Danser's Monograph on Nepenthes: Nepenthes alata



1. Nepenthes alata BLANCO, Fl. Fil., ed. 1, p. 805 (1837 BL., Mus., II, p. 10 (1852): HOOK. F., in D.C. Prodr., XVII, p. 99 (1873) ; BLANCO, Fl. Fil., ed. 3, III, p. 214 (1879) ; FERN.-VILL., Nov. app., p. 173 (1880) ; BECC., Mal., III, p. 4 (1886) ; BECK, Wien Ill. Gartenz., 1895, p. 221 (1895) ;MERR., Phil. Journ. Sc., 1, suppl. 1, p. 59 (1906) ; MACF., in ENGL, Pflanzenr., IV, 111, p. 71 (1908) ; MERR. & MERRITT, Phil. Journ. Sc., bot, V, p. 350 (1910) ; MERR., Sp. Blanc., p. 160 (1918) ; MACF., in BAIL., Cycl., IV ; p. 2129 (1919) ; MERR., Enum. Phil., II, p. 214 (1923) ; MACF., Phil. Journ. Sc., XXXIII, p. 136 (1927): ? N. Blancoi BL., Mus., II, p. 10 (1852) ; HOOK. F., in D.C., Prodr., XVII, p. 105 (1873): BECC., Mal., III, p. 5 (1886) ; MACF., Pflanzenr., IV, 111, p. 40 (1908) ; Phil. Journ. Sc., XXXIII, p. 129 (1927) ; N. eustachya MIQ., Fl., I, 1, p. 1074 (1858) ; suppl., p. 151 (1860) ; Journ. Bot. Néerl, I, p. 277 (1861) ; TEYSM. & BINN., Cat., p. 99 (1866) ; MIQ, Ill., p. 3 & 7, t. III (1870) ; HOOK. F., in D.C., Prodr., XVII, p. 99 (1873) ; BECC., Mal., III, p. 4 (1886) ; BECK, Wien. Ill. Gartenz., 1895, p. 217 (1895) ; (eustachia) BOERL., Handl., III, 1, p. 54 (1900) ; MACF., in ENGL., Pflanzenr., IV, 111, p. 51 (1908) ; N. melamphora FERN.-VILL., Nov. app., p. 173 (1880) ; non BL,, Cat., p. 111 (1823) &c.; ? N. Copelandii MACF., in ENGL., Pflanzenr., IV, 111, p. 51 (1908) ; MERR. Enum. Phil., II, p. 214 (1923) ; MACF., Phil. Journ. Sc., XXXIII, p. 131 (1927) ; N. graciliflora ELM., Leafl., IV, p. 1494 (1912).

Icon: MIQ., Ill., t. III (1870), optima.

Folia mediocria petiolata, lamina lanceolata v. spathulato-lanceolata, nervis longitudinalibus utrinque c. 3, vagina 1/3-1/2 caulis amplectente ; ascidia rosularum parte inferiore ovata, os versus cylindrica v. leviter infundibuliformia, alis 2 saepe fimbriatis ; peristomio operculum versus acuminato v. in collum breve elongato, cylindrico v. applanato, 1-3 mm lato, costis 1/3-1/4 mm distantibus, dentibus 0 ; operculo orbiculari v. ovato, facie inferiore plerumque carinato v. appendice lateraliter applanata ; ascidia inferiora magnitudine mediocria, parte inferiore anguste ovata, medio angustata, os versus leviter infundibuliformia, alis 2 fimbriatis ; perisitomio operculum versus acuto v. acuminato, cylindrico v. applanato, ad 3 mm lato, costis c. 1/3 mm distantibus, dentibus fere 0 ; operculo ovato-rotundato, facie inferiore prope basin saepe carinato v. appendice lateraliter applanata, rarius plano ; ascidia superiora magnitudine mediocria, forma fere ut inferiora, costis 2 prominentibus, raro anguste alata ; peristomio operculum versus acuto v. acuminato, elevato, cylindrico v. applanato, 2-7 mm lato, costis c. 1/3 mm distantibus, dentibus fere 0 ; operculo cordato-ovato v. suborbiculari, facie inferiore prope basin carinato v. appendice lateraliter applanata v. rarius plano ; inflorescentia racemus longus pedicellis inferioribus l-2-floris, 10 -15 mm longis, superioribus 1-floris ; indumentum iuventute passim densum velutino-tomentosum, denique parcius v. in partibus vegetatibus saepe 0.

