"Santalaceae" s. lat.

Family Description

Distribution Map

List of Genera

Photographs

Molecular phylogenetic work by Der and Nickrent (2008) highlighted the existence of several clades in what is traditionally called "Santalaceae".  The following list of genera is arranged according to these clades. The genus Anthobolus has been moved to Opiliaceae and Arjona and Quinchamalium to Schoepfiaceae.

Comandra Clade

Comandra

1. Comandra umbellata

• Flowering shoot. Virginia, USA. Photo by L. J. Musselman.
• Close-up of flowers. Virginia, USA. Photo by L. J. Musselman.
• Sectioned flower showing "receptacular" inferior ovary -- an infrequent type in angiosperms. Champaign Co., Illinois, USA. Photo by Ken Robertson.
• Flower close-up. Note 5-merous and 6-merous flowers. Photo taken May 27, 1996 in Butte Co., CA by Arthur H. Bazell (from Cal Photo website).
• Close-up of inflorescence. Albany Pine Bush, New York. Photo by Carol Gracie.

Geocaulon

1. Geocaulon lividum
   • Habit of the plant. Banff National Park, Alberta, Canada. July 1985. Photo by Job Kuijt.
   • Habit of plant with flowers, Alaska. Photo by L. J. Musselman.
   • Plant with flowers. Note the flower color changes during development. Alaska. Photo by L. J. Musselman.
   • Close-up of flowers, Alaska. Photo by L. J. Musselman.

Thesium Clade

Buckleya

1. Buckleya distichophylla
   • Shoot with staminate flowers. Buckleya is a rare, Tertiary relictual shrub of the Appalachians that is parasitic on Tsuga (hemlock trees). [DLN 2692]. Virginia, USA. Photo by D. L. Nickrent.
   • Staminate flowering shoots. Virginia, USA. Photo by L. J. Musselman.
   • Staminate flowers, close-up.Virginia, USA Photo by Ken Robertson.
   • Carpellate flower.Virginia, USA. Photo by L. J. Musselman.
   • Carpellate plants showing "torpedo-like" fruits. Virginia, USA. Photo by L. J. Musselman.
2. Buckleya lanceolata
   • Plant showing carpellate flower. Japan. Photo by Kazuo Yamasaki.
   • Fruits. Yamanashi, Japan. Photograph by Kihara Hiroshi, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).
   • Photos of the male flowers and fruits. Link goes to Y. Hada's pages, Okayama University of Science.

Kunkeliella

1. Kunkeliella canariensis
   • Habit of plant. Gran Canaria, Canary Islands. Photo by Isabel Santana López, November 2003.
   • Closer view of plant showing developing fruits. Gran Canaria, Canary Islands. Photo by Isabel Santana López, November 2003.
   • Close-up of flower. Gran Canaria, Canary Islands. Photo by Isabel Santana López, November 2003.
   • Illustration from Stearn, W. T. 1972 (Cuad. Bot. Canar. XVI: 11-26).
2. Kunkeliella subsucculenta
   • Habit of plants in vegetative condition. Tenerife, Canary Islands. Photographed in September 2003 by José Naranjo Suárez.
   • Same as above showing close-up of succulent stems. Tenerife, Canary Islands. Photographed in September 2003 by José Naranjo Suárez.
3. Kunkeliella retamoides
   • Habit of plant in fruit. Canary Islands. Photo by Peter Schönfelder. See Flora Canaria web site.
   • Close-up showing flowers. Canary Islands. Photo by Peter Schönfelder.

Osyridocarpos

1. Osyridocarpos schimperianus
   • Shoots. Near Grahamstown, South Africa. [DLN 4110]. Photo by D. L. Nickrent.
   • Flowering and fruiting stems, close-up. Near Grahamstown, South Africa. [DLN 4110]. Photo by D. L. Nickrent.
   • Flowering shoot.  Vumba Botanic Garden, Zimbabwe. From Flora of Zimbabwe web site.  Photo by B. Wursten.
   • Shoot with young fruits.  Ndundu Lodge, Vumba.  Zimbabwe.  From From Flora of Zimbabwe web site.  Photo by B. Wursten.

Thesidium

1. Thesidium fragile
   • Habitat of the plant. De Hoop National Park, South Africa. [DLN 4102]. Photo by D. L. Nickrent.
   • Closer view of the plant. Same as above. Photo by D. L. Nickrent.
   • Inflorescence. Same as above. Photo by D. L. Nickrent.
   • Close-up of plant showing squamate habit (i.e. with scale leaves), the tiny flowers arranged in compact spikes, and a developing fruit with a white inferior ovary and persistent orange sepals. Same as above. Photo by D. L. Nickrent.

