The Icacinaceae, also called the white pear family, are a

family Family (from la, familia) is a Social group, group of people related either by consanguinity (by recognized birth) or Affinity (law), affinity (by marriage or other relationship). The purpose of the family is to maintain the well-being of its ...

flowering plant Flowering plants are plants that bear flowers and fruits, and form the clade Angiospermae (), commonly called angiosperms. The term "angiosperm" is derived from the Greek words ('container, vessel') and ('seed'), and refers to those plants th ...

s,"Icacinaceae" At: Angiosperm Phylogeny Website At: Missouri Botanical Garden Website (see ''External links'' below). consisting of

tree In botany, a tree is a perennial plant with an elongated stem, or trunk, usually supporting branches and leaves. In some usages, the definition of a tree may be narrower, including only woody plants with secondary growth, plants that are ...

shrub A shrub (often also called a bush) is a small-to-medium-sized perennial woody plant. Unlike herbaceous plants, shrubs have persistent woody stems above the ground. Shrubs can be either deciduous or evergreen. They are distinguished from trees ...

s, and

liana A liana is a long- stemmed, woody vine that is rooted in the soil at ground level and uses trees, as well as other means of vertical support, to climb up to the canopy in search of direct sunlight. The word ''liana'' does not refer to a ta ...

s, primarily of the

tropics The tropics are the regions of Earth surrounding the Equator. They are defined in latitude by the Tropic of Cancer in the Northern Hemisphere at N and the Tropic of Capricorn in the Southern Hemisphere at S. The tropics are also referred to ...

. The family was traditionally

circumscribed In geometry, the circumscribed circle or circumcircle of a polygon is a circle that passes through all the vertices of the polygon. The center of this circle is called the circumcenter and its radius is called the circumradius. Not every polyg ...

quite broadly, with around 55

genera Genus ( plural genera ) is a taxonomic rank used in the biological classification of living and fossil organisms as well as viruses. In the hierarchy of biological classification, genus comes above species and below family. In binomial nomenclat ...

totalling over 400

species In biology, a species is the basic unit of classification and a taxonomic rank of an organism, as well as a unit of biodiversity. A species is often defined as the largest group of organisms in which any two individuals of the appropriate s ...

. In 2001, though, this circumscription was found to be

polyphyletic A polyphyletic group is an assemblage of organisms or other evolving elements that is of mixed evolutionary origin. The term is often applied to groups that share similar features known as homoplasies, which are explained as a result of converg ...

, and the family was split into four families in three different

orders Order, ORDER or Orders may refer to: * Categorization, the process in which ideas and objects are recognized, differentiated, and understood * Heterarchy, a system of organization wherein the elements have the potential to be ranked a number of d ...

: Icacinaceae ''

sensu stricto ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

'' (then unplaced at order rank),

Pennantiaceae ''Pennantia'' is the sole genus in the plant family Pennantiaceae. In older classifications, it was placed in the family Icacinaceae. Most authorities have recognised three or four species, depending on whether they recognised ''Pennantia baylis ...

(

Apiales The Apiales are an order of flowering plants. The families are those recognized in the APG III system. This is typical of the newer classifications, though there is some slight variation and in particular, the Torriceliaceae may also be divide ...

Stemonuraceae Stemonuraceae is a eudicot family of flowering plants. Genera This listing was sourced from the Angiosperm Phylogeny Website and Royal Botanic Gardens, Kew * '' Cantleya'' * '' Codiocarpus'' * '' Discophora'' * '' Gastrolepis'' * ''Gomphand ...

(

Aquifoliales The Aquifoliales are an order of flowering plants, including the Aquifoliaceae (holly) family, and also the Helwingiaceae (2-5 species of temperate Asian shrubs) and the Phyllonomaceae (4 species of Central American trees and shrubs). In 2001, t ...

) and

Cardiopteridaceae Cardiopteridaceae is a eudicot family of flowering plants. It consists of about 43 species of trees, shrubs, and woody vines, mostly of the tropics, but with a few in temperate regions.Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alas ...

(also Aquifoliales). Other genera have later been moved to

Metteniusaceae Metteniusaceae are a family of flowering plants, the only family in the order Metteniusales. It consists of about 10 genera and 50 species of trees, shrubs, and lianas, primarily of the tropics. The family was formerly restricted to just '' Mette ...

( Metteniusales),Stull, G. W., R. Duno de Stefano, D. E. Soltis, and P. S. Soltis (2015). Resolving Basal Lamiid Phylogeny and the Circumscription of Icacinaceae with a Plastome-Scale Data Set. American Journal of Botany 102, no. 11: 1794–1813. doi:10.3732/ajb.1500298. so that Icacinaceae now include c. 23 genera and 160 species. One genus, ''Sleumeria'', was described as late as 2005.Timothy M.A. Utteridge, Hidetoshi Nagamasu, Stephen P. Teo, Lydia C. White, and Peter Gasson. 2005. "''Sleumeria'' (Icacinaceae): A New Genus from Northern Borneo". ''Systematic Botany'' 30(3):635-643. Icacinaceae belongs to the order

Icacinales Icacinales is an order of Angiosperms. The order did not exist in the 2009 APG III system, and was added in the 2016 APG IV system, including two families, Icacinaceae The Icacinaceae, also called the white pear family, are a family of flow ...

along with

Oncotheca ''Oncotheca'' is a genus of tree endemic to New Caledonia. There are two species, '' Oncotheca balansae'' and '' Oncotheca humboldtiana''. ''Oncotheca'' is the sole genus of the Oncothecaceae, which is one of three families of flowering plants ...

ceae. ''Icacina senegalensis'' extracts have shown activity against malaria parasites.Sarr SO, Perrotey S, Fall I, Ennahar S, Zhao M, Diop YM, Candolfi E, Marchioni E.,"Icacina senegalensis (Icacinaceae), traditionally used for the treatment of malaria, inhibits in vitro Plasmodium falciparum growth without host cell toxicity." ''Malar J''. 2011 Apr 11;10(1):85

Genera

List according to Stull et al. (2015): Icacineae * ''

Alsodeiopsis ''Alsodeiopsis'' Oliver is a genus of plants in the family Icacinaceae native to tropical Africa. There are about 11 species. Seventeen species names have been published in ''Alsodeiopsis'',The International Plant Names Index. (see ''External li ...