Stems climbing, up to 4 m high, sometimes short or prostrate, slender, the part with adult leaves cylindrical or irregularly and obtusely trigonous, 4 to 8 mm thick, the internodes 1 to 10 cm long ; at the base of older plants often short shoots and rosettes. Rosette leaves thin-coriaceous, sessile, spathulate, very small or up to 13 cm long, up to 3 1/2 cm broad, obtuse to acute, broadest at about 1/3 of the length, gradually attenuate towards the rather broad base or more abruptly contracted, dilated again near the stem and clasping more than 4/5 of it ; pennate nerves running obliquely towards the margin, irregularly reticulate, the parallel ones up to 5 (mostly less) on each side, originating from the basal part of the midrib, running parallel in the outer half of the leaf blade, approaching the margin towards the apex ; tendrils of the smallest leaves about as long as the leaf blade, those of the larger leaves shorter than half the blade, curved downwards, up to 2 mm thick. Leaves of the short shoots scattered, petiolate, thin-coriaceous, lanceolate or lanceolate-spathulate, 5 to 15 cm long including the petiole, 2 to 3 cm broad, acute, gradually attenuate into the petiole 1 to 5 cm long and narrowly winged, which forms a laterally flattened, almost wholly amplexicaul sheath ; nervation distinct, the pennate nerves numerous, running obliquely towards the margin, irregularly reticulate, the longitudinal ones mostly 3, originating from the basal part of the midrib, imperfectly parallel ; tendril shorter than the leaf blade, robust, curved downwards without curl. Leaves of the climbing stem mostly scattered, rarely alternate, petiolate, thin-coriaceous, lanceolate or lanceolate-spathulate, mostly 15 to 25 cm long including the petiole, 2 to 5 cm broad, acute or slightly emarginate, gradually attenuate into the 3 to 6 cm long, narrowly winged petiole, which forms a laterally flattened sheath, clasping 1/3 to 1/2 of the stem ; nervation distinct, the pennate nerves numerous, running irregularly and very obliquely towards the margin, irregularly reticulate, the longitudinal ones about 3 (often less) on each side, originating from the basal 1/3 to 1/5 part of the midrib, running imperfectly parallel in the outer 1/3 to 1/2 of the lamina, the outermost ones often ending before the innermost ones begin ; tendril about 1 to 1 1/2 times as long as the leaf, those with pitchers always with curl. Rosette pitchers very shortly incurved at the base, the lower 1/3 to 1/2 ovate, gradually narrowed into the cylindrical upper part, sometimes slightly infundibulate, with 2 wings over the whole length, the wings fringed or not, 2 to 3 mm broad, the fringe segments up to 3 mm long and 2 to 5 mm apart ; mouth oblique, peristome cylindrical to flattened, 1 to 3 mm broad, the ribs 1/3 to 1/4 mm apart, the interior margin entire ; inner surface of the pitcher with not-overarched glands in the ventricose part ; lid orbicular to rotundate-elliptical, up to 3 cm long and broad, with a low keel on the midrib below, with numerous small, rimmed but not deepened glands ; spur inserted close to the lid, flattened, mostly divided into few branches, 2 to 3 mm long. Lower pitchers shortly incurved at the base, ovate, up to 10 cm high, up to 2 1/2 cm wide in the lower part, narrowed in the middle to 2/3 of its width, dilated again towards the mouth, with 2 fringed wings over the whole length, the wings narrow beneath, up to 5 mm broad above, the fringe segments up to 3 mm long and 2 to 4 mm apart ; mouth very oblique, acute or acuminate and elevated towards the lid ; peristome cylindrical, up to 2 mm broad, the ribs about 1/3 mm apart, the teeth of the interior margin not so long as broad, the inner surface of the pitcher in the ventricose part with little or not overarched glands, about 500 to 600 on 1 cm2 ; lid ovate to orbicular, the midrib more or less carinate on the underside in the basal part, mostly with a laterally flattened appendage, rarely flat, with numerous glands ; spur 2 to 5 mm long, not branched, flattened, attenuate. Pitchers of the climbing stems only slightly different from the lower ones, but often higher and more slender, mostly 8 to 16, rarely up to 25 cm high, 2 1/2 to 5 cm, rarely up to 6 cm wide, narrowly ovate in the lower part, gradually narrowed towards the middle, widened again towards the mouth, mostly without wings but with 2 prominent ribs ; mouth very oblique, ovate, acute or acuminate and strongly elevated towards the lid, the peristome flattened, 2 to 7 mm broad, rarely broader, the ribs about 1/3 mm apart, the teeth of the interior margin less long than broad ; inner surface of the pitcher with not or little overarched glands in the ventricose part, about 200 to 700 on 1 cm2 ; lid ovate to orbicular, slightly cordate at the base, rounded at the apex, rarely flat on the underside mostly keeled or with a laterally flattened appendage on the midrib near the base, with small glands over the whole surface ; spur without branches, inserted close to the lid, 3 to 8 mm long, flattened, attenuate. Male inflorescence a raceme, the peduncle 10 to 30 cm long, usually 3 to 5 mm thick, the axis 10 to 30 cm long, rarely longer, gradually attenuate, the pedicels thin, 1- or 2-flowered, the lower ones up to 15 mm long, sometimes with a bract at the base, the upper ones somewhat shorter, always without bract, tepals oval, about 3 mm long. Staminal column longer than the tepals, about 5 mm long including the 1-seriate anthers. Female inflorescence chiefly like the male one but shorter on the average. Ovary sessile. Fruit 15 to 25 mm long. The valves lanceolate, 2 to 4 mm broad, attenuate towards both ends. Seeds filiform, about 8 to 10 mm long, the nucleus with transverse wrinkles. Indumentum of young parts dense and short, stellate-tomentose, later disappearing from the stems and the underside of the leaves, permanent on the inflorescences, especially on the pedicels, the inner side of the perigone always glabrous, the staminal column hairy at the base, the rest glabrous, the ovary very densely covered with tomentum and longer appressed hairs, the fruit sparingly covered by these same hairs. Colour of the pitchers light-green, often with light or dark-red or violet spots ; colour of herbarium specimens fallow-dun in different hues, the upper surface of the leaves more fallow than the underside and the pitchers. (Description after all the plants seen by the author.)

