Balanophoraceae

Family Description

Distribution Map

List of Genera

Photographs

Balanophora

For a list of synonyms of Balanophora according to Hansen (1972, 1999), click HERE for the pdf file.

For a key to species of Balanophora according to Hansen (1972), click HERE for the pdf file.

1. Balanophora abbreviata
   • Inflorescence.  Benara mountain (alt 600 m), Island of Mayotte (a French territory in the Comoros archipelago, between Madagascar and Africa).  Photo by Fabien Barthelat.
   • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
2. Balanophora dioica - no photos
   • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
3. Balanophora elongata.
   • Habit of young male inflorescences. Parasitic on Pandanus klossii. Gunung Ulu Kli, Pahang, Malaysia. Photo by E. Soepadmo (Nature Malesiana, 1978, 3:24-31; name given as B. papuana).
4. Balanophora fungosa ssp. fungosa.
   • Habit of plant. Note this is a monoecious subspecies. Kingfisher Park, near Mossman, Queensland, Australia. Photograph by D. L. Nickrent [DLN 4456'].
   • Close-up of plant. Mossman Gorge, Queensland, Australia. Photograph by D. L. Nickrent [DLN 2825'].
   • SEM of pollen grain. Photo by D. L. Nickrent.
   • SEM of female flowers. Photo by D. L. Nickrent.
   • Excavated plants. Link goes to Australian National Botanic Gardens.
   • Excavated plants showing host root and tuber connection. Link goes to Australian National Botanic Gardens.
   • Inflorescence with ants. Ishigaki-island in Okinawa, Japan. Photograph by Tomoda Hitoshi, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
5. Balanophora fungosa ssp. indica var. indica
   • Young male plant with inflorescence just emerging from subtending bracts. Peninsular Malaysia. Photo by E. Soepadmo (Nature Malesiana, 1978, 3:24-31).
   • Male inflorescence, fully open. Perak, Malaysia.  Photo by W. K. Fletcher & D. M. Baylis, from Wildside’s Searchable Album.
   • Female plant in hand (listed as B. pentamera). No voucher information. Links to Missouri Botanical Gardens Tropicos Image Library.
   • Young inflorescences just emerging from the tuber and a close-up.
   • Male plants and a close-up showing the flowers fully open.
   • Female inflorescences emerging from soil.
   • Female inflorescence (left) and immature male inflorescence (right). Cebu Island, Philippines. Photo Feb. 2003 by Jeremy Holden.
   • Female inflorescences (listed as B. yakushimensis). Yakushima in Kagoshima, Japan. Photograph by Murata Hitoshi, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
   • One view and a second of the parasite growing on Quercus sp. in high altitude peat forrest near the peak of Doi Inthanon. Doi Inthanon National Park, Prov. Chiang Mai, northern Thailand, ca. 2400 m alt. Photo September 2005 by A. S. Fleischmann.
   • Tuber with host root washed out from soil after heavy rainfall.  Doi Mon Khom Long, Chiang Mai, northern Thailand, ca. 1400 m alt. Photo September 2005 by A. S. Fleischmann.
6. Balanophora fungosa ssp. indica var. minor
   • Male and female inflorescences, southern western Ghats of southern India. Photo by Rengaian Ganesan (ATREE, Bangalore, India).
7. Balanophora harlandii
   • Male inflorescences. Victoria Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
   • Young female inflorescences. Sichuan Province, China, near Yangjuan (Yanyuan?), N 27 41.584, E 101 25.764. Photo Oct. 1, 2002 by Elizabeth A. Kristy.
   • Older female inflorescence, excavated showing tuber attachment to host root. Sichuan Province, China, near Yangjuan (Yanyuan?), N 27 41.584, E 101 25.764. Photo Oct. 1, 2002 by Elizabeth A. Kristy.
   • Two male and one female inflorescence arising from tubers. Sichuan Province, China, near Yangjuan (Yanyuan?), N 27 41.584, E 101 25.764. Photo Oct. 1, 2002 by Elizabeth A. Kristy.
   • Female inflorescences. Taiwan. Photograph by C. I. Peng.
   • Female inflorescence. In a primary upper montane, closed canopy, tropical monsoon, evergreen broad-leaf forest. Altitude ca. 2,300 m in the Hoang Lien mountain range, the highest in NW Vietnam. Photo by Jeremy Holden and Fauna & Flora International (FFI).
   • Close-up of female inflorescence. In a primary upper montane, closed canopy, tropical monsoon, evergreen broad-leaf forest. Altitude ca. 2,300 m in the Hoang Lien mountain range, the highest in NW Vietnam. Photo by Jeremy Holden and FFI.
   • Close-up of male inflorescence. In a primary upper montane, closed canopy, tropical monsoon, evergreen broad-leaf forest. Altitude ca. 2,300 m in the Hoang Lien mountain range, the highest in NW Vietnam. Photo by Jeremy Holden and FFI.
8. Balanophora involucrata
   • Habit of plants. From Lijiang, Yunnan, China. Photo by Thomas Schoepke (see his Plant Image Gallery HERE)
   • Close-up of monoecious plants showing inflorescences. From Lijiang, Yunnan, China. Photo by Thomas Schoepke
   • Excavated plants showing the connate, verticillate leaves in one whorl at middle of stem. From Lijiang, Yunnan, China. Photo by Thomas Schoepke.
   • A group of inflorescences. Gaoligongshan Mt. range in Yunnan Province, China. Plant seen by team from California Academy of Science, China Natural History Project expedition. Photo by Dong Lin.
   • Two inflorescences. Tibet (Gyala Peri region, north of Namcha Barwa). Photo taken in 1994 by Walter Obermayer.
   • Habit of several plants, excavated. Yunnan Province, Yangbi: W side of Diancang Shan mountain range. Malutang in the vicinity of Chang Shan. Elevation 2700 m. 1984 Sino-Amer. Bot. Exped. (B. Bartholomew, D. E. Boufford, H. W. Li, C. G. Ma, D. H. Nicolson, T. S. Ying, & S. W. Yu) specimen no. 510. Photographed 25 June 1984 by David Boufford. [as B. fargasii]
   • Same as above, showing close-up of inflorescences. For more on the Biodiversity of the Hengduan Mountains Region, China, click HERE. [as B. fargasii]
9. Balanophora japonica
   • Inflorescences. From Hiroshima, Japan. Local name Tsuchi-torimochi. Photo by Kazuo Yamasaki (see his image gallery HERE).
   • Group of inflorescences. Kitagawa village, Akigun in Kochi, Japan. Photograph by Nagata Hideo, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
10. Balanophora latisepala
    • Population of male and female plants.  Thailand.  Photo January 1999 by W. Meijer.
    • Close-up of male inflorescence. Same loc. Photo by W. Meijer.
    • Young and old female inflorescences (foreground) and male inflorescences (background). Same loc. Photo by W. Meijer.
    • Male and female plants. Common on Phyllanthus emblica (Euphorbiaceae) in high elevation forrest on Doi Mon Khom Long Mountain, Prov. Chiang Mai, Northern Thailand, alt. 1400 m. Photo October 2005 by Andreas Fleischmann.
    • Female inflorescence. Location same as above. Photo by Stefan Kattari.
    • Male inflorescence. Location same as above. Photo by Stefan Kattari.
    • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
11. Balanophora laxiflora. Hansen has not published on the recently described taxon B. hongkongensis which occurs only in an extremely restricted area in Hong Kong. The plant was named by K. M. Lau, N. H. Li, and S. Y. Hu. [2003. Harvard Papers in Botany, 7 (2): 439-441]. These authors note that this taxon is very closely related to B. laxiflora and B. spicata. Note that Hansen considers B. spicata a synonym of B. laxiflora. The authors cite a few morphological characters that supposedly distinguish these species: the number of floral segments in the perianth and the presence of raised granules on the haustorial tuber. Balanophora honkongensis is described as having 4-9 segments, B. laxiflora 4-6, and B. spicata 6. Hansen clearly discussed the polymorphic nature of this character stating that B. laxiflora is 4-5-merous or 7-12-merous especially towards the proximal and distal parts of the inflorescence. Thus, the perianth segment ranges for both B. spicata and B. hongkongensis are clearly within the total range of B. laxiflora. With regard to surface features on the tuber, Hansen (1972) states: "I am of the opinion that far too much importance has been attached to the systematic value of the presence or absence of stellate warts and to the form of the surface cells of the tubers, as well as to the form of the actual tubers." Given that Lau et al. (2003) did not cite Hansen's 1972 monograph of Balanophora, an extremely serious omission, it is not clear whether they were unaware of it or chose to ignore it. I tend to agree with Hansen (1999) who stated "My purpose is, as it always was, to aim at a stable and lasting species delimitation. To reach this goal I have always tried to see how few taxa I could recognize, not how many."
    • Habit of male and female plants in this dioecious species. Photo by J.-M. Hu, Taiwan. [as B. spicata]
    • Close-up of male inflorescence. Photo by J.-M. Hu, Taiwan. [as B. spicata]
    • Habit of a young male inflorescence, red color form, flowers still in bud. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar. [all below as B. hongkongensis]
    • Habit of male inflorescences, red color form, with fully opened flowers. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Male inflorescences, orange color form, with flowers begining to open at the base. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Male flower, close-up, showing the synandrium. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Female inflorescences, orange color form. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Female inflorescences, red color form. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Bracts on male inflorescence referred to by Hansen as "short, blunt bodies" (left). Figure on right shows an apparent extrafloral nectary associated with the male flower buds. Castle Peak, Hong Kong. Photo taken November 2004 by Shek-Shing Mar.
    • Male inflorescences. Hoang Lien Mountains, Vietnam (far north on the Chinese border in Lao Cai province). Photo December 2004 by Jeremy Holden and FFI.
12. Balanophora lowii
    • Young inflorescence, with bracts still unfolded. Trus Madi Range, Sabah, Borneo, Malaysia. Photo by Troy Davis.
    • Inflorescence, opening to show the staminate flowers. Trus Madi Range, Sabah, Borneo, Malaysia. Photo by Troy Davis.
    • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
13. Balanophora papuana
    • Excavated male and female plants. Sirunke near Wabag, Western Highlands District, Papua New Guinea. Photo taken June 1962 (slide no. 1357), archived at Lae Herbarium.
    • Male and female plants with ruler. Sirunke near Wabag, Western Highlands District, Papua New Guinea. Photo taken June 1962 (slide no. 1357), archived at Lae Herbarium.
    • Female plant and closeup; male plant close-up. Sirunke near Wabag, Western Highlands District, Papua New Guinea. Photos taken by R. A. Howard, June 1962, archived at Lae Herbarium, PNG (slide no. 1357).
    • Male plants. Link goes to the Vascular Plant Families web page of Gerald Carr (Univ. Hawaii). Photo by Tim Motley.
    • Male plants, in full flower. Kalinga, Philippines. Photo May 2003 by Julie Barcelona.
    • Male plants, partially excavated. Kalinga, Philippines. Photo May 2003 by Julie Barcelona.
    • Female plant. Northern Sumatra, near Sibolga. Photo August 2003 by Jeremy Holden and Fauna and Flora International (FFI).
    • Male plant, and closer view showing flower buds. Northern Sumatra, near Sibolga. Photo August 2003 by Jeremy Holden and FFI.
    • Young male plant with inflorescence just emerging from the large subtending bracts. Kerinci mountains, Sumatra, Indonesia. Photo December 2004 by Jeremy Holden and FFI.
14. Balanophora polyandra - no photos
    • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
15. Balanophora reflexa
    • Close-up of male plant. Borneo: Sarawak, Gunong Mulu. Photo by J. P. Lewis 5 Oct, 1977, archived at Kew. [J. P. Lewis 280]
    • Close-up of male flower. Borneo: Sabah, Ranan District, Crocker Range. Photo by J. Beaman 29 Dec, 1983, archived at Kew. [Beaman et al. 8114]
    • Close-up male flower in bud. Borneo: Sabah, Ranan District, Crocker Range. Photo by J. Beaman 29 Dec, 1983, archived at Kew. [Beaman et al. 8114].
    • Habit of plant. Borneo: Sabah, Ranan District, Crocker Range. Photo by J. Beaman 29 Dec, 1983, archived at Kew. [Beaman et al. 8114]
    • Close-up of male flower. Borneo: Sabah, Ranan District. Photo by J. Beaman, 14 April, 1984, archived at Kew. [Beaman et al. 9378].
    • Several tubers which have not yet produced inflorescences.  Note the leafy liverworts colonizing the tubers and the prominent stellate warts.  Hansen (1972) does not indicate this species has such warts, so this could be another species of Balanophora. Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
16. Balanophora wilderi - no photos
    • Illustration from Hansen (1972) The genus Balanophora, a taxonomic monograph. Dansk Botanisk Arkiv, Vol. 28 (1): 1-188.
17. Balanophora wrightii
    • Inflorescences of plant from Japan. Name given as B. tobiracola Makino, placed in synonomy by Hansen (1972). From web pages of Eizi Suzuki, Kagoshima Unversity, Japan.
    • Close-up of young inflorescence showing open male flowers. Kaimondake, in Kagoshima, Japan. Photograph by Kawakubo Nobumitsu, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
    • Inflorescences.  Name given as B. tobiracola. Photo taken 2 November 1993, Nagasaki prefecture Nagasaki city. From "Praising Flowers" website, Japan. 
18. Balanophora yuwanensis
    • Inflorescences. Kagoshima, Japan. Local name Yuwan-tuchi-torimochi. Photo by Kazuo Yamasaki.
    • Another image of this species.