Thesium

1. Four (of the many!) species of Thesium from South Africa. All obtained in general vicinity of Capetown, South Africa. Photo by D. L. Nickrent.From left to right:
   • T. spicatum [DLN 4095]
   • T. strictum [DLN 4093]
   • T. carinatum [DLN4 4094]
   • T. nudicaule [DLN 4098]
2. Thesium alpinum
   • Inflorescence and close-up of flowers. Note that the flowers are arranged in a one-sided raceme, which is characteristic for this species. This species normally bears 4-merous flowers, 5-petaled ones occur only occasionally. Elev. about 1600 m, Brauneck, Bavaria, Germany. Photo taken 21st June 2005 by Andreas S. Fleischmann.
   • Close-up of flowers. Hochschwab, Austria. Photo by G. Glatzel.
3. Thesium arvense (= T. ramosum)
   • Shoots. A very widespread species in Asia and Eastern Europe. Note the white, swollen fruiting pedicels. The plant is also quite scabrid, specially on the inflorescence. Photographed near Pambak, Sevan Lake, Armenia, by Miguel A. García.
4. Thesium bavarum
   • Habit of plant. Growing up to 60 cm in height, this is one of the tallest European Thesium species. Landsberg, Bavaria, altitude 580 m. Photo June 2006 by A. S. Fleischmann.
   • Close-up of flowers. Landsberg, Bavaria, altitude 580 m. Photo June 2006 by A. S. Fleischmann.
5. Thesium bergeri
   • Flower close-up. Fayraya. Lebanon. Photographed April 18, 2002 by L. J. Musselman.
6. Thesium brachyphyllum
   • Flowering shoots. Photographed at Meghri Pass, Armenia at 2550 m. not far from the Iranian border, by Miguel A. García.
7. Thesium carinatum
   • Close-up of inflorescence. Bain's Kloof, just above Montagu Rocks, South Africa [DLN 4094]. Photo by D. L. Nickrent.
8. Thesium chinense
   • Close-up of flower. Y. Hada's pages, Okayama University of Science. Link to more photos of this species HERE.
   • Fruiting plant. Photo by J.-M. Hu, May 25, 1993. Hua-Lien, Taiwan. [Hu 93-1211].
9. Thesium funale
   • Flowering shoots. Link goes to Plant Web, plants of South Africa by Peter Swart.
10. Thesium humifusum
    • Several photographs of flowers, fruits and pollen. From BIOIMAGES: The Virtual Field-Guide (UK).
11. Thesium lineatum
    • Habit of the plant. This shrub stands out among the shorter vegetation at this site south of Southerland, South Africa. Photo by D. L. Nickrent.
    • Closer view of the shoots showing flowers and developing fruits. Photo by D. L. Nickrent.
12. Thesium nudicaule
    • Close-up of flowers and fruits. Goedverwag, South Africa [DLN 4098]. Photo by D. L. Nickrent.
13. Thesium pyrenaicum
    • Plant in flower. Landsberg, Bavaria, Germany. Photo June 2004 by Andreas S. Fleischmann.
    • Plant in fruit. Landsberg, Bavaria, Germany. Photo June 2004 by Andreas S. Fleischmann.
14. Thesium rostratum
    
    • Habit of plant. Starnberg, Bavaria. Photo June 2006 by A. S. Fleischmann.
    • Close-up of flowers. Starnberg, Bavaria. Photo June 2006 by A. S. Fleischmann.
    • Fruits. This species is easily recognized by its conspicious yellow, berry-like fruits that are topped by a persistent green petals and the presence of one (vs. three) floral bract. Starnberg, Bavaria. Photo June 2006 by A. S. Fleischmann.
15. Thesium spicatum
    • Habit of plant. Boskloof Farm, South Africa. [DLN 4095] Photo by D. Nickrent.
    • Inflorescences. Boskloof Farm, South Africa. Photo by D. Nickrent.
    • Close-up of inflorescence. Boskloof Farm, South Africa. Photo by D. Nickrent.
16. Thesium strictum
    • The plant is a common shrub just over one meter in height. Kirstenbosch Botanical Garden, Capetown, South Africa. [DLN 4093]. Photo by D. L. Nickrent.
17. Thesium szovitsii
    • Photos of flowers and fruits. This plant grows in subdesertic places near Yerevan and close to the Turkish and Azerbaijan borders. It is a glaucous, erect plant with relatively large flowers and in many cases is somewhat succulent.Photo by Miguel A. García.
18. Thesium triste
    
    • Habit of plant.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
19. Thesium viridifolium
    • Nine beautiful photographs showing flowers, etc. Link goes to Plant Web, plants of South Africa by Peter Swart.