PHILIPPINE ISLANDS. (As the here after mentioned localities sufficiently show, the distribution in the Philippine Islands, I have not recorded the localities given by: BLANCO, Fl. Fil., ed. 1. p. 806 ; MACF., in ENGL., Pflanzenr., IV, 111, p. 72 ; Phil. Journ, Sc., XXXIII, p 129-137: MERR., Enum. Phil., II, p. 214 and by some other authors.) Without island: CUMING 1337, H. L. B. (0) ; 1906, LOHER 5467, H. B. (0) (cfr. LOHER 5466, further on) ; Luzon: Burgos, 23-25 XI 1916, FENIX 26726, H. B. (m), H. S. (0) ; Pauai, IV-VI 1918, SANTOS, 31877, H. B. (f) ; Benquet, Com. flor. forest. Fil. 1671, H. L. B. (0) ; XII 1908, RAMOS 5372, H. B. (0) ; Pinaglubo, 1906, LOHER 5466, H. B. (m) ; Mt. Maguiling, 27 I 1906, LOHER 5465, H. L. B. (0): VI 1914, SULIT (MERR., Sp. Blanc. no. 507), H. B. (0) ; Los Banos, VI-VII 1917, ELMER 17766 (authentic specimen of N. graciliflora ELM.) H. B. (0), H. A. R. T. (0) ; Umiray, VI 1914, LOHER 13988 H. B. (f) ; Irosin, Mt. Bulusan, IV 1916, ELMER 15847. H. B. (m, f) ; H. A. R. T. (m) ; Culion: X 1922, RAMOS 41301, H. B. (m) ; Sibuyan: Mt. Giting-giting, Magallenes, V 1910, ELMER 12465 (type of N. graciliflora ELM.) H. B. (m) ; H. A. R. T. (m). Panay: Antique, V-VIII 1918, MCGREGOR 32313, H. B. (0) ; Negros: Cuernos Mts., Dumaguete, IV 1908, ELMER 9725, H. B. (m) ; V 1908, ELMER 10073, H. B. (f) ; Mindanao: Camiguin de Mindanao, III-IV 1912, RAMOS 14650 H. B. (m) ; Mt. Urdaneta, Cabadbaran, X 1912, ELMER 14248, H. B. (0) ; Mt. Apo, Todaya. VII 1909, ELMER 11523, H. B. (0).

MALAY PENINSULA. Pahang: G. Tahan VII 1911, RIDLEY 16097, H. S. (f).