Chlamydophytum - no photos

1. Chlamydophytum aphyllum
   • Illustration from Harms (1935). Legend (translated from German): A. male inflorescence. B. Female flower, C longitudinal section. D. Group of two flowers, E cut through. F group of three flowers, G cut through. H tepals with anthers. J anthers longitudinal section, K in cross section. L female inflorescence longitudinal section, with the covering still enclosed. M part of the female inflorescence. N surface of a small female capitulum, O such in longitudinal section.
   • Illustration from Hallé (1978). Legend (translated from French): Plate 1. 1, 2, young male spike at the beginning of the opening of the spathe, 30 X 12.5 X 9 cm; 3, male stipe near to anthesis, 23 X 13 cm; 4, detail of the epidermis of the rachis of the male inflorescence, lenticels of 0.2 mm high; 5 paired male flowers close to anthesis, face and profile. (Sita 3819).
   • Illustration from Hallé (1978). Legend (translated from French): Plate 2. 1, opened up male spike, inflorescence 15 cm broad; 2, details of lenticel of inflorescence axis; 3, detail of bract; 4, detail of floriferous branch; 5, 6, opened up male flowers, profile and face, diam. 15 mm; 7, diagram of male flower having 2 supernumerary stamens; 8, longitudinal section of 2 flowers, 12 X 12 mm; 9, detail of the insertion of the anther at the base of a lobe. (Sita 3819).
   • Illustration from Hallé (1978). Legend (translated from French): Plate 3. 1, opened up female inflorescence, total width 17 cm with all the ramifications; 2, lenticel of the rachis 0.3 mm high; 3, branch of inflorescence; 4, cut of multiflowered female capitulum, diam. 12 mm; 5, stylar channel and ovule in longitudinal section; 6, stigma; 7, detail of a trilobed stigma, diam. 1.3 mm. (Sita 3819).
   • Illustration from Hallé (1978). Legend (translated from French): Plate 6. 1, 2, old desiccated and parasitized male spike, height 20-30 cm (Sita 3679); 3, provision of female flowers on a capitulum, d = dextral parastichies, s = sinistral parastichies (Sita 3819); 4, Cochineal Stictococcus, env. 3.5 X 3 mm; 5, Acarien Thiroglyphe 0.6 mm; 6, Acarien Oribate (a mite), 0.2 mm; 7, dipteran larva Orthorrhaphe, 7 mm; 8, id., above the brain of 0.9 mm in length.
   • Images 1, 2 and 3 of this remarkable species, photographed (possibly for the first time!) November 2007 in west Africa (Gabon?) by Alexander Georgiev.  The local people believe (erroneously) that this plant is a mushroom and that it is poisonous to touch. The description of this species in Hansen's key to Balanophoraceae of Continental Africa says that there are 6-7 stamens, but this specimen has fewer.