Cervantesia Clade

Acanthosyris

1. Acanthosyris annonagustata
   • Habit of plant in fruit. Eastern Ecuador. Web page at Missouri Botanical Garden where Ulloa and Jørgensen describe this new taxon. Photo by V. Persson, N. Garwood & G. Villa.
   • Fruit cut lengthwise. Fruit and nut. Photo by V. Persson, N. Garwood & G. Villa.
2. Acanthosyris asipapote
   • Flowers from the type specimen (Vargas 3234). Santa Cruz, Bolivia. Photo by M. Nee.
   • Leaves and fruits. Santa Cruz, Bolivia. Photo by M. Nee.
   • Sectioned fruits. Santa Cruz, Bolivia. Photo by M. Nee.
3. Acanthosyris falcata
   • Plant with flowers. Bolivia. Photo by A. Gentry (No. 75283). Link goes to TROPICOS database, MO.
4. Acanthosyris glabrata.
   • Plant with flowers, Ecuador. Photo by A. Gentry (No. 72366). Link goes to TROPICOS database, MO.
5. Acanthosyris spinescens
   • Close-up view1 and view2 of flowers. Florida: Río Viejo, Uruguay. Photo by Mauricio Bonifacino.
   • Plant in fruit. Called "granaditas" or "quebrachillos." Plants of the Chaco Orientale, Paraguay. Photo and website by Juan Antonio Alberto.

Cervantesia

1. Cervantesia tomentosa
   • Plant with fruits. Colombia. Photo by A. Gentry (No. 65359). Link goes to TROPICOS image library MO.
   • Habit of plant. Ecuador, Loja. Voucher (No. 2283) and photographs by Jørgensen, Ulloa, Caranqui and Sánchez. Link goes to TROPICOS image library MO.
   • Close-up of flower, face view. Same as above. Link goes to TROPICOS image library MO.
   • Close-up of flower. Same as above.
   • Base of fruit. Same as above
   • Fruits, various views. Same as above.
   • Herbarium specimen, from Missouri Botanical Gardens (MO).

Jodina

1. Jodina rhombifolia
   • Habit of plant. Photo by L. J. Musselman.
   • Flowering plant. Photo by L. J. Musselman.
   • Close-up of flowers. Photo by L. J. Musselman.
   • Close-up of open flower. Maldonado, Route 81, Uruguay.  Photo by M. Bonifacino.
   • Fruit.  Maldonado, Route 81, Uruguay.  Photo by M. Bonifacino.
   • Fruit.  Note, when the fruit is ripe the outer exocarp dehisces leaving a clear to whitish inner exocarp. This is the stage where the fruit is attractive to seed dispersing birds. Planted tree in Parque Saavedra, La Plata, Buenos Aires province, Argentina. Photo taken October (spring) 2005 by Gabriela Ruellan.
   • Fruit, closer view. Planted tree in Parque Saavedra, La Plata, Buenos Aires province, Argentina. Photo taken November (spring) 2007 by Gabriela Ruellan.
   • Bark.  Maldonado, Route 81, Uruguay.  Photo by M. Bonifacino.

Okoubaka

1. Okoubaka aubrevillei
   • Habit of the plant. It's hard to believe that this large (up to 40 m tall) tree is actually a root parasite in the sandalwood family! But indeed the hemiparasitic nature of this species has been documented (see Veenendaal et al., 1996, New Phytol. 134:487-493 and Swaine & Hall, 1986 listed therein). Atewa Range Forest Reserves, Ghana. Photo. 1977 by M.D. Swaine.
   • Aspect of entire tree showing abundant epiphytes. It is from this tree that haustoria were first recorded for the species. Atewa Range Forest Reserves, Ghana. Photo. 1977 by M.D. Swaine.
   • Seed with millimeter scale above. The dry mass of the nut (seed plus bony mesocarp) averages 43 g, the largest reported for a hemiparasitic plant. Atewa Range Forest Reserves, Ghana. Photo 1977 by M. D. Swaine.
   • Herbarium specimen, showing inflorescence, Missouri Botanical Gardens (MO).

