Heliamphora

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Heliamphora
H chimantensis2.jpg
Heliamphora chimantensis
Scientific classification OOjs UI icon edit-ltr.svg
Kingdom: Plantae
Clade: Tracheophytes
Clade: Angiosperms
Clade: Eudicots
Clade: Asterids
Order: Ericales
Family: Sarraceniaceae
Genus: Heliamphora
Benth. (1840)
Heliamphora distribution.svg
Heliamphora distribution

The genus Heliamphora ( /hɛliˈæmfərə/ or /hiːliˈæmfərə/ ; Greek: helos "marsh" and amphoreus "amphora") contains 24 species of pitcher plants endemic to South America. [1] The species are collectively known as sun pitchers, based on the mistaken notion that the heli of Heliamphora is from the Greek helios, meaning "sun". The name instead derives from the Greek helos, meaning "marsh", so a more accurate translation of their scientific name would be marsh pitcher plants. [2] Species in the genus Heliamphora are carnivorous plants that consist of a modified leaf form that is fused into a tubular shape. They have evolved mechanisms to attract, trap, and kill insects; and control the amount of water in the pitcher. At least one species ( H. tatei ) produces its own proteolytic enzymes that allows it to digest its prey without the help of symbiotic bacteria.

Morphology

All Heliamphora species are herbaceous perennial plants that grow from a subterranean rhizome. Heliamphora species form stemless rosettes and leaf height ranges from a few centimeters ( H. minor , H. pulchella ) up to more than 50 cm (20 in) ( H. ionasi , H. tatei ). [1] Heliamphora possess tubular traps formed by rolled leaves with fused edges. Marsh pitcher plants are unusual among pitcher plants in that they lack lids (opercula), instead having a small "nectar spoon" on the upper posterior portion of the leaf. This spoon-like structure secretes a nectar-like substance, which serves as a lure for insects and small animals. Each pitcher also exhibits a small drainage hole (W, E2a, E2b, and E3 Clades) or slit (E1 Clade) in its side that allows excess rainwater to drain away, similar to the overflow on a sink. [1] It was inferred that the drainage structure first evolved as drainage hole in ancestral Heliamphora populations and further modified into drainage slit in the ancestors of E1 clade. [3] This allows the marsh pitcher plants to maintain a constant maximum level of rainwater within the pitcher. The pitchers' inner surface is covered with downward-pointing hairs to force insects into the pitchers' lower parts. The morphological diversification of Heliamphora pitchers is both convergent and divergent, likely as a result of adaptive radiation in the geographically complex Guiana highland. [4]

Phylogeny of Heliamphora inferred from nuclear DNA and the evolution of morphological characters. The drainage slit is only found in the E1 clade. In the rest of Heliamphora, the drainage hole is present. The study suggested that the drainage hole is the ancestral state in Heliamphora and was further modified into drainage slit in the E1 clade. ASRHeli.png
Phylogeny of Heliamphora inferred from nuclear DNA and the evolution of morphological characters. The drainage slit is only found in the E1 clade. In the rest of Heliamphora, the drainage hole is present. The study suggested that the drainage hole is the ancestral state in Heliamphora and was further modified into drainage slit in the E1 clade.
Adaptive Radiation of Heliamphora Pitchers Morphological Diversity of Heliamphora Pitchers.png
Adaptive Radiation of Heliamphora Pitchers
Illustration of Heliamphora nutans from John Muirhead Macfarlane's 1908 monograph on the genus (A: entire plant; B: androecium; C: stamen; D: pistil; E: transverse section of the ovary; F: seed, with the testa; G: vertical section of the seed; H: embryo) Heliamphora nutans Macfarlane illustration.jpg
Illustration of Heliamphora nutans from John Muirhead Macfarlane's 1908 monograph on the genus (A: entire plant; B: androecium; C: stamen; D: pistil; E: transverse section of the ovary; F: seed, with the testa; G: vertical section of the seed; H: embryo)