Corynaea

1. Corynaea crassa
   • Photo. Habit of plants with young inflorescences and black peltate scales still enclosing the inflorescence. Mauseth no. 7790. August 1980. Photo by J. Mauseth.
   • Photo Excavated plant, with peltate scales still enclosing the inflorescence. Volcan Barva, Costa Rica. Specimen Nickrent3011. Photograph by D. L. Nickrent.
   • Photo. Habit of plants with black peltate scales still enclosing the inflorescence. Cuericí Biological Station, Costa Rica. Photo by Joel McNeil.
   • Photo. Inflorescence after loosing peltate scales, emerging from the haustorial tuber. Note the ring-like structure (volva) formed midway on the inflorescence axis. Younger inflorescences are emerging from the tuber below. Cuericí Biological Station, Costa Rica. Photo by Joel McNeil.
   • Photo. Left inflorescence, without peltate scales, emerging from the large tuberous haustorium. Cloud forest habitat (ca. 10,000 ft.) in the Talamancas, Cuericí Biological Station, Costa Rica. Photo by Kyle Williams.
   • Photo. Same plant as above, close-up, showing lower portion of inflorescence without peltate bracts, and styles of female flowers emerging from the mass of filiform clavate hairs. Cuericí Biological Station, Costa Rica. Photo by Kyle Williams.
   • Photo. Close-up of inflorescence surface after loosing peltate scales showing the mass of filiform clavate hairs (= articulated filaments of Hooker). Cuericí Biological Station, Costa Rica. Photo by Joel McNeil.
   • Photo. Inflorescence with male flowers and close-up. Unisexual inflorescences are found in about 30% of the plants. Male flowers have a tubular perianth and three stamens whose filaments are united into a column. Cuericí Biological Station, Costa Rica. Photo by Joel McNeil.
   • Photo. Excavated plants. Costa Rica. Photo by Barry Hammel. Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Photo. Excavated plants. Costa Rica. Photo by Barry Hammel. Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Photo. Close-up of inflorescence, peltate scales dehisced, inflorescence beginning to deteriorate, possibly with young fruits on the surface. Costa Rica. Photo by Barry Hammel. Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Illustration. This is C. crassa var. sprucei, which is distinguished from the typical variety by its 3-lobed (vs. unlobed) perianth. Flowers not shown. Illustration by Roland Eberwein.

Dactylanthus

1. Dactylanthus taylori
   • Photo A monotypic genus endemic to New Zealand. These three inflorescences, showing hundreds of small male flowers laden with white pollen, will each have a pool of up to 1.5 ml of nectar inside the protecting bracts. Photo from Ecroyd (1995, Nature Conservation 4: The Role of Networks, Surrey Beatty & Sons. Used with permission).
   • Photo. Shown is a short-tailed bat (Mystacina tuberculata) feeding on the nectar from a male inflorescence of the root parasite. Bats appear to be the primary pollinators of Dactylanthus. Photo by Eric Anderson and Chris Ecroyd.
   • Photo. Germinating seed with long hairs growing from the 'radicle.' Photo by B. O'Brien and S. Holzapfel. From Ecroyd (1996, New Zealand Journal of Ecology, 20:81-100).
   • Photo. Scanning electron micrograph of cellular structure of seedling. Note intracellular fungal hypha (indcated by arrows. Photo from Ecroyd (1996, New Zealand Journal of Ecology, 20:81-100).
   • Photo. Woodrose formed by the host root in response to the root parasite. Photo from Ecroyd (1995).
   • Dactylanthus taylorii is under threat by several mammals that feed on the inflorescences. The introduced Australian brush-tailed possum (Trichosurus vulpecula), shown in the upper picture, is attempting to get through a netting exclosure. The lower picture shows the ship rat (Rattus norvegicus) is responsible for some pollination but also destroys flowers.
   • Plate 1 of four images from the New Zealand Plant Conservation Network. A. young tuber (Avi Holzapfel); B. Tuber partly exposed and inflorescence (Avi Holzapfel); C. group of inflorescences (Chris Ecroyd); D. inflorescence (NZ Department of Conservation).
   • Plate 2 of four images from the New Zealand Plant Conservation Network. A. female plant with inflorescences (Nick Singers); B. plant with lighter (yellow) inflorescence bracts (Nick Singers); C. inflorescence in hand (Avi Holzapfel); D. infructescence (Avi Holzapfel).

Ditepalanthus

1. Ditepalanthus malagasicus
   • Photo. Madagascar: Hiaraka. Photo by Bogner 28 Jan., 1969 [Bogner 269]. Slide no. 6477 archived at Kew (as Rhopalocnemis malagasica).
   • Illustration. By Roland Eberwein

Exorhopala

1. Exorhopala ruficeps
   • Photo. Inflorescence emerging from soil. Peninsular Malaysia, Penang, Penang Hill (Bukit Penara), 14. Aug., 1986, Photo by Anton Weber.
   • Photo. Two older inflorescences. Peninsular Malaysia, Penang, Penang Hill (Bukit Penara), 14. Aug., 1986, Photo by Anton Weber.
   • Photo. Excavated plant showing haustorial connection. Peninsular Malaysia, Penang, Penang Hill (Bukit Penara), 14. Aug., 1986, Photo by Anton Weber.

