The earliest known collection of N. tobaica was made by Johannes Elias Teijsmann on February 8, 1856, probably from the Batak regions. This specimen, which includes female floral material, is H. L. B. 908,155-1106.[2]
"N. tobaica has only been found on the plateau north, east and south of Lake Toba. It is most closely related to N. Reinwardtiana and I am not quite certain, whether it is perhaps a form of this species, but up till now no intermediate forms have been discovered."
In the scientific literature, N. tobaica has been confused with the closely related N. angasanensis on several occasions. Nepenthes tobaica as described in Danser's 1940 description of N. densiflora[3] is actually N. angasanensis.[5] Specimens identified as N. tobaica in Rusjdi Tamin and Mitsuru Hotta's 1986 monograph[4] on SumatranNepenthes actually represent both N. angasanensis and N. tobaica.[5]
Nepenthes tobaica is a climbing plant. The stem can attain a height of up to 7 m and is up to 6 mm in diameter. Internodes are up to 25 cm long and often round in cross section. However, mature plants may have angular stems because of a groove that originates at the node and extends across most of the internode's length.[5]
Leaves are coriaceous in texture and range from sessile to sub-petiolate. The shape of the lamina, or leaf blade, varies from oblong to spathulate. It measures up to 20 cm in length by 4 cm in width. The apical end of the lamina is typically rounded, but may be narrowed and obtuse. Robust plants sometimes have a sub-peltate tendril insertion. The lamina may be gradually or abruptly contracted towards the amplexicaul base, which clasps the stem for around half of its circumference. One to three longitudinal veins are present on either side of the midrib, although they are only prominent in robust specimens. Pinnate veins are indistinct. Tendrils grow to 30 cm in length.[5]
Rosette and lower pitchers are ovoid in the lower portion and cylindrical above. They are up to 20 cm high by 4 cm wide. A pair of wings runs down the pitcher's ventral surface, often bearing fringe elements either throughout the whole length or only in the upper part. The glandular region of the pitcher's inner surface is confined to the ovoid portion. A pair of "eye spots" is sometimes present on the inner surface, below the lid attachment. The pitcher mouth is ovate and has an oblique insertion. The narrow peristome (≤5 mm wide) is cylindrical or slightly expanded and bears indistinct teeth. The pitcher lid or operculum is ovate to sub-orbicular and has a somewhat cordate base. It lacks appendages. A spur measuring up to 5 mm in length is inserted near the base of the lid. It may be unbranched, bifid, or trifid.[5]
Upper pitchers are typically somewhat infundibular in the lowermost part, becoming narrowly ovoid in the lower third, and finally cylindrical and slightly narrower above. They may be quite large, reaching 25 cm in height by 5 cm in width.[5] The waxy zone of the inner surface is well developed.[8] A pair of remnant ribs is present in place of the ventral wings.[5] The inner portion of the peristome accounts for around 40% of its total cross-sectional surface length.[8] Other parts of aerial pitchers are similar to their lower counterparts.[5]
Nepenthes tobaica has a racemoseinflorescence. The peduncle and rachis can each grow to 20 cm in length. Partial peduncles are two-flowered and lack bracteoles. In male inflorescences, the sepals are elliptic-obtuse, being slightly narrower in female ones.[5] Around the town of Prapat, plants have been observed to come into flower in April, although mature fruits are not common at this time.[9]
Nepenthes tobaica is characterised by an uneven indumentum. Most mature vegetative parts are glabrous, although the midrib may bear persistent hairs. Groups of white, stellate hairs are often present in the leaf axils. The sepals of this species are densely tomentose, but the rest of the inflorescence has a sparser covering of short hairs.[5]
In some forms, the underside of the lid is a vivid red, making the plants particularly easy to spot amongst other vegetation.[9]
Ecology
A lithophytic rosette plant with dark purple lower pitchers
Nepenthes tobaica commonly grows among dense populations of ferns
This species typically grows terrestrially in exposed areas.[5]Nepenthes tobaica commonly grows in forest edges, where it is often sympatric with species of Leptospermum and Rhodomyrtus.[1]
The pitchers of N. tobaica play host to a wide variety of infaunal organisms.
