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Nepenthes Baramensis (Nepenthaceae) – a New Species from North-Western Borneo

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Nepenthes Baramensis (Nepenthaceae) – a New Species from North-Western Borneo Blumea 56, 2011: 229–233 www.ingentaconnect.com/content/nhn/blumea RESEARCH ARTICLE http://dx.doi.org/10.3767/000651911X607121 Nepenthes baramensis (Nepenthaceae) – a new species from north-western Borneo C. Clarke1, J.A. Moran2, C.C. Lee3 Key words Abstract Nepenthes baramensis, a new species from peat swamp and heath forests in north-western Borneo, is described. It is distinguished from related species on the basis of its modified pitchers, which facilitate a faculta- Borneo tive mutualistic interaction with Hardwicke’s Woolly Bat, Kerivoula hardwickii, which roosts in its pitchers and may ecology provide the plant with a substantial proportion of its foliar nitrogen. Nepenthaceae Nepenthes baramensis Published on 12 October 2011 Nepenthes rafflesiana taxonomy INTRODUCTION ent resource for the plant: on average, 34 % of foliar N may be derived from this source; in contrast, pitchers of the typical form Pitcher plants of the genus Nepenthes (Nepenthaceae) are are not used as roosting sites by bats and faecally-derived N carnivorous, producing jug-shaped leaves that serve primarily is not present in the leaves (Grafe et al. 2011). Not all pitchers as traps for arthropods and other small animals (Clarke 1997). of the elongate form are host to the bats, and it is possible that Raffles’ pitcher plant, N. rafflesiana Jack (Nepenthaceae), the current facultative relationship represents the early stage occurs in Borneo, Peninsular Malaysia and Sumatra (Clarke of a nascent mutualism. 1997, 2001), where it commonly grows in disturbed vegetation Prior to the research outlined above, the taxonomic importance and ecotones over nutrient-deficient substrates up to 1200 of the morphological differences between these two taxa was m altitude. It is very common in northern Borneo and the debatable (Clarke 1997, Phillipps et al. 2008). Given that: i) Malaysian state of Johor in the Malay Peninsula, but patchy similar degrees of variation exist within some other species elsewhere throughout its range. The plant is a robust climber, (e.g. N. mirabilis (Lour.) Druce); ii) this level of intraspecific producing vines that may exceed 15 m in length. The pitchers variation is not unusual in Nepenthes; and iii) there has been a of the rosettes and climbing stems demonstrate pronounced tendency among taxonomists not to recognise subspecific taxa dimorphism which enables this species to target crawling and within the genus (Danser 1928, Jebb & Cheek 1997, Cheek flying arthropods in those parts of the habitat where these prey & Jebb 2001, Clarke 2001), there has been some degree of are most abundant (Moran 1996, Moran et al. 1999, Di Giusto uncertainty about how to treat the elongate form. Furthermore, et al. 2008, 2010). Nepenthes rafflesiana displays considerable there appears to be some disparity among ecologists regarding morphological variation (Cheek & Jebb 2001, Clarke 2001, its identity. Moran (1996), Gaume & Di Giusto (2009) and Grafe Phillipps et al. 2008) and in Borneo, a sub-specific taxon that et al. (2011) referred to it as N. rafflesiana var. elongata Hort., produces, narrow, elongated pitchers has been recorded on a whereas Bauer et al. (2011) used the terms ‘elongate form’ and number of occasions (Clarke 1997, Phillipps et al. 2008). There ‘typical form’ to distinguish between them. The name N. raf- are several consistent morphological differences between this flesiana var. elongata derives from plants that were raised elongate taxon and ‘typical’ N. rafflesiana (Moran 1996, Gaume in cultivation at Kew (Anon. 1897) and has become popular & Di Giusto 2009, Bauer et al. 2011; Fig. 1, 2; Table 1, 2), how- in horticulture as a practical means of distinguishing the two ever, their functional significance has not been addressed in taxa. However, no detailed description of N. rafflesiana var. recent taxonomic revisions of the genus (Jebb & Cheek 1997, elongata was ever published and no type is extant at Kew Cheek & Jebb 2001). (Cheek & Jebb 2001). Accordingly, there is no evidence that Researchers have been intrigued by ecological differences the name N. rafflesiana var. elongata refers to the ‘elongate between the ‘typical’ and elongate forms of N. rafflesiana for form’ of N. rafflesiana from Borneo. Clarke (1997) and Phillipps some time – these are summarised in Table 1. Notably, Grafe et al. (2008) discussed this taxon under their comments for et al. (2011) have demonstrated that the elongate form utilises N. rafflesiana, mentioning the use of the name ‘N. rafflesiana a unique, facultative nitrogen (N) acquisition strategy, in which var. elongata’ in horticulture, but did not suggest that this was Hardwicke’s Woolly Bats (Kerivoula hardwickii Horsfield, the correct name for the taxon. (Chiroptera: Vespertilionidae)) roost and defecate in its upper On the basis of the findings of Moran (1996), Gaume & Di Giusto pitchers. Bat faeces potentially represents an important nutri- (2009), Grafe et al. (2011) and Bauer et al. (2011), we contend that the differences between these two taxa are of sufficient 1 School of Science, Monash University Sunway Campus, Jalan Lagoon importance for the elongate form to be distinguished from Selatan, 46150 Bandar Sunway, Selangor, Malaysia; N. rafflesiana at specific rank. Although the visible morpho- corresponding author e-mail: [email protected]. 