Nepenthes gracilis

Nepenthes gracilis
A pitcher of Nepenthes gracilis from Terengganu, Peninsular Malaysia
Scientific classification Edit this classification
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Order: Caryophyllales
Family: Nepenthaceae
Genus: Nepenthes
Species:
N. gracilis
Binomial name
Nepenthes gracilis
Korth. (1839)[2]
Synonyms

Nepenthes gracilis (/nɪˈpɛnθz ˈɡræsɪlɪs/; from Latin: gracilis "slender"), or the slender pitcher-plant,[5] is a common lowland pitcher plant that is widespread in the Sunda region. It has been recorded from Borneo, Cambodia,[6] Peninsular Malaysia, Singapore, Sulawesi, Sumatra, and Thailand.[7][8][9][10][11] The species has a wide altitudinal distribution of 0 to 1100 m[12] (and perhaps even 1700 m[13]) above sea level, although most populations are found below 100 m and plants are rare above 1000 m.[13] Despite being a widespread plant, natural hybrids between N. gracilis and other species are quite rare.

Nepenthes gracilis was formally described by Pieter Willem Korthals in his 1839 monograph, "Over het geslacht Nepenthes".[2]

Nepenthes abgracilis from the Philippines is named for its superficial similarity to this species.[14]

Carnivory

The small, elongated pitchers of N. gracilis appear relatively unremarkable and have a very thin peristome. Nevertheless, the species is unusual (and possibly unique) in that the underside of the pitcher lid bears an uneven layer of wax crystals. This layer is not as thick as, and structurally distinct from, that found in the waxy zone of the pitcher interior, and insects can easily adhere to it in dry conditions. During downpours, however, it functions as part of a trapping mechanism, whereby the impact of raindrops striking the lid causes insects to lose their footing and fall into the pitcher cup below.[15]

Left: Nepenthes gracilis upper pitcher with a Polyrhachis pruinosa ant (A and B), showing the waxy zones of the pitcher interior and lower lid surface.

Centre: Scanning electron micrographs revealing the ultrastructure of N. gracilis wax crystals on the inner surface of the pitcher (A and B) and underside of the lid (C and D).

Right: Prey capture efficiency of the two waxy layers and peristome of N. gracilis under 'dry', 'raining' and 'wet' conditions.

Distribution

Nepenthes gracilis habitat in Pahang, Peninsular Malaysia

One of the most widespread Nepenthes species, N. gracilis is native to Borneo, Cambodia,[6] Peninsular Malaysia, Singapore, Sumatra, central Sulawesi, and southernmost Thailand.[7] It has also been recorded from many smaller islands, including Bangka, Batu Islands, Belitung,[16] Bengkalis, Ko Lanta, Ko Tarutao, Labuan,[17] Langkawi, Mendol, Mentawai Islands (Siberut), Meranti Islands (Padang, Rangsang, and Tebing Tinggi), Musala, Nias, Penang, Phuket, Riau Islands (Lingga Islands, Natuna Islands,[18] and Riau Archipelago), and Rupat.[19]

Evolutionary origins

In 2001, Charles Clarke performed a cladistic analysis of the Nepenthes species of Sumatra and Peninsular Malaysia using 70 morphological characteristics of each taxon. The following is a portion of the resultant cladogram, showing part of "Clade 6", which includes N. gracilis.[20]

unnamed

N. gracilis

N. reinwardtiana

unnamed

When molecular genetic DNA sequence data instead of morphological characters are used to perform similar analyses,[21][22] it becomes evident that N. gracilis is not closely related to any of these 4 species. Rather, N. gracilis is the basal lineage within a "N. mirabilis group", a clade that also includes N. bicalcarata, a group of species around N. tomoriana (several species from Sulawesi, Papua and New Caledonia), N. ampullaria, N. mirabilis and N. rafflesiana sensu lato. The contrast between molecular and morphological phylogenies implies that the morphological characters evolved quite fast and repeatedly, and do not necessarily reflect shared ancestry of the species.

Infraspecific taxa

Despite varying little across its range,[20] N. gracilis has a number of infraspecific taxa. Most of these are no longer considered valid.

  • Nepenthes gracilis f. angustifolia (Mast.) Hort.Westphal (1993)
  • Nepenthes gracilis var. angustifolia (Mast.) Hort.Weiner in sched. (1985)
  • Nepenthes gracilis var. arenaria Ridl. ex Macfarl. (1908)[23]
  • Nepenthes gracilis var. elongata Blume (1852)[24]
  • Nepenthes gracilis var. longinodis Beck (1895)[4]
  • Nepenthes gracilis var. major Hort.Van Houtte ex Rafarin (1869)
  • Nepenthes gracilis var. teysmanniana (Miq.) Beck (1895)[4]

Natural hybrids

The following natural hybrids involving N. gracilis have been recorded.