Carnivory

Though often counted among the various carnivorous plants, with the exception of Heliamphora tatei, the vast majority of plants in the genus Heliamphora do not produce their own digestive enzymes (i.e. proteases, ribonucleases, phosphatases, etc.), relying instead on the enzymes of symbiotic bacteria to break down their prey. [7] They do, however, attract prey through special visual and chemical signals and trap and kill the prey through a typical pitfall trap. Field studies of H. nutans , H. heterodoxa, H. minor, and H. ionasi have determined that none of these species produce their own proteolytic enzymes. [8] H. tatei is one of the few species observed to produce both digestive enzymes and wax scales, which also aid in prey capture. [8] The pattern of carnivory among Heliamphora species, combined with habitat data, indicates that carnivory in this genus evolved in nutrient-poor locations as a means to improve absorption of available nutrients. Most Heliamphora typically capture ants, while H. tatei can capture and absorb nutrients from more flying insects. The carnivorous habit among these species is lost in low light conditions, which suggests that certain nutrient concentrations (specifically nitrogen and phosphorus) are only limiting during periods of fast growth under normal light conditions, thus rendering most of the carnivorous adaptations inefficient and not energy cost effective. [8]

Distribution

All Heliamphora species are endemic to the tepuis of the Guiana Highlands and their surrounding uplands. Most are found in Venezuela, with a few extending into western Guyana and northern Brazil. Many of the tepuis have not yet been explored for Heliamphora, and the large number of species described in recent years suggests that many more species may be awaiting discovery.

Botanical history

Flowers of H. pulchella Flower H pulchella1.jpg
Flowers of H. pulchella

The first species of the genus to be described was H. nutans, which George Bentham named in 1840 based on a specimen collected by Robert Hermann Schomburgk. This remained the only known species until Henry Allan Gleason described H. tatei and H. tyleri in 1931, also adding H. minor in 1939. Between 1978 and 1984, Julian Alfred Steyermark and Bassett Maguire revised the genus (to which Steyermark had added H. heterodoxa in 1951) and described two more species, H. ionasi and H. neblinae , as well as many infraspecific taxa. Various exploratory expeditions as well as review of existing herbarium specimens has yielded many new species in recent years, mainly through the work of a group of German horticulturalists and botanists (Thomas Carow, Peter Harbarth, Joachim Nerz and Andreas Wistuba). [9]

Care in cultivation

Heliamphora are regarded by carnivorous plant enthusiasts and experts as one of the more difficult plants to maintain in cultivation. The genus requires cool (the "highland" species) to warm (the "lowland" species) temperatures with a constant and very high humidity. [10] An amateur botanist in New York City has shown that cultivation of the genus can be achieved with an inexpensive setup consisting of a large plastic crate, a fan, egg cartons, and water bottles filled with ice. [11] The highland species, which originate from high on the humid tepui mountaintops, include H. nutans, H. ionasi, and H. tatei. The lowland Heliamphora, such as H. ciliata and H. heterodoxa have migrated to the warmer grasslands at the foot of the tepuis.

Shredded, long-fibered, or live sphagnum moss is preferred as a soil substrate, often with added horticultural lava rock, perlite, and pumice. The substrate must always be kept moist and extremely well drained. Misting Heliamphora with purified water is often beneficial to maintain high humidity levels.

Propagation through division only has a limited rate of success, as many plants that are divided go into shock and eventually die. Germination of Heliamphora seed is achieved by scattering it on milled sphagnum moss and keeping in bright light and humid conditions. Seed germination begins after many weeks.

Classification

The genus Heliamphora contains the most species in the Sarraceniaceae family and is joined by the cobra lily (Darlingtonia californica) and the North American pitcher plants (Sarracenia spp.) in that taxon.

Species

Twenty-four species of Heliamphora are currently recognized. [1] Unless otherwise stated, all information and taxonomic determinations in the table below are sourced from the 2011 work Sarraceniaceae of South America authored by Stewart McPherson, Andreas Wistuba, Andreas Fleischmann, and Joachim Nerz. [1] Authorities are presented in the form of a standard author citation, using abbreviations specified by the IPNI. [12] Years given denote the year of the species's formal publication under the current name, not the earlier basionym date of publication if one exists.