Hachettea

1. Hachettea austro-caledonica
   • Photo. Male inflorescence. Link goes to the Vascular Plant Families web page of Gerald Carr (Univ. Hawaii). Photo by Tim Motley.
   • Photo. Male flowers, close-up. Link goes to the Vascular Plant Families web page of Gerald Carr (Univ. Hawaii). Photo by Tim Motley.
   • Photo. Young inflorescence emerging from the soil, not yet open. Mt. Ouin, New Caledonia. Photo by Sarah Mathews.
   • Photo.  Young inflorescences just beginning to open.  New Caledonia.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)

Helosis

1. Helosis cayennensis
   • Photo. Beautiful photograph of several inflorescences from Peru. Photographer unknown. From "Parasites, looking for a free lunch" by Jennifer Ackerman, National Geographic Magazine, October 1997, Vol. 192, No. 4, p. 80.
   • Photo. Immature inflorescence. Peru, Loreto: Maynas Iquitos. Photo by Mac Alford, 2002.
   • Photo. Young inflorescence, with peltate scales just starting to separate. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. Showing flowers emerging from the dehiscing peltate scales (inflorescence on the right). Bolivia, Department La Paz, Province Inquisivi, mountain rainforest along road Inquisivi-Circuata, 2 km beyond Polea. Photo by Marie-Stéphanie Samain, 13 January 2008.
   • Photo. Excavated plant showing subterranean rhizomes. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. Close-up of rhizomes. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. Male flower buds emerging from the mass of clavate hairs on the inflorescence. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. Inflorescence with peltate scales already dehisced and the male flower buds emerging from the mass of clavate hairs. Peru, Loreto: Maynas Iquitos. Photo by Mac Alford, 2002.
   • Photo. Habit of plant showing male flower dehiscence. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. Another plant showing male flower dehiscence. Wesern Ecuador. Photo by Nancy Garwood.
   • Male flowers and close-up of a single flower. Note the three tepals, the fused staminal column, the free filaments at the top that bear the anthers which are coherent into a synandrium. Wesern Ecuador. Photo by Nancy Garwood.
   • Photo. A beautiful photograph showing close-ups of flowers. Saül, French Guiana. Photograph by C. Gracie.
   • Photo. Habit of emerging inflorescences. La Milpa, Belize. Photo by Nancy Garwood.
   • Photo. Young inflorescences excavated. La Milpa, Belize. Photo by Nancy Garwood.
   • Photo. Female inflorescences. La Milpa, Belize. Photo by Nancy Garwood.
   • Photo. Female inflorescences showing the silky styles. Equitos, Peru. Photo by G. Amico.
   • Photo . Inflorescences. Photo by A. Gentry (voucher no. 14018). Links to Missouri Botanical Gardens Tropicos Image Library.
   • Photo . Inflorescences in situ. Photo by A. Gentry (voucher no. 14018). Links to Missouri Botanical Gardens Tropicos Image Library.
   • Photo. This one is considered H. cayennensis var. mexicana by Hansen. Veracruz, Mexico. Photograph by Dave Lorence.
   • Illustration. H. cayennensis var. cayennensis by Roland Eberwein
   • Illustration. H. cayennensis var. mexicana by Roland Eberwein.

Langsdorffia

1. Langsdorffia hypogaea
   • Photo. Immature inflorescences just emerging from the soil. Las Alturas, Costa Rica. Photograph by Shu-Chuan Hsiao.
   • Photo. Immature inflorescences. Monteverde, Costa Rica. Photo by Kevin Nixon 1991. Archived at PlantSystematics.org.
   • Photo. Somewhat older inflorescences, with male flowers emerging from the subtending bracts. Monteverde, Costa Rica. Photo by Kevin Nixon 1991.
   • Photo. Top view of inflorescences. Note greenish color on male flowers! Peru. Photograph by Alwyn Gentry, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
   • Photo. Intact male and female inflorescences. 6 km west of El Junquito, Venezuela. Nee and Whalen (voucher no.16864). Photo July 26, 1979 by M. Nee.
   • Photo Female (above) and male (below) inflorescences both sectioned longitudinally. 6 km west of El Junquito, Venezuela. Nee and Whalen (voucher no.16864). Photo July 26, 1979 by M. Nee.
   • Photo. Close-up of male inflorescence, sectioned longitudinally. 6 km west of El Junquito, Venezuela. Nee and Whalen (voucher no.16864). Photo July 26, 1979 by M. Nee.
   • Photo. Male inflorescence. Teresópolis, Rio de Janeiro, Brazil. Photo by Leandro Cardoso.
   • Photo. Male flower. From fixed material. Teresópolis, Rio de Janeiro, Brazil. Photo by Leandro Cardoso.
   • Photo. Conical bodies (derived from bracts) that subtend the male flowers. From fixed material. Teresópolis, Rio de Janeiro, Brazil. Photo by Leandro Cardoso.
   • Photo. Fruits from herbarium specimen by B.M.T. Walter et al. (1973). Uruaçu, Goiás, Brazil. Photo by Leandro Cardoso.
   • Photo. Nine female inflorescences and pruners. Central Brazilian plateau. Photo by David Hunt, 17 Dec. 1966 [D. R. Hunt 6027]. Slide no. 11416 archived at Kew.
   • Photo. Infructescence. Pirre. June 30, 1980. Photo by J. Mauseth
   • Photo. Female inflorescences and infructescences. Photograph by Al Gentry (voucher 14019). Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Photo. Female inflorescences. Photograph by Al Gentry (voucher 14019). Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Photo. Male inflorescences - bright red!. Photograph by Al Gentry (voucher 53282). Links to Missouri Botanical Gardens TROPICOS Image Library.
   • Photo. Male (above) and female (below) inflorescences. Photograph by Al Gentry (voucher 74997). Links to Missouri Botanical Gardens TROPICOS Image Library.
2. Langsdorffia papuana
   • Photo. Young inflorescence emerging from soil. Yamap, Bulolo Valley, Papua New Guinea. Photo taken May 1970, archived at Lae Herbarium.
   • Photo. Older inflorescence and rhizome. Yamap, Bulolo Valley, Papua New Guinea. Photo taken May 1970, archived at Lae Herbarium.