Pilgerina

1. Pilgerina madagascariensis
   
   • Habit of the plant. Toliara, Mandena Forestry Reserve. Rogers et al. 890, type collection. Photo by Zach Rogers.
   • Inflorescence in hand. Toliara, Mandena Forestry Reserve. Rogers et al. 890, type collection. Photo by Zach Rogers.
   • Inflorescence showing closer view of flowers. Madagascar, Toliara, Mandena Forestry Reserve. Rogers et al. 890, type collection. Photo by Zach Rogers.
   • Fruiting branch. Toliara, Sainte Luce Forestry Reserve. Rogers et al. 967. Photo by Zach Rogers.
   • Fruits in section, photo 1, photo 2. Madagascar, Toliara, Sainte Luce Forestry Reserve. Rogers et al. 976. Photos by Zach Rogers.
   • For detailed drawings of the plant, see Rogers et al. (2008), pdf file HERE.

Pyrularia

1. Pyrularia edulis
   • Line drawing of this Chinese species from Flora of China. Web page HERE.
2. Pyrularia pubera
   • Habit of plant with male inflorescence. West Virginia, USA. Photo by L. J. Musselman.
   • Close-up of a possibly bisexual flower. Virginia, USA. [DLN 2737]. Photo by Ken Robertson.
   • Plant with inflorescences. From Flora of Southwest Virginia. © Craig Van Boskirk. Web page HERE.
   • Inflorescence and close-up of inflorescence, from Flora of Southwest Virginia. © Craig Van Boskirk.
   • Flowers, dissected. Virginia, USA. [DLN 2737]. Photo by Ken Robertson.
   • Fruits. Photo by Dan Busemeyer, Illinois Natural History Survey.
   • Fruiting branch removed. West Virginia, USA. Photo by L. J. Musselman.
   • Close-up showing apex of fruit with persistent perianth. West Virginia, USA. Photo by L. J. Musselman.
   • Fruit dissected and nut removed. South Carolina, USA. Photo by L. J. Musselman.
   • Fruit in cross section (left) and nut exposed (right). Photo by L. J. Musselman.
   • Haustoria, connected to host root (Abies fraseri, Fraser fir). West Virginia, USA. Photo by L. J. Musselman.
   • A Fraser fir tree parasitized by Pyrularia in a Christmas tree plantation. West Virginia, USA. Photo by L. J. Musselman.

Scleropyrum

1. Scleropyrum maingayai
   
   • Inflorescence.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
   • Closer view of flowers.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
2. Scleropyrum wallichianum.
   • Trunk of the tree showing the characteristic spines. Singapore. Photo by Joseph Lai. Visit his website HERE.
   • Branches with young male inflorescences. From Flora of Western Ghats, India. Photo by K. A. Subramanian, Centre for Ecological Sciences, IISc, Bangalore. Website HERE.
   • Young male inflorescence and open flower. Thailand. Photo taken by Somran Suddee and provided by J. Macklin.
   • Fruits. Black and white photo from Rao, L. N. (1942. Studies in the Santalaceae. Ann. Bot., N.S. 6: 151-175.).
   • Herbarium specimen, Missouri Botanical Gardens (MO).
   • Herbarium specimen, Missouri Botanical Gardens (MO). Close-up of inflorescence.
   • Herbarium specimen, Missouri Botanical Gardens (MO). Fruits.
   • Line drawing from Flora of China. Website HERE.

A NOTE ABOUT SCLEROPYRUM PENTANDRUM. I was informed by Boonying Tubtim, faculty of pharmacy at Chiang Mai University in Thailand, that there are reports of people eating this plant by mistake (thinking it is Melientha suavis in Opiliaceae) and dying from poisoning. Any information about the toxic substances present in this species should be relayed to Mr. Tubtim at: Boony-ta@suthep.pharmacy.cmu.ac.th. Jill Macklin provided information she gleaned from herbarium labels that the flowers of S. aurantiacum are eaten in New Guinea, so the toxic components may not be present in all species of Scleropyrum.

Staufferia

1. Staufferia capuronii
   
   • Fruiting branch.  Madagascar, Antsiranana Province, Anivorano Nord. Ratovoson et al. 877. Photo by Fidisoa Ratovoson.
   • Fruiting branch, closer view.  Madagascar, Antsiranana Province, Ambilobe. Callmander et al. 317. Photo by Martin Callmander.
   • Immature fruit. RFA 877.  Madagascar, Antsiranana Province, Anivorano Nord. Ratovoson et al. 877. Photo by Fidisoa Ratovoson.
   • For detailed drawings of the plant, see Rogers et al. (2008), pdf file HERE.

Nanodea Clade

Mida

1. Mida salicifolia
   • Plant in vegetative condition. The "shiny-leaf maire" of the North Island, New Zealand. Photo by Tony Foster.
   • Herbarium specimen, Missouri Botanical Gardens (MO).