SpeciesAuthorityYearImageDistributionAltitudinal distribution
Heliamphora arenicola Wistuba, A.Fleischm., Nerz & S.McPherson [1] 2011 Venezuela Below 2000 m
Heliamphora ceracea Nerz, Wistuba, Grantsau, Rivadavia, A.Fleischm. & S.McPherson [1] 2011 Brazil Highland
Heliamphora chimantensis Wistuba, Carow & Harbarth [13] 2002 H chimantensis3.jpg Venezuela 1900–2100 m
Heliamphora ciliata Wistuba, Nerz & A.Fleischm. [14] 2009 Venezuela 900 m
Heliamphora collina Wistuba, Nerz, S.McPherson & A.Fleischm. [1] 2011 Venezuela 1700–1825 m
Heliamphora elongata Nerz [15] 2004 Venezuela 1800–2600 m
Heliamphora exappendiculata (Maguire & Steyermark) Nerz & Wistuba [16] 2006 Venezuela 1700–2100 m
Heliamphora folliculata Wistuba, Harbarth & Carow [17] 2001 H folliculata 2.jpg Venezuela 1700–2400 m
Heliamphora glabra (Maguire) Nerz, Wistuba & Hoogenstrijd [18] 2006Borderlands of Brazil, Guyana, and Venezuela 1200–2750 m
Heliamphora heterodoxa Steyerm. [19] 1951 Guyana?, Venezuela 1200–2200 m
Heliamphora hispida Nerz & Wistuba [20] 2000Border between Brazil and Venezuela 1800–3014 m
Heliamphora huberi A.Fleischm., Wistuba & Nerz [14] 2009 Venezuela 1850–2200 m
Heliamphora ionasi Maguire [21] 1978 H ionasii.jpg Venezuela 1800–2600 m
Heliamphora macdonaldae Gleason [22] 1931 Venezuela 1500–2300 m
Heliamphora minor Gleason [23] 1939 Heliminor.JPG Venezuela 1650–2500 m
Heliamphora neblinae Maguire [21] 1978Border between Brazil and Venezuela 860–2200 m
Heliamphora nutans Benth. [24] 1840 Heliamphora nutans.jpg Borderlands of Brazil, Guyana, and Venezuela 2000–2700 m
Heliamphora parva (Maguire) S.McPherson, A.Fleischm., Wistuba & Nerz [1] 2011 Venezuela 1750–2200 m
Heliamphora pulchella Wistuba, Carow, Harbarth & Nerz [25] 2005 H pulchella1.jpg Venezuela 1850–2550 m
Heliamphora purpurascens Wistuba, A.Fleischm., Nerz & S.McPherson [1] 2011 Heterodoxa pt12.JPG Venezuela 2400–2500 m
Heliamphora sarracenioides Carow, Wistuba & Harbarth [26] 2005 Venezuela 2400–2450 m
Heliamphora tatei Gleason [22] 1931 Heliamphora tatei.JPG Venezuela 1700–2400 m
Heliamphora uncinata Nerz, Wistuba & A.Fleischm. [14] 2009 Venezuela 1850 m

Incompletely diagnosed taxa

A further two incompletely diagnosed taxa are known that may represent distinct species in their own right. [1]

SpeciesDistributionAltitudinal distribution
Heliamphora sp. 'Akopán Tepui' Venezuela 1800–1900 m
Heliamphora sp. 'Angasima Tepui' Venezuela 2200–2250 m

Varieties

Two varieties are currently recognised in the genus: H. minor var. pilosa and H. minor var. minor. [27] Additionally, an undescribed variant of H. pulchella , with traps lacking long retentive hairs is known from Amurí Tepui. [1]

Cultivars

There are currently four registered Heliamphora cultivars including Heliamphora 'Cyclops' (A. Smith), Heliamphora 'Patasola' (B. Tincher & J. Lei), Heliamphora 'Red Mambo' (F. Boulianne), and Heliamphora 'Scylla' (I. Bogdanow). [28] [29]

