Nepenthes gracilis

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.{{cite journal | author = Bauer U., Di Giusto B., Skepper J., Grafe T.U., Federle W. | year = 2012 | title = With a flick of the lid: a novel trapping mechanism in Nepenthes gracilis pitcher plants | journal = PLOS ONE | volume = 7 | issue = 6| page = e38951 | doi = 10.1371/journal.pone.0038951 | pmid = 22719998 | pmc = 3374759 | bibcode = 2012PLoSO...738951B | doi-access = free }}

{{multiple image

| align = center

| footer = 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.

| image1 = Nepenthes gracilis with ant.png

| width1 = 400

| image2 = Microstructure of N. gracilis waxy surfaces.png

| width2 = 251

| image3 = Contribution of the individual N. gracilis pitcher surfaces to prey capture under different environmental conditions cropped.png

| width3 = 203

}}

{{Gallery

|title=Trapping function of N. gracilis pitcher lid under experimental conditions

|width=320

|align=center

|File:Nepenthes gracilis video1.ogv|Ants being flicked off the waxy lower surface of the lid by the action of simulated rain

|File:Nepenthes gracilis video2.ogv|The same experiment carried out with an isolated lid held in place with a paper clip

|File:Nepenthes gracilis video3.ogv|High-speed recording showing a house fly (Musca domestica) falling into a pitcher following raindrop impact (recording frame rate: 428 s⁻¹, playback frame rate: 10 s⁻¹)

}}

File:Nepenthes gracilis13.jpg, Peninsular Malaysia]]

One of the most widespread Nepenthes species, N. gracilis is native to Borneo, Cambodia, Peninsular Malaysia, Singapore, Sumatra, central Sulawesi, and southernmost Thailand. It has also been recorded from many smaller islands, including Bangka, Batu Islands, Belitung,{{in lang|id}} Hidayat, S., J. Hidayat, Hamzah, E. Suhandi, Tatang & Ajidin 2003. [http://biodiversitas.mipa.uns.ac.id/D/D0402/D040205.pdf 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. Bengkalis, Ko Lanta, Ko Tarutao, Labuan,{{cite journal | author = Burbidge F.W. | year = 1882 | title = Notes on the new Nepenthes | url = https://www.biodiversitylibrary.org/page/25598125 | journal = The Gardeners' Chronicle | volume = 17 | issue = 420| page = 56 }} Langkawi, Mendol, Mentawai Islands (Siberut), Meranti Islands (Padang, Rangsang, and Tebing Tinggi), Musala, Nias, Penang, Phuket, Riau Islands (Lingga Islands, Natuna Islands,{{in lang|id}} 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. [http://digilib.biologi.lipi.go.id/view.html?idm=80840 Abstract] {{webarchive|url=https://web.archive.org/web/20130921054353/http://digilib.biologi.lipi.go.id/view.html?idm=80840 |date=2013-09-21 }} and Riau Archipelago), and Rupat.Clarke, C.M. 2001. Appendix C: Distribution Maps. In: Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu. pp. 299–307.

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.Clarke, C.M. 2001. Nepenthes of Sumatra and Peninsular Malaysia. Natural History Publications (Borneo), Kota Kinabalu.