The mite species Creutzeria tobaica was described from the pitchers of a JavaneseNepenthes identified as N. tobaica.[11][12] However, N. tobaica has not been recorded outside of Sumatra.[5]
Differences between N. angasanensis, N. mikei and N. tobaica (Salmon & Maulder, 1999)
Character
N. angasanensis
N. mikei
N. tobaica
Habit
Produces offshoots from underground rhizomes
No rhizomes
No rhizomes
Spur
Forked
Fasciculate
Filiform
Inner margin of peristome
Teeth to 1.5–2 mm long
Teeth to 0.2-0.4 mm long
Teeth < 0.2 mm
Stem cross section
Cylindrical
Cylindrical
Cylindrical to obtusely triangular
Bracteoles
Sometimes near base of lowest pedicel only
Half way up every pedicel
At base or slightly below pedicel attachment, few
Pitcher glands
300 / cm2
150–180 / cm2
200–250 / cm2
Pedicels
1-flowered
1-flowered
2-flowered
Inflorescence (female)
55–125 mm long, 9-17 flowers
40–80 mm long, 4-10 flowers
195–400 mm long, 30-50 flowers
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". The sister pair of N. angasanensis and N. mikei has 79% support.[5]
a.^ The Latin description of N. tobaica from Danser's monograph reads:[2]
Folia mediocria sessilia, lamina lineari-lanceolata, nervis longitudinalibus utrinque 0-1, raro 2, basi lata semiamplexicauli ; ascidia rosularum ignota ; ascidia inferiora parte inferiore anguste ovata, medio angustata, os versus infundibuliformia, alis 2 fimbriatis, peristomio operculum versus acuto, cylindrico, 1/2-1 mm lato, costis 1/3-1/4 mm distantibus, saepe indistinctis, dentibus fere 0 ; operculo rotundato-elliptico, facie inferiore plano ; ascidia superiora parva, e basi infundibuliformi parte inferiore paulum ventricosa, medio leviter angustata, os versus paulum infundibuliformia, costis 2 prominentibus, peristomio operculum versus acuto, cylindrico, 1/2-1 mm lato, costis 1/3-1/4 mm distantibus, saepe indistinctis, dentibus fere 0 ; operculo orbiculari v. rotundato-elliptico facie interiore plana ; inflorescentia racemus pedicellis 13-15 mm longis fere omnibus 2-floris ; indumentum in inflorescentiis et ascidiis parcum stellatum, ceterum 0.
b.^ In addition to the earliest specimen, collected by Teijsmann in 1856, Danser lists 11 separate collections of N. tobaica.
^ abDanser, B.H. 1940. A new Nepenthes from Sumatra. Bulletin du Jardin Botanique de Buitenzorg, Série III, 16: 268–271.
^ ab(in Indonesian) Tamin, R. & M. Hotta 1986. Nepenthes di Sumatera: The genus Nepenthes of the Sumatra Island. In: M. Hotta (ed.) Diversity and Dynamics of Plant Life in Sumatra: Forest Ecosystem and Speciation in Wet Tropical Environments. Part 1: Reports and Collection of Papers. Kyoto University, Kyoto. pp. 75–109.
^Catalano, M. 2009. Nepenthes. In: Growing Carnivores — an Italian perspective. Prague. pp. 50–57.
^ abBauer, U., C.J. Clemente, T. Renner & W. Federle 2012. Form follows function: morphological diversification and alternative trapping strategies in carnivorous Nepenthes pitcher plants. Journal of Evolutionary Biology25(1): 90–102. doi:10.1111/j.1420-9101.2011.02406.x
Bonhomme, V., H. Pelloux-Prayer, E. Jousselin, Y. Forterre, J.-J. Labat & L. Gaume 2011. Slippery or sticky? Functional diversity in the trapping strategy of Nepenthes carnivorous plants. New Phytologist191(2): 545–554. doi:10.1111/j.1469-8137.2011.03696.x
Meimberg, H., P. Dittrich, G. Bringmann, J. Schlauer & G. Heubl 2000. Molecular phylogeny of Caryophyllidae s.l. based on matK sequences with special emphasis on carnivorous taxa. Plant Biology2(2): 218–228. doi:10.1055/s-2000-9460
Meimberg, H., A. Wistuba, P. Dittrich & G. Heubl 2001. Molecular phylogeny of Nepenthaceae based on cladistic analysis of plastid trnK intron sequence data. Plant Biology3(2): 164–175. doi:10.1055/s-2001-12897
Meimberg, H. & G. Heubl 2006. Introduction of a nuclear marker for phylogenetic analysis of Nepenthaceae. Plant Biology8(6): 831–840. doi:10.1055/s-2006-924676
Meimberg, H., S. Thalhammer, A. Brachmann & G. Heubl 2006. Comparative analysis of a translocated copy of the trnK intron in carnivorous family Nepenthaceae. Molecular Phylogenetics and Evolution39(2): 478–490. doi:10.1016/j.ympev.2005.11.023
Renner, T. & C.D. Specht 2011. A sticky situation: assessing adaptations for plant carnivory in the Caryophyllales by means of stochastic character mapping. International Journal of Plant Sciences172(7): 889–901. doi:10.1086/660882
Thornhill, A.H., I.S. Harper & N.D. Hallam 2008. The development of the digestive glands and enzymes in the pitchers of three Nepenthes species: N. alata, N. tobaica, and N. ventricosa (Nepenthaceae). International Journal of Plant Sciences169(5): 615–624. doi:10.1086/533599