Nanodea

1. Nanodea muscosa
   • Close-up of the tiny flowering plants. Tierra del Fuego. Photograph by Javier Puntieri.
   • Bright red fruit, huge in comparison to the size of the entire plant! Tierra del Fuego. Photograph by Javier Puntieri.

Santalum Clade

Antidaphne

1. Antidaphne punctulata
   • Flowering plant parasitic on Desfontainia spinosa. Nahuelbuta Chile. January 2002. Photo by G. Amico.
   • Close-up of flowering shoot. Plant parasitic on Desfontainia spinosa. Nahuelbuta Chile. Photo by G. Glatzel.
2. Antidaphne viscoidea
   • Shoots of mistletoe with inflorescence buds. Monte Verde, Costa Rica. Photo by D. L. Nickrent.
   • Close-up of shoot with floral buds. San Ramon, Costa Rica. Photograph by Wolfgang Forsteuter, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).

Colpoon

1. Colpoon compressum

• Habit of plant with flowers and fruits. Goedverwag, South Africa. Photo by D. L. Nickrent.
• View1 and View 2 of flowering and fruiting shoots. Goedverwag, South Africa. Photo by D. L. Nickrent.
• Close-up of flowering and fruiting shoots. Kirstenbosch Botanical Garden, Capetown, South Africa. Photo by D. L. Nickrent.
• Habit of plant with mature fruits. Roberg Nature Reserve, South Africa. Photo by D. L. Nickrent.

Eubrachion

1. Eubrachion ambiguum
   • The mistletoe on its host tree, Pimenta (Myrtaceae). Santana Peak, Costa Rica. [DLN 2699]. Photo by D. L. Nickrent.
   • Closer view of the mistletoe shoots. Same as above. Photo by D. L. Nickrent.
   • Close-up of the "ament-like" staminate spikes. Same as above. Photo by D. L. Nickrent.
   • Mistletoe on host branch. Southern Brazil. Photograph by Wolfgang Forsteuter, published in "World of Plants" article by Mitsuru Hirota (Weekly Asahihyakka, Jan. 22, 1995).

Exocarpos

1. Exocarpos aphyllus
   • Habit. Link goes to Australian National Botanic Gardens Photo Collection.
2. Exocarpos casuarinoides
   • Habit of plant. Mt. des Sources, New Caledonia. September 3, 1981. Photo by Job Kuijt.
   • Close-up of fruiting shoots. Mt. des Sources, New Caledonia. September 3, 1981. Photo by Job Kuijt.
3. Exocarpos cupressiformis
   • This parasitic plant is a fairly large tree. NSW, Australia. Photo by D. L. Nickrent.
   • Plant showing receptacular "false fruit." The Cherry Ballart. Photo from Costermans (1983).
   • Shoots with fruits. This is the only one of the five species of Exocarpos to reach tree size in Tasmania. Photo by Greg Jordan.
4. Exocarpos gaudichaudii
   • Habit Link goes to University of Hawaii Vascular Plant Families.
   • Close-up of flowers. Link goes to University of Hawaii Vascular Plant Families.
   • Plant in fruit.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
5. Exocarpos humifusus
   • Close-up showing the fruit and swollen receptacle, both red, of this alpine shrubby species. Tasmania. Photo by Greg Jordan.
6. Exocarpos latifolius
   • Plant with immature fruit. Queensland, Australia. [DLN 2801]. Photo by D. L. Nickrent.
   • Fruit. Link goes to Australian National Botanic Gardens Photo Collection.
7. Exocarpos luteolus
   • Habit of the plant, fruits. Link goes to University of Hawaii Vascular Plant Families.
   • Plant in fruit.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
8. Exocarpos menziesii
   • Habit of plant. South Kona District, Hawai'i, Hawai'i. [DLN 4297]. Photo by D. L. Nickrent.
   • Close-up of inflorescence. Same as above. Photo by D. L. Nickrent.
   • Two pictures of fruits and two of plant habit. Link goes to University of Hawaii Vascular Plant Families.
9. Exocarpos neocaledonicus
   
   • Habit of plant.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
10.  Exocarpos strictus
    • Fruiting branches. Link goes to Australian National Botanic Gardens Photo Collection.
    • Flowering branch. Shrub of the dry forests of Tasmania. Photo by Greg Jordan (see also Key to Tasmanian Dicots).
    • Fruiting branch. Tasmania. Photo by Greg Jordan
11. Exocarpos sparteus
    • Habit of plant. No voucher information. Link goes to TROPICOS image library MO.
    • Branches of plant. No voucher information. Link goes to TROPICOS image library MO.
    • Close-up of branches of plant. No voucher information. Link goes to TROPICOS image library MO.
12. Exocarpos syrticola
    • Habit of the plant, growing on the exposed, rocky coasts of Tasmania. Photo by Greg Jordan.