Natural hybrids

At least eleven natural hybrids have been recorded: [1]

Additionally, putative complex hybrids occur on the Neblina Massif among populations of H. ceracea , H. hispida , H. neblinae , and H. parva . [1] Putative crosses between H. macdonaldae and H. tatei have also been recorded in the southern part of Cerro Duida. [30]

Phylogeny and Diversification

Closely related species tend to be geographically closely distributed. Major Heliamphora clades probably emerged through both geographical separation and dispersal in the Guiana Highlands during Miocene with more recent diversification driven by vertical displacement during the Pleistocene glacial-interglacial thermal oscillations. [31]

Evolutionary History of Heliamphora inferred from nuclear DNA. The evolutionary time is represented in million of years. The node bars represent uncertainty in ancestral age estimations. Heli phylo.jpg
Evolutionary History of Heliamphora inferred from nuclear DNA. The evolutionary time is represented in million of years. The node bars represent uncertainty in ancestral age estimations.

Related Research Articles

<span class="mw-page-title-main">Andreas Wistuba</span>

Andreas Wistuba is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. More than half of all known Heliamphora species have been described by Wistuba.

Dr. Joachim Nerz is a German taxonomist and botanist specialising in the carnivorous plant genera Heliamphora and Nepenthes. Nerz has described several new species, mostly with Andreas Wistuba.

<i>Heliamphora chimantensis</i> Species of carnivorous plant

Heliamphora chimantensis is a species of marsh pitcher plant endemic to the Chimantá Massif in Venezuela. Specifically, it has been recorded from Apacará and Chimantá Tepuis. It is thought to be more closely related to the southern growing H. tatei and H. neblinae than to any of the other species found in the Gran Sabana and its tepuis. All other species known from this region have between 10 and 15 anthers, while H. tatei, H. neblinae and H. chimantensis have around 20. However, the anthers of H. tatei and the closely related H. neblinae are 7–9 mm long, while those of H. chimantensis only reach 5 mm in length.

<i>Heliamphora folliculata</i> Species of carnivorous plant

Heliamphora folliculata is a species of Marsh Pitcher Plant endemic to the Aparaman group of tepuis in Venezuela. It grows on all four mountains: Aparaman Tepui, Murosipan Tepui, Tereke Tepui and Kamakeiwaran Tepui.

Heliamphora glabra is a species of marsh pitcher plant native to Serra do Sol in Venezuela. It was for a long time considered a form of H. heterodoxa, but has recently been raised to species rank.

<i>Heliamphora heterodoxa</i> Species of Marsh Pitcher Plant native to Gran Sabana, Venezuela

Heliamphora heterodoxa is a species of marsh pitcher plant native to Venezuela and adjacent Guyana. It was first discovered in 1944 on the slopes interlinking Ptari-tepui and Sororopan-tepui and formally described in 1951.

<i>Heliamphora hispida</i> Species of carnivorous plant

Heliamphora hispida is a species of Marsh Pitcher Plant endemic to Cerro Neblina, the southernmost tepui of the Guiana Highlands at the Brazil-Venezuela border.

<i>Heliamphora ionasi</i> Species of carnivorous plant

Heliamphora ionasi is a species of marsh pitcher plant thought to be endemic to the plateau that lies between the bases of Ilu Tepui and Tramen Tepui in Venezuela. It produces the largest pitchers in the genus, which can be up to 50 cm in height.

<i>Heliamphora minor</i> Species of carnivorous plant

Heliamphora minor is a species of marsh pitcher plant endemic to Auyán-tepui in Venezuela. As the name suggests, it is one of the smallest species in the genus. It is closely related to H. ciliata and H. pulchella.

