Haniffia santubongensis S.Y. Wong & P.C. Boyce, sp. nov. — Type: Malaysia, Sarawak, Kuching, Gunung Santubong, Summit Trail, just after F4, 04 44 12.1 N 110 19 30.6 E, 2 Sept. 2005, P.C. Boyce & S.Y. Wong ZI-22 (holotype: SAR; isotype: SAR (spirit), FRIM (spirit)) Figure 2.

Diagnosis

Haniffia santubongensis most closely resembles H. cyanescens var. cyanescens (Negeri Sembilan) but is readily distinguished by the oblique-tipped, bifid lateral staminodes, and labellum distally notched (not deeply split) with a yellow median band extending from the entrance to the basal spur to the innermost extent of the lip distal division.

Description

Terrestrial, clumping herb ca 50 cm tall; rhizomes ca 10 mm diam., shallowly buried in soil; roots fibrous. Leafy pseudostem 5–25 per clump, closely spaced, composed of leafless sheaths and leaf sheaths, base ca 1 cm diam.; leafless sheaths ca 6 per shoot, occupying the lower 1/3-1/2, reddish, glabrous, membranous on apex and margins. Leaves 10– 20 per shoot, sub-sessile, sheaths very short, to ca 1 cm long; ligule obtuse to almost truncate, slightly emarginate, ca 3 mm long, soon turning black, glabrous; petiole reddish; lamina narrowly elliptic to lanceolate, 8.5-15 × 2.5-3 cm, base cuneate, apex acuminate, acumen 6–20 mm long, glabrous, bright green, abaxially very slightly paler with deep green slender primary lateral veins. Inflorescences borne at base of leafy shoot, usually solitary, rarely two together, 5–7 cm long, with ca 5 flowers per inflorescence, peduncle to 3.5 cm long, reddish where exposed, otherwise pale green, basal sheaths 1–2, 1–2 cm, pink. Bracts boat-shaped, 15–25 × 6–10 mm, spirally arranged, membranous, reddish green, glabrous, each subtending one flower. Calyx tubular, 20–30 mm long, apex unequally tridentate, greenish, translucent, hairy at tip. Floral tube 36–70 mm long, greenish white, slightly glandular-hairy inside and out; dorsal corolla lobe elliptic, 20–26 × ca. 6 mm, membranous, greenish white with darker veins, lateral corolla lobes narrowly elliptic, 20–25 × ca 4 mm, membranous, greenish white with darker veins, glabrous. Lateral staminodes spatulate, apex oblique bifid, 15–20 × 4–6 mm, greenish white with darker veins, both surfaces covered with dense, greenish white glandular hairs. Labellum obovate, 21–25 × 10–14 mm, swollen at base where adnate to staminodes, apex distally notched, distal lobes curved downwards, white with blue staining, lobes suffused purple or dark blue-violet, centrally with a yellow median callus, upper surface covered with dense, white glandular hairs, lower surface sub-glabrous. Stamen white, covered with dense, glandular hairs; filament 5–6 mm long; anther 6–10 mm long; anther crest 2–2.5 mm long, apex bidentate, white. Style 46–54 mm long, glabrous; stigma cup-shaped, ca1.8 mm long, ciliate. Ovary 3-locular, ovules many. Epigynous glands two, free, linear, 2–2.4 mm long, yellow. Fruit a capsule, dehiscent, somewhat 3-sided, globose with prominent terminal floral remains, ca. 2 × 2 cm, semi-glossy brownish, heavily stained with reddish brown speckles on a cream base, exterior very sparsely warty, splitting longitudinally into 3 valves, valves fleshy, usually with 3-locules well developed. Seeds ellipsoid-obovoid, 6–7 × 3–4 mm, glossy brown, turning greyish-black, aril thick, white when fresh.

Ecology

Partially shaded, deep sandy peat podzols of ridge kerangas in Dryobalanops-dominated hill forest; ca 200–250 m asl.

Distribution

Haniffia santubongensis is known only from the type locality where it occurs as two separate, dense, populations.

Etymology

The species epithet is derived from the name of the type locality.