Lathrophytum

1. Lathrophytum peckoltii
   • Photo1 and Photo2. Plants attached to Paullinia sp. (Sapindaceae). Parque Estadual da Pedra Branca, Rio de Janeiro, Brazil. Photo by Leandro Cardoso [Cardoso 261].
   • Photo. Entire plant (fixed) showing haustorial tuber and monoecious condition (male flowers above, female below).
   • Photo. Composite of photographs showing fixed male and female flowers. A - close-up of male flower; B - surface view of male inflorescence showing male flowers (each with two anthers) subtended by a fleshy bract; C - female flowers arranged on several secondary axes; D - close-up of one secondary axis with female flowers. Photo by Leandro Cardoso.
   • Photo. Young inflorescence, just emerging from the soil, with peltate bracts still covering flowers. Brasil, RJ, Rio de Janeiro, Recreio dos Bandeirantes. Photo by Leandro Cardoso December 2003.
   • Photo. An older inflorescence. Brasil, RJ, Rio de Janeiro, Recreio dos Bandeirantes. Photo by Leandro Cardoso, December 2003.
   • Photo. Older inflorescence, excavated and in hand. Brasil, RJ, Rio de Janeiro, Recreio dos Bandeirantes. Photo by Leandro Cardoso, December 2003.
   • Illustration from Hansen (1980). Legend of Fig. 22. A-H, Peckolt sn 1886 after Hansen (1976), I and K (Pereira 5645) original. A, habit of young specimen, tuber lost, inflorescence covered by peltate parts of bracts X1; B, bract subtending female branch, side view X6; C, the same, front view of pelta X6; D, bract subtending male flower, from below X6; E, male flower, side view X6; F, the same, from above X6; G, female branch, side view X6; H, female flower, side view X17.5; I, habit of flowering specimen with host root, tuber, volva lobes, lower female part of inflorescence showing peltas of branches, upper male part with anthers opened, some 30 flowers have lost the anthers X1, four male flowers with anthers open, three anthers lost, seen from distal end X2.5.
   • Illustration from Hansen (1976). Legend of Figure 1: a: tuber, volva with lobes broken off, and lower part of stem, presumably old specimen (herb. M); b: stem with volva-lobes, inflorescence still covered by bracts, younger specimen (herb. C); c: longitudinal section of young specimen, inflorescence still covered by volva (herb. K); d: same, upper part enlarged, lower part of inflorescence with female-secondary axes, upper part with male-flowers; e: bract supporting male-flower, lateral view slightly from behind; f: same, from below; g: male-flower, from above; h: same, lateral view; i: bract supporting female-secondary axis, lateral view; k: same, from behind; l: same, front view; m: female-secondary axis, lateral view, flowers removed in lower part, still in position in upper part, covered outwards by the enlarged, peltate top part of branch; n: female-flower with 2 styles; o: same, longitudinal section. - Enlargement; a-c: scale 5 cm; d-m: scale 5 mm; n, o: scale 1 mm. - Material: a, b, e-o: PECKOLT s. n., leg. 1886; c, d: GLAZIOU s. n., leg. 1886.

Lophophytum

1. Lophophytum leandrii
   • Photo,. Entire plant, showing female and male inflorescences on same individual (hence monoecious). Note the dehisced pollen on the rock below. Sete Barras, state of São Paulo, Brazil. Photo 23 August, 2004 by R. Goldenberg (voucher 664 ).
   • Photo,. Close-up of male inflorescence showing male flowers. Sete Barras, state of São Paulo, Brazil. Photo 23 August, 2004 by R. Goldenberg (voucher 664 ).
   • Photo,. Close-up of male flowers. Sete Barras, state of São Paulo, Brazil. Photo 23 August, 2004 by R. Goldenberg (voucher 664 ).
   • Photo,. Base of plant showing inflorescence bracts, female inflorescences, and male inflorescences. Sete Barras, state of São Paulo, Brazil. Photo 23 August, 2004 by R. Goldenberg (voucher 664 ).
   • Photo,. Close-up of female inflorescences. Sete Barras, state of São Paulo, Brazil. Photo 23 August, 2004 by R. Goldenberg (voucher 664).
2. Lophophytum mirabile
   • Photo. Young inflorescence emerging from soil, covered with bracts  - Brasil, RJ, Angra dos Reis. Photo by Leandro Cardoso, August 2004.
   • Photo. Mature inflorescence showing the open male flowers above and the female flowers below on the spike. Brasil, Espírito Santo. Photo by Leandro Cardoso, November 2004.
   • Photo 1. Photo 2. Inflorescences. Perú. From Neotropical Live Plant Photos.   Photo IDs 12269a, 12271 by Robin Foster.
   • Illustration of L. mirabile and L. leandrii from Harms (1894). Figure 161 legend, translation from German. A-C Lophophytum mirabile Schott et Endl. A a group of young plants in different stages of the development, on a strongly swollen host root (Nw); a = very young tubers; b = somewhat older; c tuber with the beginning of the development of leaves. d = somewhat older stage of a tuber growing into an inflorescence, without cataphylls at the lower part; e = older growing rhizomes with cataphylls covering also basal parts. B longitudinal section through a young inflorescence, showing the shieldlike stalked bracts and the individual spadices of the second order. C flowering plants, in which the bracts have been shed - D part of a female spadix of L. leandri Eichl., showing the bracts of the flowers. - E-K L. mirabile. E male flower from the side and from the front, with p = pistil vestige. F pollen (240/1). G female flower with two staminodia. H ovary in longitudinal section, ov = ovule., skl = sclerenchyma group. J longitudinal section of a somewhat older ovary, showing further advanced development of the upper ovary rim, with ovules completely adhered to the ovary wall .; spt = the septum-like extension of the placenta, es = embryosac. K longitudinal section of a fruit, pt = endocarp, s = seed. (after Eichler.).
3. Lophophytum weddellii
   • Illustration from Hooker (1856). Fig. 1. Portion of section of male inflorescence. Fig. 2. Portion of section of female inflorescence.