SUMATRA. Govt. Eastcoast: Bila on the Mana Lesé (Aèk Boero), 200 m, 25 IV 1925, LÖRZING 11603, H. B. (m), vern. name, Batak: dahoel-dahoel ; Res. Tapiannoeli: Sibaraboeai, HAGAN, H. B. (0): Sibolga, on the coast, II 1856, TEYSMANN 529, H. B. (m) ; H. A. R. T. (m) ; authentic specimen of N. eustachya MIQ., vern. name, Minangkabau: katoepat baroek ; Res. Westcoast: Padang Uplands, BURCK, H. B. (0) ; Berani, BÜNNEMEIJER 3366, H. B. (0) ; Mangani, 1200 m, 15 IV 1924, DE BOER 11, H. B. (0) ; Bt. Tinggi, near Mangani, 1100 m, 15 VI 1918, BÜNNEMEIJER 3054, H. B. (0) ; Tanang Taloe, 1600 m, 15 VI 1917, BÜNNEMEIJER 1049, H. B. (0).

? MOLUCCAS. Res. Amboina: Amboina, BOTTER, H. B. (0) ; for this see the discussion.

N. eustachya MIQ., only recorded from Sumatra and still distinguished by MACFARLANE, is united with N. alata in the above. In his monograph, MACFARLANE places N. alata in the group with carinate lid, N. eustachya among the species without keel on the lid ; yet he distinguishes a N. alata var. ecristata, without keel. For the rest there is hardly any difference to be stated between these two species and especially the inflorescences are strikingly alike. The specimen recorded by me from the Malay Peninsula deviates more, especially by the long, narrow inflorescence and 2-flowered pedicels, but also in the Philippines and Sumatra forms with 2-flowered pedicels have been found (RAMOS 14650, LÖRZING 11603).

N. Blancoi is based by BLUME upon the Nepenthes sp. "BLANCO Fl. Fil., p. 808 in nota". However, BLANCO mentions 2 unnamed species, one from the mountains of Agoo and Pangasinan, of which he has only seen female flowers, and another from Cebu, of which he has only seen male flowers. lt is possible, that both are forms of N. alata, but since this cannot be proved, it remains uncertain whether N. Blancoi is synonymous to N. alata. This seems to have never been noticed by any author except by MACFARLANE, in his last publication, in which however, he keeps N. Blancoi apart from N. alata on insufficient arguments.

N. alata is not very polymorphous ; the form, however, of the operculum varies much more than in other species: it is orbicular to rather narrowly-ovate, carinate or without any keel. The Sumatra plants mostly have a suborbicular and often an ecarinate lid, the Philippine plants usually an ovate, carinate one. The varieties ecristata and biflora, distinguished by MACFARLANE, are not more than extreme variations.

The distribution of N. alata is very peculiar. Though it has a continuous area, it is remarkable that it is not recorded from Borneo. If N. philippinensis has to be united with N. alata, the latter at least occurs in Palawan. That it grows in Amboina is very improbable ; the only pitcher, found in the Buitenzorg Herbarium among plants of N. mirabilis, is probably intermixed unintentionally.

N. alata, like most species of Nepenthes is a mountain plant. MERRILL first (Sp. Blanc. p. 160) says, that it occurs in the mossy forest between 800 and 2000 m above the sea level, but later (Enum. Phil., II, p. 214) ; above 1400 m, whereas MACFARLANE in his monograph gives: up to 2400 m. In Sumatra it grows from the rocks along the coast up to a height of 1600 m ; its habitat is the forest or its margin, rarely open ground.

Vernacular names. Near Aèk Boero (Batak): tahoel-tahoel ; near Sibolga (Minangkabau): katoepat baroek ; TEYSMANN says all species of Nepenthes near Sibolga bear the names: katoepat baroek, tjalong baroek and tahoel-tahoel ; cf. N. Treubiana. Katoepat baroek means monkey's rice packet: perhaps BURBIDGE gives the explication of this name (Journ. Roy. Hort. Soc., XXI, p. 256): 'When I was staying with the headman of the Kadyans on the Lawas River, his people often gave me delicious rice cooked in the pitchers of N. Hookeriana, as a sweetmeat to be eaten with jungle fruit and bananas." With this N. Hookeriana is probably meant N. Rafflesiana. Tjalong baroek means monkeys' water scooper. The name katoepe baroek as given by DE CLERCQ seems to be wrong.


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