<i>Heliamphora neblinae</i> Species of carnivorous plant

Heliamphora neblinae is a species of marsh pitcher plant endemic to Cerro de la Neblina, Cerro Aracamuni and Cerro Avispa in Venezuela. It is one of the most variable species in the genus and was once considered to be a variety of H. tatei. It is unclear whether or not there is a consensus regarding its status as a species, with at least a few researchers supporting the taxonomic revision that would elevate both H. tatei var. neblinae and H. tatei f. macdonaldae to full species status.

<i>Heliamphora nutans</i> Species of carnivorous plant

Heliamphora nutans is a species of marsh pitcher plant native to the border area between Venezuela, Brazil and Guyana, where it grows on several tepuis, including Roraima, Kukenán, Yuruaní, Maringma, and Wei Assipu. Heliamphora nutans was the first Heliamphora to be described and is the best known species.

<i>Heliamphora pulchella</i> Species of carnivorous plant

Heliamphora pulchella is a species of marsh pitcher plant endemic to the Chimanta Massif and surrounding tepuis in Venezuela. It is one of the smallest species and closely related to H. minor.

Heliamphora sarracenioides is a species of marsh pitcher plant endemic to Ptari Tepui in Bolívar state, Venezuela. Approximately 200 mature plants were observed in the type locality, however this site's true location and information regarding sympatric species has not been disclosed for conservation reasons. The species differentiates itself from others by the extremely wide pitcher lid, which resembles Sarracenia species. It is closest to H. heterodoxa and H. folliculata, from which it can be distinguished by the large lid glands and width of the pitcher lid.

<i>Heliamphora tatei</i> Species of carnivorous plant

Heliamphora tatei is a species of marsh pitcher plant endemic to Cerro Duida, Cerro Huachamacari and Cerro Marahuaca in Venezuela. It is closely related to H. macdonaldae, H. neblinae, and H. parva, and all three have in the past been considered forms or varieties of H. tatei. Like H. tatei, these species are noted for their stem-forming growth habit.

<i>Heliamphora exappendiculata</i> Species of carnivorous plant

Heliamphora exappendiculata is a species of marsh pitcher plant native to the Chimantá and Aprada Massifs of Bolívar state, Venezuela. It was for a long time considered a variety of H. heterodoxa, but has recently been raised to species rank. Pitchers collect insects on flattened pitcher mouths which function as 'landing platforms' upon which prey falls from surrounding vegetation. Also, the pitcher shape effectively collects leaf litter and organic debris which may serve as additional source of nutrition for plants, similarly to H. ionasi.H. exappendiculata hybridizes naturally with H. pulchella and H. huberi in areas within which they grow together. This species occurs in shaded conditions, apparently preferring them over other habitats. In addition, plants upon Chimanta and Amuri Tepui grow directly upon the walls of gorges and ravines where surfaces are permanently wet. In contrast to those populations, on all other tepuis and massif regions the species grows on summit savannahs and stunted or shrubby forests, though these individuals represent a minority in habitat choice.

Heliamphora uncinata is a species of Marsh Pitcher Plant endemic to Venezuela. This species of carnivorous plant is known as a pitcher plant. Individuals use tube like leaves to trap insects that slip into the bottom. At the bottom of the "pitcher" there are digestive juices which slowly digest the prey item to give the plant additional nutrients. The pitchers of this species are around 25–35 cm long, and are 8–10 cm wide at the opening. The pitcher mouth is circular in shape and the back is raised to form the lid. It is known only from the type collection, which was made in a narrow canyon on Amurí-tepui. There it grows at an elevation of approximately 1850 m on sandstone cliff faces in shady conditions. It is also found in humus pockets and cracks at this location. The only other species in the genus known to have a similar growth habit is H. exappendiculata. These two taxa also share a number of morphological features and appear to be closely related. These shared morphological features include: the shape of pitchers, the general growth pattern, and appearance of nectaries.

Heliamphora huberi is a species of Marsh Pitcher Plant endemic to the Chimantá Massif in Venezuela, where it grows at elevations of 1850–2200 m in a variety of habitats. It has thus far been recorded from Angasima-tepui, Apacará-tepui, Amurí-tepui, Acopán-tepui, and the border of Torono- and Chimantá-tepui. Due to its intermediate appearance between species related to H. minor and H. heterodoxa, it is suspected to be of hybridogenic origin.