References

  1. ^ Clarke, C.M. (2018). "Nepenthes gracilis". IUCN Red List of Threatened Species. 2018: e.T39663A143960417. doi:10.2305/IUCN.UK.2018-1.RLTS.T39663A143960417.en. Retrieved 19 November 2021.
  2. ^ a b (in Dutch) Korthals, P.W. 1839. Over het geslacht Nepenthes. In: C.J. Temminck 1839–1842. Verhandelingen over de Natuurlijke Geschiedenis der Nederlandsche overzeesche bezittingen; Kruidkunde. Leiden. pp. 1–44, t. 1–4, 13–15, 20–22.
  3. ^ Masters M.T. (1881). "New garden plants. Nepenthes angustifolia (Mast.), sp. nov". The Gardeners' Chronicle. 16 (408): 524.
  4. ^ a b c (in German) Beck, G. 1895. Die Gattung Nepenthes. Wiener Illustrirte Garten-Zeitung 20(3–6): 96–107, 141–150, 182–192, 217–229.
  5. ^ Phillipps, A. & A. Lamb 1996. Pitcher-Plants of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  6. ^ a b Mey, F.S. 2016. The beautiful Nepenthes kampotiana x bokorensis. Strange Fruits: A Garden's Chronicle, 5 October 2016.
  7. ^ a b c d e McPherson, S.R. 2009. Pitcher Plants of the Old World. 2 volumes. Redfern Natural History Productions, Poole.
  8. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sulawesi. Redfern Natural History Productions, Poole.
  9. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Borneo. Redfern Natural History Productions, Poole.
  10. ^ (in Italian) Catalano, M. 2010. Nepenthes della Thailandia: Diario di viaggio. Prague.
  11. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Sumatra and Java. Redfern Natural History Productions, Poole.
  12. ^ McPherson, S.R. & A. Robinson 2012. Field Guide to the Pitcher Plants of Peninsular Malaysia and Indochina. Redfern Natural History Productions, Poole.
  13. ^ a b Adam, J.H., C.C. Wilcock & M.D. Swaine 1992. "The ecology and distribution of Bornean Nepenthes" (PDF). Archived from the original (PDF) on 2011-07-22. Journal of Tropical Forest Science 5(1): 13–25.
  14. ^ Cheek M., Jebb M. (2013). "The Nepenthes micramphora (Nepenthaceae) group, with two new species from Mindanao, Philippines". Phytotaxa. 151 (1): 25–34. doi:10.11646/phytotaxa.151.1.2.
  15. ^ a b Bauer U., Di Giusto B., Skepper J., Grafe T.U., Federle W. (2012). "With a flick of the lid: a novel trapping mechanism in Nepenthes gracilis pitcher plants". PLOS ONE. 7 (6): e38951. Bibcode:2012PLoSO...738951B. doi:10.1371/journal.pone.0038951. PMC 3374759. PMID 22719998.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  16. ^ (in Indonesian) Hidayat, S., J. Hidayat, Hamzah, E. Suhandi, Tatang & Ajidin 2003. Analisis vegetasi dua jenis tumbuhan pemakan serangga di Padang Pinang Anyang, Pulau Belitung. [Vegetation analysis of two insectivorous plants in Padang Pinang Anyang, Belitung Island.] Biodiversitas 4(2): 93–96.
  17. ^ Burbidge F.W. (1882). "Notes on the new Nepenthes". The Gardeners' Chronicle. 17 (420): 56.
  18. ^ (in Indonesian) Mansur, M. 2012. Keanekaragaman jenis tumbuhan pemakan serangga dan laju fotosintesisnya di Pulau Natuna. [Diversity on insectivorous plants and its photosynthetic rate in Natuna Island.] Berita Biologi 11(1): 33–42. Abstract Archived 2013-09-21 at the Wayback Machine
  19. ^ Clarke, C.M. 2001. Appendix C: Distribution Maps. In: Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu. pp. 299–307.
  20. ^ a b c Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.
  21. ^ Scharmann, Mathias; Wistuba, Andreas; Widmer, Alex (October 2021). "Introgression is widespread in the radiation of carnivorous Nepenthes pitcher plants". Molecular Phylogenetics and Evolution. 163: 107214. doi:10.1016/j.ympev.2021.107214. hdl:20.500.11850/494608. ISSN 1095-9513. PMID 34052438.
  22. ^ Murphy, Bruce; Forest, Félix; Barraclough, Timothy; Rosindell, James; Bellot, Sidonie; Cowan, Robyn; Golos, Michal; Jebb, Matthew; Cheek, Martin (March 2020). "A phylogenomic analysis of Nepenthes (Nepenthaceae)". Molecular Phylogenetics and Evolution. 144: 106668. doi:10.1016/j.ympev.2019.106668. ISSN 1095-9513. PMID 31682924.
  23. ^ Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. Das Pflanzenreich IV, III, Heft 36: 1–91.
  24. ^ (in Latin) Blume, C.L. 1852. Ord. Nepenthaceae. In: Museum Botanicum Lugduno-Batavum, sive stirpium exoticarum novarum vel minus cognitarum ex vivis aut siccis brevis expositio. Tom. II. Nr. 1. E.J. Brill, Lugduni-Batavorum. pp. 5–10.
  25. ^ a b c d Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
  26. ^ Fretwell S (2013). "Back in Borneo to see giant Nepenthes. Part 2: Mt Tambuyukon and Poring". Victorian Carnivorous Plant Society Journal. 108: 6–15.
  27. ^ Lee, C.C. 2004. Nepenthes. In: Sarawak Bau Limestone Biodiversity. H.S. Yong, F.S.P. Ng and E.E.L. Yen (eds). The Sarawak Museum Journal Vol. LIX, No. 80; Special Issue No. 6: 71–77.
  28. ^ Tan W.K., Wong C.L., Frazier C.K. (1996). "Nepenthes × (rafflesiana and gracilis)?". Nature Malaysiana. 21: 82–85.{{cite journal}}: CS1 maint: multiple names: authors list (link)
  29. ^ Bednar, B.L. 1985. "Nepenthesdominii and var. intermedia" (PDF). Carnivorous Plant Newsletter 14(4): 105–106.

Further reading