2 School of Environment and Sustainability, Royal Roads University, 2005 logical differences between them are comparatively slight, Sooke Road, Victoria, BC, V9B 5Y2, Canada. there are additional differences in ontogeny, physiology and 3 Peti Surat 2507, 93750 Kuching, Sarawak, Malaysia. chemistry (see Table 1, 2) that are consistent and appear to be © 2011 Nationaal Herbarium Nederland You are free to share - to copy, distribute and transmit the work, under the following conditions: Attribution: You must attribute the work in the manner specified by the author or licensor (but not in any way that suggests that they endorse you or your use of the work). Non-commercial: You may not use this work for commercial purposes. No derivative works: You may not alter, transform, or build upon this work. For any reuse or distribution, you must make clear to others the license terms of this work, which can be found at http://creativecommons.org/licenses/by-nc-nd/3.0/legalcode. Any of the above conditions can be waived if you get permission from the copyright holder. Nothing in this license impairs or restricts the author’s moral rights. 230 Blumea – Volume 56 / 3, 2011 Table 1 List of differences in pitcher characteristics and ecology of N. rafflesiana and N. baramensis. Differences are categorised on the basis of cause and effect. ‘Causal’ differences include characteristics that have roles in the detectable ecological differences between the taxa (i.e., the ‘effects’). N. rafflesiana N. baramensis Names used in previous studies: ‘Typical’ form; var. typica ‘Elongate’ form; var. elongata ‘Causal’ differences Epicuticular wax zone on upper part of inner pitcher wall reduced as plant Epicuticular wax zone retained throughout ontogeny1 matures; absent from upper pitchers of mature plants1 Copious nectar secretion by extrafloral nectaries of the peristome2 Little nectar secreted (up to seven times less)2 Complex colour contrast pattern corresponding to insect visual sensitivity Simple colour contrast pattern3 maxima in ultraviolet, blue and green wavebands3 Upper pitchers produce insect-attracting fragrance4,5,6 No fragrance produced4,5,6 ‘Effect’ differences High diversity of invertebrate prey1,4,5 Low diversity of invertebrate prey1,4,5 High prey capture rate1,5 Low prey capture rate1,5 Primary prey-trapping structure: peristome2 (plus viscoelastic pitcher fluid4) Primary prey-trapping structure: epicuticular wax zone2 No facultative mutualism with bats7 Facultative mutualism with bats7 1 Gaume & Di Giusto 2009; 2 Bauer et al. 2011; 3 See Fig. 2, this paper; 4 Di Giusto et al. 2008; 5 Moran 1996; 6 Di Giusto et al. 2010; 7 Grafe et al. 2011. Table 2 Diagnostic characteristics for distinguishing N. rafflesiana and N. baramensis. Characteristic N. rafflesiana N. baramensis Texture of the upper pitchers Coriaceous Chartaceous Position of hip in upper pitchers Absent, or immediately beneath the mouth 1/3–1/2 of the way between the mouth and the base of the pitcher Structure of peristome at Prominently raised into a flat, horizontal ‘ridge’ Not raised at all at the front, or occasionally with a small kink near the front of the pitcher mouth the centre at the front Shape of upper pitchers Funnel-shaped throughout Funnel-shaped in the lower 1/2–2/3, cylindrical in the upper 1/2–1/3 Lid shape of upper pitchers Circular, strongly cordate at the base Ovate, not (or only slightly) cordate at the base Waxy zone on upper inner surface Reduced or entirely absent Always present of upper pitchers on mature plants Strong fragrance associated Present Absent with upper pitchers involved in divergent N-acquisition strategies. Accordingly, we Nepenthes baramensis C.Clarke, J.A.Moran & C.C.Lee, here describe the elongate form as a new species, Nepenthes sp. nov. baramensis sp. nov. Nepenthi rafflesianae similis sed ascidiis aeriis in parte superiore cylindricae et texturae chartaceae differt. — Holotypus: Hotta M. 12419 (SAR), Brunei, Materials AND METHODS Belait District, Seria, en route K. Badas to Seria along the railway of B.S.C., 7 Dec. 1963. Spectroradiometric analysis was carried out on the reflectance Etymology. The specific epithet elongata was used by G. Beck to describe characteristics
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