Heliamphora arenicola is a species of marsh pitcher plant known only from the western side of the Ilu–Tramen Massif in Venezuela's Gran Sabana, where it grows at elevations of less than 2000 m. It may also occur on Karaurin Tepui.

<i>Heliamphora purpurascens</i> Species of carnivorous plant

Heliamphora purpurascens is a species of marsh pitcher plant known only from the summit area of Ptari Tepui in Venezuela, where it grows at elevations of 2400–2500 m.

<i>Sarraceniaceae of South America</i>

Sarraceniaceae of South America is a monograph on the pitcher plants of the genus Heliamphora by Stewart McPherson, Andreas Wistuba, Andreas Fleischmann, and Joachim Nerz. It was published in September 2011 by Redfern Natural History Productions and covered all species known at the time.

References

  1. 1 2 3 4 5 6 7 8 9 10 11 12 13 14 McPherson, S., A. Wistuba, A. Fleischmann & J. Nerz 2011. Sarraceniaceae of South America . Redfern Natural History Productions, Poole.
  2. ↑ Mellichamp, T.L. 1979. "The Correct Common Name for Heliamphora" (PDF). (196 KB) Carnivorous Plant Newsletter 8(3): 89.
  3. ↑ Liu, Sukuan; Smith, Stacey D. (2021-01-01). "Phylogeny and biogeography of South American marsh pitcher plant genus Heliamphora (Sarraceniaceae) endemic to the Guiana Highlands" (PDF). Molecular Phylogenetics and Evolution. 154: 106961. doi:10.1016/j.ympev.2020.106961. ISSN   1055-7903. PMID   32956799.
  4. ↑ S Liu, SD Smith (2023). "Replicated radiations in the South American marsh pitcher plants (Heliamphora) lead to convergent carnivorous trap morphologies". American Journal of Botany. 110 (10): e16230. doi:10.1002/ajb2.16230. PMID   37807697 – via Wiley.
  5. 1 2 3 Liu, Sukuan; Smith, Stacey D. (2021-01-01). "Phylogeny and biogeography of South American marsh pitcher plant genus Heliamphora (Sarraceniaceae) endemic to the Guiana Highlands". Molecular Phylogenetics and Evolution. 154: 106961. doi:10.1016/j.ympev.2020.106961. ISSN   1055-7903. PMID   32956799. S2CID   221844433.
  6. ↑ Macfarlane, J.M. 1908. Sarraceniaceae. In: A. Engler Das Pflanzenreich IV, 110, Heft 36: 1–91.
  7. ↑ ISBN   0-88192-356-7 Carnivorous Plants of the World a. Pietropaolo p. 72
  8. 1 2 3 Jaffe, K., Michelangeli, F., Gonzalez, J.M., Miras, B., and Ruiz, M.C. (1992). Carnivory in Pitcher Plants of the Genus Heliamphora (Sarraceniaceae). New Phytologist, 122(4): 733-744. (First page available online: JSTOR PDF of first page and HTML text of abstract
  9. ↑ Information on dates and authors of descriptions
  10. ↑ Rice, Barry A. (2006). Growing Carnivorous Plants. Timber Press: Portland, Oregon. ISBN   0-88192-807-0
  11. ↑ Beylin, Gregory (September 2023). "An Inexpensive Growth Chamber for Heliamphora". Journal of the International Carnivorous Plant Society. 52 (3): 162. doi: 10.55360/cpn523.gb111 .
  12. ↑ Author Query. International Plant Names Index.
  13. ↑ Wistuba, A., T. Carow & P. Harbarth 2002. Heliamphora chimantensis, a new species of Heliamphora (Sarraceniaceae) from the ‘Macizo de Chimanta’ in the south of Venezuela. Carnivorous Plant Newsletter 31(3): 78–82.