Lepidoceras

1. Lepidoceras chilense
   • Male plant in full flower. Nahuelbuta Chile. Photo by G. Glatzel.
   • Close-up of male flowers and young inflorescence buds still enclosed by bracts. Nahuelbuta Chile. Photo by G. Glatzel.
   • Haustorial attachment to host. Puyehue, Chile. Photo by Delbert Wiens.
   • Haustorial connection to host. Nahuelbuta Chile. Photo by G. Glatzel.
   • Habit of plant in fruit. Osorno, Chile. February 2003. Photo by Guillermo Amico.
   • Close-up of fruiting stem. Note the sclerotic "nail" at the apex of the leaves. Puyehue, Chile. Photo by Delbert Wiens.
   • Habit of plant and close-up of leaves. Parasitic on Laurelia sempervirens. Voucher: J. Solomon no. 4574 (given as L. kingii). Link goes to TROPICOS image library MO.

Myoschilos

1. Myoschilos oblongata
   • Habit of plant. Trail to Mt. Fitzroy, Argentina. .Photo1996 by Job Kuijt.
   • Habit of plant. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo by Norlan Tercero Bucardo.
   • Close-up of leaves. Close-up of leaves. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo by Norlan Tercero Bucardo.
   • Plant with young fruits forming on terminal portions of the spike. The yellow structures below the fruit are undeveloped flowers and bracts. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo taken in November, 2002 by Norlan Tercero Bucardo.
   • Another view of the flowering/fruiting plant. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo by Norlan Tercero Bucardo.
   • Fruiting plant. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo taken January 2003 by Norlan Tercero Bucardo.
   • Fruits forming on short shoots arising from the stems. Arroyo Blanco, Bariloche, Nahuel Huapi National Park, Argentina. Photo by Norlan Tercero Bucardo.
   • Herbarium specimen, Missouri Botanical Gardens (MO). Showing small leaves present on flowering branches. Photo by D. L. Nickrent.
   • Herbarium specimen, Missouri Botanical Gardens (MO). Showing larger leaves present on older branches. Photo by D. L. Nickrent.

Nestronia

1. Nestronia umbellula
   • Habit of plant. Banister River, Virginia, USA. Photo by L. J. Musselman.
   • Plant with male flowers. Banister River, Virginia, USA. Photo by L. J. Musselman.
   • Staminate flowers, some in section. [DLN 2736]. Virginia, USA. Photo by K. Robertson.
   • Fruit. Banister River, Virginia, USA. Photo by L. J. Musselman.

Omphacomeria

1. Omphacomeria acerba
   • Herbarium specimen, Missouri Botanical Gardens (MO).
   • Flowering branches. Australia. Link goes to Australian National Botanic Gardens Photo Collection.

Osyris

1. Osyris alba (including O. quadripartida; see Stauffer 1961)
   • Plant with staminate flowers. Bonifacio, Corsica, France. From Plant Image Gallery by Thomas Schoepke.
   • Habit of staminate plant in flower. Greece (Samos). Photo by Thomas Zumbrunn. For more, see the Botanical Image Database.
   • Close-up of staminate flowers. Greece (Samos). Photo by Thomas Zumbrunn. For more, see the Botanical Image Database.
   • Staminate plant. Cadiz, Spain. [DLN 2830] Photo by D. L. Nickrent.
   • Carpellate plant with fruit. Cadiz, Spain. [DLN 2830] Photo by D. L. Nickrent.
   • Fruit. Greece (Samos). Photo by Thomas Zumbrunn. For more, see the Botanical Image Database.
   • Habitat showing numerous plants in winter condition (February 2004). San Agustin de Guadalix, north of Madrid, Spain. Photo by D. L. Nickrent.
   • Fruiting plant in winter condition. San Agustin de Guadalix, north of Madrid, Spain. Photo by D. L. Nickrent.
   • Closer view of fruiting plant. San Agustin de Guadalix, north of Madrid, Spain. Photo by D. L. Nickrent.
2. Osyris lanceolata
   • Close-up of stem with male flowers. Witwatersrand Botanical Garden, South Africa. Photo by L. J. Musselman.
   • Seedling. Arrow indicates the cotyledons inside the fruit. Plant is native to S. Africa, here cultivated in the SIUC greenhouse, Illinois. [DLN 2830] Photo by D. L. Nickrent.
   • Potted plant. These plants were grown in pots from seeds (above photo) without a host. They flowered for several seasons and then unexpectedly died. Photo by D. L. Nickrent.