{{clade| style=font-size:100%;line-height:80%

|label1=

|1={{clade

|label1=unnamed

|1={{clade

|1=N. gracilis

|2=N. reinwardtiana

}}

|label2=unnamed

|2={{clade

|label1=

|1={{clade

|1=N. tobaica

|label2= 79%

|2={{clade

|1=N. angasanensis

|2=N. mikei

}}

}}

}}

}}

}}

When molecular genetic DNA sequence data instead of morphological characters are used to perform similar analyses,{{Cite journal |last1=Scharmann |first1=Mathias |last2=Wistuba |first2=Andreas |last3=Widmer |first3=Alex |date=October 2021 |title=Introgression is widespread in the radiation of carnivorous Nepenthes pitcher plants |url=https://pubmed.ncbi.nlm.nih.gov/34052438/ |journal=Molecular Phylogenetics and Evolution |volume=163 |pages=107214 |doi=10.1016/j.ympev.2021.107214 |issn=1095-9513 |pmid=34052438|hdl=20.500.11850/494608 |hdl-access=free }}{{Cite journal |last1=Murphy |first1=Bruce |last2=Forest |first2=Félix |last3=Barraclough |first3=Timothy |last4=Rosindell |first4=James |last5=Bellot |first5=Sidonie |last6=Cowan |first6=Robyn |last7=Golos |first7=Michal |last8=Jebb |first8=Matthew |last9=Cheek |first9=Martin |date=March 2020 |title=A phylogenomic analysis of Nepenthes (Nepenthaceae) |url=https://pubmed.ncbi.nlm.nih.gov/31682924/ |journal=Molecular Phylogenetics and Evolution |volume=144 |pages=106668 |doi=10.1016/j.ympev.2019.106668 |issn=1095-9513 |pmid=31682924}} 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.

Despite varying little across its range, 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)Macfarlane, J.M. 1908. Nepenthaceae. In: A. Engler. Das Pflanzenreich IV, III, Heft 36: 1–91.
• Nepenthes gracilis var. elongata Blume (1852){{in lang|la}} Blume, C.L. 1852. [https://www.biodiversitylibrary.org/page/444841 Ord. Nepenthaceae.] In: [https://dx.doi.org/10.5962/bhl.title.274 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.
• Nepenthes gracilis var. longinodis Beck (1895)
• Nepenthes gracilis var. major Hort.Van Houtte ex Rafarin (1869)
• Nepenthes gracilis var. teysmanniana (Miq.) Beck (1895)

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

• N. albomarginata × N. gracilis
• N. ampullaria × N. gracilis [=N. × trichocarpa]Clarke, C.M. 1997. Nepenthes of Borneo. Natural History Publications (Borneo), Kota Kinabalu.
• (N. ampullaria × N. gracilis) × N. bicalcarata [=N. × trichocarpa × N. bicalcarata]
• N. bicalcarata × N. gracilis [=N. × cantleyi]
• ? N. eustachya × N. gracilis
• N. gracilis × N. mirabilis [=N. × sharifah-hapsahii, N. × ghazallyana, N. × grabilis, N. neglecta?]{{cite journal | author = Fretwell S | year = 2013 | title = Back in Borneo to see giant Nepenthes. Part 2: Mt Tambuyukon and Poring | journal = Victorian Carnivorous Plant Society Journal | volume = 108 | pages = 6–15 }}
• N. gracilis × N. northiana [=N. × bauensis]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.
• N. gracilis × N. rafflesiana{{cite journal | author = Tan W.K., Wong C.L., Frazier C.K. | year = 1996 | title = Nepenthes × (rafflesiana and gracilis)? | journal = Nature Malaysiana | volume = 21 | pages = 82–85 }}Bednar, B.L. 1985. {{cite web|url= http://www.carnivorousplants.org/cpn/articles/CPNv14n4p105_106.pdf |title=Nepenthes ✕ dominii and var. intermedia. }} Carnivorous Plant Newsletter 14(4): 105–106.
• N. gracilis × N. reinwardtiana
• N. gracilis × N. sumatrana

File:N. albomarginata x N. gracilis.jpg|N. albomarginata × N. gracilis

Image:Ntrichocarpa1.jpg|N. ampullaria × N. gracilis

Image:Nepenthes x cantleyi.jpg|N. bicalcarata × N. gracilis

Image:Nepentheseustachyagracilis.jpg|? N. eustachya × N. gracilis

File:N. gracilis x N. mirabilis.jpg|N. gracilis × N. mirabilis

File:Bauensis1.jpg|N. gracilis × N. northiana

File:Nepenthes gracilis x Nepenthes rafflesiana.jpg|N. gracilis × N. rafflesiana

Image:Nepenthesgracilissumatrana2.jpg|N. gracilis × N. sumatrana

{{Reflist|2}}

{{Commons category|Nepenthes gracilis}}

{{Refbegin|2}}

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{{Refend}}

{{Nepenthes}}

{{Taxonbar|from=Q150492}}

Category:Carnivorous plants of Asia

gracilis

Category:Flora of Borneo

Category:Flora of Malaya

Category:Flora of Sulawesi

Category:Flora of Sumatra

Category:Flora of Thailand

Category:Plants described in 1839

Category:Least concern plants

Category:Articles containing video clips