Mystropetalon

1. Mystropetalon polemannii
   • Photo. Group of inflorescences. Betty's Bay, South Africa. Photo by L. J. Musselman
   • Photo. Close-up of inflorscence. Betty's Bay, South Africa. Photo by L. J. Musselman
   • Photo. Close-up of flower. Betty's Bay, South Africa. Photo by L. J. Musselman
   • Photo. Three photographs showing inflorescence, fruits and seeds. Link goes to Plant Web, plants of South Africa by Peter Swart.
2. Mystropetalon thomii
   • Photo Individual plant with older genets toward outside, young genet on inside of ring. The young shoot is in the female (pistillate stage). Near Kleinmond, South Africa. (voucher no. DLN 4091). Photo by D. L. Nickrent.
   • Photo Same plant as above, but excavated to show that all genets arise from the same stem. Note the new shoots being formed. A curious finding was the presence of liquid surounding the central stems. Is this produced by the parasite? What is its function? Photo by D. L. Nickrent.
   • Photo Old (left) and new (right) shoots in male (staminate) flower phase. Photo by D. L. Nickrent.
   • Photo Succession of floral stages present on one inflorescence. The shoot on the right has female flowers forming fruits below, old male flowers in the middle, and young male flowers at the top. Photo by D. L. Nickrent.
   • Photo Close-up of very young shoot forming female flowers. Note the long styles that project from the inflorescence. Photo by D. L. Nickrent.
   • Photo Female (top) and male (bottom) flowers. The ring-shaped, white elaisome that sits below the ovary can be seen enclosed within bracts of the female flower. (Note: the fruit/elaiosome is upside down). The elaiosome is harvested by ants which disperse the one-seeded fruits. The male flowers contain two rather typical stamens (as compared to other Balanophoraceae!) that are attached to the perianth (tepals). The flower is then subtended by bracts and bracteoles. Photo by D. L. Nickrent.

Ombrophytum

1. Ombrophytum peruvianum
   • Photo. Inflorescences. Perú. From Neotropical Live Plant Photos.   Photo ID 12360 by Robin Foster.
   • Photo. Plant with inflorescence. Departamento Santa Cruz, Bolivia. (voucher no. 37346 M. Nee). Photo modified from one taken by M. Nee.
2. Ombrophytum subterraneum
   • Photo. Plant parasitic on Tessaria absinthioides, Chiuchiu, Chile. Photo by James Mauseth.
   • Photo. Close-up of inflorescence emerging from volva. Parasitizing cultivated alfalfa (Medicago). Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. Inflorescence scales. Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. Young inflorescence. Chiuchiu, Chile. May 1992. Photo by J. Mauseth.
   • Photo. Whole plants in hand. On Tessaria absinthioides (Asteraceae). May 1991. Chiuchiu, Chile. Photo by J. Mauseth.
   • Photo. Excavated plant showing close-up of tuber. Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. Excavated plant showing tuber. Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. Whole plant, excavated, showing attachment to host root. Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. Locale of subterranean Ombrophytum, whose shoot are just visible protruding from the soil. Chiuchiu, Chile. August 1987. Photo by J. Mauseth.
   • Photo. SEM photograph of female flower. White bar = 1.0 mm. Photo by J. Mauseth.
   • Photo. Transverse section of vessels of the parasite. The lumen of the narrower vessels are almost occluded by the ingrowths whereas wider vessels are relatively unobstructed. See Mauseth et al. (1992). Photo by J. Mauseth.
   • Photo. A parasite cell embedded within Tessaria tuber tissue. This cell is still close to the interface (note O. subterraneum cells above) and appears healthy. See Mauseth et al. (1992). Photo by J. Mauseth.
   • Photo. Host-parasite interface. Photo by J. Mauseth.
   • Photo. SEM showing a transverse section of vessels. The secondary walls of the vessels have knobby ingrowths whose function is unknown. See Mauseth and Montenegro (1992). Photo by J. Mauseth.
3. Ombrophytum violaceum
   • Photo. Inflorescences. Perú. From Neotropical Live Plant Photos.   Photo ID 12266 by Robin Foster.
4. Ombrophytum sp.
   • Photo. Inflorescences. Photograph by Al Gentry (voucher 39680). Links to Missouri Botanical Gardens TROPICOS Image Library.