Rhoiacarpos

1. Rhoiacarpos capensis
   • Fruiting shoots. Near Grahamstown, South Africa. [DLN 4117]. Photo by D. L. Nickrent.
   • Flowering and fruiting shoots. Same as above. Photo by D. L. Nickrent.
   • Closer view of flowering and fruiting shoots. Same as above. Photo by D. L. Nickrent.

Santalum

Santalum species of Hawaii. Link goes to Australian National Botanic Gardens Photo Collection.

1. Santalum acuminatum
   
   • Habit of plant.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
   • Close-up of fruiting branches.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
   • Fruiting branches. Link goes to Australian National Botanic Gardens Photo Collection.
2. Santalum album
   • Sandalwood objects. The aromatic wood of this species is used for many purposes, such as these bookmarks, pens, and key chains.
3. S. ellipticum
   • Flowers. Link goes to University of Hawaii Vascular Plant Families.
4. S. freycinetianum var. freycinetianum
   • Habit. Link goes to University of Hawaii Vascular Plant Families.
   • Close-up of flowers (beautiful!). Link goes to University of Hawaii Vascular Plant Families.
   • Flowering shoots.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
5. S. freycinetianum var. pyrularium
   • Flowers, fruits. Link goes to University of Hawaii Vascular Plant Families.
6. Santalum haleakalae
   • Habit. Link goes to University of Hawaii Vascular Plant Families.
   • Flowers. Link goes to University of Hawaii Vascular Plant Families.
   • Flowering shoots.  Photo by R. A. Howard (see Plant Image Collection, Smithsonian Inst.)
7. Santalum lanceolatum
   • Flowering & fruiting shoots. Link goes to Australian National Botanic Gardens Photo Collection.
8. Santalum macgregorii
   • Close-up of flowers. Cultivated at the National Botanical Gardens, Lae. Original seed collected by John Beko near Apanaipi, Gulf District, Papua New Guinea. [DLN 4499]. Photo by D. L. Nickrent.
9. Santalum murrayanum
   • Fruiting shoot. Link goes to Australian National Botanic Gardens Photo Collection.
10. Santalum paniculatum
    • Shoot with young inflorescence. Foster Botanical Garden, Honolulu, Hawaii. [DLN 2830] Photo by D. L. Nickrent.
    • Flowering shoots. North Kohala, Hawai'i, Hawai'i. [DLN 4296]. Photo by D. L. Nickrent.
    • Inflorescence. North Kohala, Hawai'i, Hawai'i. [DLN 4296]. Photo by D. L. Nickrent.
    • Fruits. Mary Greenwell Ethnobotanical Garden, Captain Cook, Hawai'i. Photo by D. L. Nickrent.

Amphorogyne Clade

Amphorogyne

1. Amphorogyne celastroides
   • Female flowers. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
   • Fruits. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
2. Amphorogyne spicata
   • Habit of plant, 2.4 m in height. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
   • Branch with male flowers. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
   • Male flowers, 2.0 - 2.2 mm wide. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
   • Fruit. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
3. Amphorogyne staufferi
   • Herbarium specimen, Missouri Botanical Gardens (MO).
4. Amphorogyne sp.
   • Vegetative shoots. Mt. des Sources, New Caledonia. September 3, 1981. Photo by Job Kuijt.

Choretrum

1. Choretrum candollei
   • Flowering branches. Link goes to Australian National Botanic Gardens Photo Collection.
2. Choretrum pauciflorum.
   • Flowering shoots. SE Australia. Photo by M. Fagg (from Flora of Australia, Vol. 22, 1984).
3. Choretrum spicatum.
   • Female flowers (left) and fruits (right). Australia. Photo from Stauffer (1969, Santalales Studien X).

Daenikera

1. Daenikera corallina

• Shoots with flowers and fruits. New Caledonia. Photo by Jérôme Munzinger.
• Shoots. New Caledonia. Photo by Porter Lowry.
• Fruit. New Caledonia. Photo by Porter Lowry.
• Habit of plant, showing shoots emerging from host tree. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
• Close-up of haustorial connection to host tree. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
• Shoots with female flowers. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).
• Shoots with fruits. New Caledonia. Photo from Stauffer (1969, Santalales Studien X).