  14. 1 2 3 Fleischmann, A., A. Wistuba & J. Nerz. 2009. Three new species of Heliamphora (Sarraceniaceae) from the Guayana Highlands of Venezuela. Willdenowia39(2): 273–283. doi : 10.3372/wi.39.39206
  15. ↑ Nerz, J. 2004. Heliamphora elongata (Sarraceniaceae), a new species from Ilu-Tepui. Carnivorous Plant Newsletter 33(4): 111–116.
  16. ↑ Nerz, J. & A. Wistuba 2006. Heliamphora exappendiculata, a clearly distinct species with unique characteristics. Carnivorous Plant Newsletter 35(2): 43–51.
  17. ↑ Wistuba, A., P. Harbarth & T. Carow 2001. Heliamphora folliculata, a new species of Heliamphora (Sarraceniaceae) from the ‘Los Testigos’ table mountains in the south of Venezuela. Carnivorous Plant Newsletter 30(4): 120–125.
  18. ↑ (in German) Nerz, J., A. Wistuba & G. Hoogenstrijd 2006. Heliamphora glabra (Sarraceniaceae), eine eindrucksvolle Heliamphora Art aus dem westlichen Teil des Guayana Schildes. Das Taublatt 54: 58–70.
  19. ↑ Steyermark, J. 1951. Sarraceniaceae. Fieldiana, Botany28: 239–242.
  20. ↑ Nerz, J. & A. Wistuba 2000. Heliamphora hispida (Sarraceniaceae), a new species from Cerro Neblina, Brazil-Venezuela. Carnivorous Plant Newsletter 29(2): 37–41.
  21. 1 2 Maguire, B. 1978. Sarraceniaceae (Heliamphora). The Botany of the Guyana Highland Part–X, Memoirs of the New York Botanical Garden29: 36–61.
  22. 1 2 Gleason, H.A. 1931. Botanical results of the Tyler-Duida Expedition. Bulletin of the Torrey Botanical Club58(6): 367–368.
  23. ↑ Gleason, H.A. & E.P. Killip 1939. The flora of Mount Auyan-Tepui, Venezuela. Brittonia3: 141–204.
  24. ↑ Bentham, G. 1840. Heliamphora nutans. The Transactions of the Linnean Society of London18: 429–432.
  25. ↑ (in German) Wistuba, A., T. Carow, P. Harbarth, & J. Nerz 2005. "Heliamphora pulchella, eine neue mit Heliamphora minor (Sarraceniaceae) verwandte Art aus der Chimanta Region in Venezuela" (PDF). Archived from the original (PDF) on 2011-07-20. Das Taublatt 53(3): 42–50.
  26. ↑ Carow, T., A. Wistuba & P. Harbarth 2005. Heliamphora sarracenioides, a new species of Heliamphora (Sarraceniaceae) from Venezuela. Carnivorous Plant Newsletter 34(1): 4–6.
  27. ↑ (in Spanish) Fleischmann, A. & J.R. Grande Allende 2012 ['2011']. Taxonomía de Heliamphora minor Gleason (Sarraceniaceae) del Auyán-tepui, incluyendo una nueva variedad. [Taxonomy of Heliamphora minor Gleason (Sarraceniaceae) from Auyán-tepui, including a new variety.] Acta Botánica Venezuelica34(1): 1–11.
  28. ↑ "Heliamphora 'Red Mambo'".
  29. ↑ "Registered Heliamphora cultivars".
  30. ↑ Rivadavia, F. (2008). Cerro Duida, Cerro Avispa, Cerro Aracamuni Archived 2014-04-14 at archive.today . CPUK Forum, 14 June 2008.
  31. ↑ Liu, Sukuan; Smith, Stacey D. (2021). "Phylogeny and Biogeography of South American Marsh Pitcher Plant Genus Heliamphora (Sarraceniaceae) Endemic to the Guiana Highlands". Molecular Phylogenetics and Evolution. 154: 106961. doi:10.1016/j.ympev.2020.106961. PMID   32956799. S2CID   221844433 – via Elsevier Science Direct.

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