Rhopalocnemis

1. Rhopalocnemis phalloides
   • Photo. Sumatra, at 1400 m elevation. Note the inflorescence, with hexagonal scales, pushing upward through the basal volva causing it to split. Photo by Jeremy Holden and FFI.
   • Photo. Young inflorescence emerging from basal volva. In a primary upper montane, closed canopy, tropical monsoon, evergreen broad-leaf forest. Altitude ca. 2,300 m in the Hoang Lien mountain range, the highest in NW Vietnam. Photo by Jeremy Holden and FFI.
   • Photo. An older inflorescence, possibly forming fruits. In a primary upper montane, closed canopy, tropical monsoon, evergreen broad-leaf forest. Altitude ca. 2,300 m in the Hoang Lien mountain range, the highest in NW Vietnam. Photo by Jeremy Holden and FFI.

Sarcophyte

1. Sarcophyte sanguinea
   • Photo. Excavated plant showing massive, tuberous haustorium attached to host root (to the left) and three male (staminate) inflorescences. Hoole farm, near Riebeek Oos, South Africa (voucher DLN 4109). Photo by D. L. Nickrent.
   • Photo. Portion of male inflorescence showing the fleshy, 3-merous flowers with prominent white anthers. These flowers have an extremely strong and disagreable odor (like rotten meat) that attracts the pollinators (flies). Photo by D. L. Nickrent.
   • Photo. Excavated female (pistillate) plant showing the globose inflorescences composed of numerous "heads" that each contain hundreds of reduced female flowers. Photo by D. L. Nickrent.
   • Photo. Female (pistillate) inflorescence sectioned to show vasculature. Photo by D. L. Nickrent.

Scybalium

1. Scybalium depressum
   • Illustration by Roland Eberwein.
2. Scybalium fungiforme
   • Photo. Sectioned inflorescence from an herbarium specimen (E.M. Santos et al. BHCB). Photo by Leandro Cardoso.
   • Photo. Same as above, showing flowers close up. Photo by Leandro Cardoso.
   • Illustration by Roland Eberwein.
3. Scybalium glaziovii
   • Photo. Plant in situ. Reserva Ecológica de Macaé de Cima, Nova Friburgo, Brazil. Photo by M. Bocayuva & L. Cardoso
   • Photo. Excavated plant showing basal tuber. Reserva Ecológica de Macaé de Cima, RJ, Brazil. Photo by Leandro Cardoso.
   • Photo. Herbarium specimen. Brade 18909. Photo by Leandro Cardoso.
4. Scybalium jamaicense
   • Photo. Immature inflorescences, with bracts still attached, emerging from rhizomes. Hardwar Gap, Jamaica. (DLN 3021) Photograph by D. L. Nickrent.
   • Photo.  Two male inflorescences on the left and a female inflorescence on the right.  The male flowers are shown emerging as buds from a thick layer of pink hairs as the bracts are being shed.  Jamaica.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
   • Photo. Longitudinal sections of inflorescences. Photograph by Al Gentry (No. 28341). Links to Missouri Botanical Gardens Tropicos Image Library.
   • Illustration by Roland Eberwein.

Thonningia

1. Thonningia sanguinea
   • Photo. Zambia: Mwinilunga district, Miombo woodland by Chingabola Dambo. Photo by C. C. Townsend (no. SPM) 16 Feb. 1975. Slide no. 9986 archived at Kew.
   • Photo. Gabon, coastal rain forest near Iguela. Photograph August, 2000 by Marc and Peggy Faucher (South Burlington, VT).
   • Photo. Gabon, near Makokou. View of plant from top. Photo by Yves Caraglio.
   • Photo. Same as above showing plant from side. Photo by Yves Caraglio.
   • Photo 1 and photo 2 of a plant from west Africa (Gabon?) photographed by Alexander Georgiev.
   • Photo.  Plant being visited by ants.  Budongo Forest Reserve, western Uganda.  Photo by Concy Olanya Acen, Makerere University Kampala, Faculty of Forestry and Nature Conservation.
   • Photo. Gabon, Haut-Ogooué. Voucher (No. 961) and photograph by Walters, Bradley, Essouma & Mbaniboua. Links to Missouri Botanical Gardens TROPICOS Image Library.

Phylogeny

1. Placement of Balanophoraceae among Flowering Plants

Determining the closest photosynthetic relatives of Balanophoracease has long intrigued botanists. Recent molecular phylogenetic analyses point toward Santalales, the sandalwood order. See paper by Nickrent et al. (2005) in BMC Evolutionary Biology. For a PDF file of this article, click HERE. For the full text online version, go HERE.

2. Infrafamilial Phylogeny of Balanophoraceae

Click HERE to see a tree generated using nuclear small-subunit (18S) rDNA sequences from 11 of the possible 17 genera of Balanophoraceae, with Amborella included as an outgroup. Bear in mind that this analysis was done before the closest photosynthetic relatives of Balanophoraceae were known (see BMC paper above). Thus, one should not interpret this tree as supporting a close relationship between Balanophoraceae and the most primitive flowering plant Amborella.

This tree shows a strong biogeographical signal, with the Neotropical taxa forming one clade that is derived from within the Paleotropical grade. The association of Dactylanthus and Hachettea follows traditional classifications (e.g. Dactylanthaceae sensu Takhtajan 1997), however, the additional association with Mystropetalon (Mystropetalaceae sensu Takhtajan 1997) is surprising. Despite the wide geographic separation (S. Africa, New Zealand, and New Caledonia), these data suggest an ancient southern hemisphere association that traces to ancestors on the Gondwanan landmass.

References