Dendromyza (including Cladomyza)

1. Dendromyza acroslera
   • Herbarium specimen, Missouri Botanical Gardens (MO).
2. Dendromyza ledermannii
   • Branch with male flowers. Photo from Stauffer (1969, Santalales Studien X).
   • Branch with female flowers and fruits. Photo from Stauffer (1969, Santalales Studien X).
3. Dendromyza sp. cf. ledermannii
   • Habit of plant. Along road below Nanduo, between Boring and Masanko Junction, Morobe District, Finschhafen Subdistrict, Papua New Guinea. 2668 ft (813 m) elev. [Banka 4473]. Photo by D. L. Nickrent.
   • Male plant. Hills above Nanduo Village, along trail to Sasaniko, Morobe District, Finschhafen Subdistrict, Papua New Guinea.3301 ft (1006 m) elev. [Banka 4466]. Photo by D. L. Nickrent.
   • Close-up of fruiting plant. Wagau Village, edge of yam field, Morobe District, Papua New Guinea. 3672 ft (1119 m) elev. [Banka 4483]. Photo by D. L. Nickrent.
4. Dendromyza reinwardtiana
   • Habit of plant. Photo from Stauffer (1969, Santalales Studien X).
5. Dendromyza uncinata
   • Branch with female flowers. Photo from Stauffer (1969, Santalales Studien X).
   • Haustorial branches on Ceratopetalum sp. Photo from Stauffer (1969, Santalales Studien X).
6. Dendromyza sp.
   • Herbarium specimen, Missouri Botanical Gardens (MO).

Dendrotrophe (including Henslowia)

1. Dendrotrophe varians
   • Plant in fruit. Sarawak, Malaysia. [DLN 4014] Photo by D. L. Nickrent.
   • Branch with male flowers. Photo from Stauffer (1969, Santalales Studien X).
2. Dendrotrophe sp. 1
   • Shoots bearing fruits. Menyamya Highway, near Okinaiwa, Papua New Guinea. 1903 m elev. Parasitic on Glochidion (Euphorbiaceae). This is likely an unnamed species. [DLN 4498] Photo by D. L. Nickrent
   • Fruit, dissected to show the bright red mesocarp inside of leathery exocarp. The seed on the right is without the mesocarp, exposing the hard endocarp and attached viscin strands. Photo by D. L. Nickrent.
3. Dendrotrophe sp. 2
   • Ripe fruits. Summit of Gunung Tahan, alt. ca. 2300 m. Photo by E. Soepadmo (Nature Malaysiana; 1978, 3:24-31).

Dufrenoya (including Hylomyza) - no photos

Leptomeria

1. Leptomeria aphylla
   • Fruiting spikes. Grampians, Victoria, Australia. Photo from Costermans (1983).
2. Leptomeria drupacea
   • Flowering spikes. Genoa R, Victoria, Australia. Photo (listed as L. acida) from Costermans (1983).
   • Flowering shoots. Tasmania. Slide 10064 archived at Kew. Photo 1965 by T. E. Burns.
3. Leptomeria lehmannii
   • Female flowers. Australia. Photo (as L. pauciflora) from Stauffer (1969, Santalales Studien X).
   • Fruits. Australia. Photo (as L. pauciflora) from Stauffer (1969, Santalales Studien X).
4. Leptomeria pachyclada
   
   • Flowering branches. Australia. Link goes to Australian National Botanic Gardens Photo Collection.
5. Leptomeria pauciflora
   
   • Flowering branch.  Australia.  Photo by Brendan Lepschi.

Phacellaria

1. Phacellaria rigidula (?)
   
   •  Habit of vegetative plant parasitic on Dentrotrophe fructescens. Hong Kong.  Photo September 2007 by Shek-shing Mar.
   • Close-up of shoot showing what appears to be caducous scale leaves.  Hong Kong.  Photo September 2007 by Shek-shing Mar.
2. Phacellaria sp.
   • A hyperparasite on Macrosolen. Inflorescence/infructescence. Malaysia: Pahang, Selonger Genting Highlands. Photograph by John Dransfield, Dec. 1977. Slide no. 3961 archived at Kew.
   • Close-up of infructescence. Same as above. Photograph by John Dransfield, Dec. 1977. Slide no. 3962 archived at Kew.

Spirogardnera

1. Spirogardnera rubescens
   • Composite photograph, from FloraBase, the Western Australian Flora. Left photo, by J. L. Roberts showing close-up of the inflorescence. Top photo, by Susan J. Patrick showing habit of plant along a roadside. Bottom photo, by Mary Hancock, showing the plant emerging along a gravel road following disturbance.
   • Flowering branches. Western Australia. Photo from Stauffer (1968, Santalales Studien IX).
   • Flowers. Western Australia. Photo from Stauffer (1968, Santalales Studien IX).
   • Fruits. Western Australia. Photo from Stauffer (1968, Santalales Studien IX).

Classifications, past and present

Phylogeny

References