'' Oliver * ''

Casimirella ''Casimirella'' is a genus of flowering plants belonging to the family Icacinaceae The Icacinaceae, also called the white pear family, are a family of flowering plants,"Icacinaceae" At: Angiosperm Phylogeny Website At: Missouri Botanical Garde ...

'' Hassler * ''

Icacina ''Icacina'' is a genus of flowering plants belonging to the family Icacinaceae. Its native range is Cape Verde, Western Tropical Africa Although tropical Africa is mostly familiar to the West for its rainforests, this biogeographic realm of ...

'' A. Jussieu * '' Lavigera'' Pierre * ''

Leretia ''Leretia'' is a monotypic genus of flowering plants belonging to the family Icacinaceae. The only species is ''Leretia cordata'' Its native range is Costa Rica to southern Tropical America. It is also found in the countries of Bolivia, Brazil, C ...

'' Vellozo * ''

Merrilliodendron ''Merrilliodendron'' is a monotypic genus of flowering plants belonging to the family Icacinaceae. It has a synonym of ''Peekeliodendron'' . The only species is ''Merrilliodendron megacarpum'' Its native range is Central Malesia to western Pacif ...

'' Kanehira * ''

Pleurisanthes ''Pleurisanthes'' is a genus of flowering plants belonging to the family Icacinaceae The Icacinaceae, also called the white pear family, are a family of flowering plants,"Icacinaceae" At: Angiosperm Phylogeny Website At: Missouri Botanical Gard ...

'' Baillon Iodeae * ''

Iodes ''Iodes'' is a genus of flowering plants belonging to the family Icacinaceae. Its native range is Tropical Africa, Madagascar, Tropical and Subtropical Asia. Species: *'' Iodes africana'' *'' Iodes cirrhosa'' *'' Iodes globulifera'' *'' Iod ...

'' Blume * †'' Iodicarpa'' Manchester * '' Mappianthus'' Handel-Mazzetti Mappieae * '' Mappia'' Jacquin * '' Nothapodytes'' Blume Phytocreneae * '' Desmostachys'' Miers * '' Hosiea'' Hemsley & E. H. Wilson * '' Miquelia'' Meisner * '' Natsiatopsis'' Kurz * '' Natsiatum'' Arnott * †'' Palaeophytocrene'' Reid & Chandler * '' Phytocrene'' Wallich * '' Pyrenacantha'' Wight * '' Rhyticaryum'' Beccari * '' Sarcostigma'' Wight & Arnott * ''

Sleumeria ''Sleumeria'' is a genus of flowering plants belonging to the family Icacinaceae. Its native range is Borneo Borneo (; id, Kalimantan) is the third-largest island in the world and the largest in Asia. At the geographic centre of Maritim ...

'' Utteridge ''et al''. * '' Stachyanthus'' Engler * '' Vadensea'' Jongkind & O. Lachenaud ''Incertae sedis'' * '' Cassinopsis'' Sonder * †'' Goweria''

History

In 1841,

George Bentham George Bentham (22 September 1800 – 10 September 1884) was an English botanist, described by the weed botanist Duane Isely as "the premier systematic botanist of the nineteenth century". Born into a distinguished family, he initially studi ...

described ''Apodytes'' and ''Pogopetalum'' as new

and united them with ''Icacina, Gomphandra'', and ''Leretia'' to create the

tribe The term tribe is used in many different contexts to refer to a category of human social group. The predominant worldwide usage of the term in English language, English is in the discipline of anthropology. This definition is contested, in p ...

Icacineae of what would later be called the family

Olacaceae Olacaceae is a family (biology), family of flowering plants in the order Santalales. They are woody plants, native throughout the tropical regions of the world. , the Circumscription (taxonomy), circumscription of the family varies; some sources ...

.George Bentham. 1841. page 679. In: "Account of two new genera allied to Olacineae". ''Transactions of the Linnean Society of London'' 18:671-686 & plates 41 and 42. (see ''External links'' below). Olacaceae was at that time, and through the 20th century, defined broadly, encompassing several families in the order

Santalales The Santalales are an order of flowering plants with a cosmopolitan distribution, but heavily concentrated in tropical and subtropical regions. It derives its name from its type genus ''Santalum'' (sandalwood). Mistletoe is the common name for a ...

.Valéry Malécot and Daniel L. Nickrent. 2008. "Molecular Phylogenetic Relationships of Olacaceae and Related Santalales". ''Systematic Botany'' 33(1):97-106. ''Pogopetalum'' was later synonymized with ''Emmotum''.Rodrigo Duno de Stefano, Diego F. Angulo, and Fred W. Stauffer. 2007. "''Emmotum harleyi'', a New Species from Bahia, Brazil, and Lectotypification of Other Icacinaceae". ''Novon'' 17(3):306-309.Richard A. Howard. 1942. "Studies of the Icacinaceae. III. A revision of ''Emmotum''". ''Journal of the Arnold Arboretum'' 23:479-494. In 1852, John Miers argued that Bentham's Icacineae did not belong in Olacaceae and he raised them to the

taxonomic rank In biological classification, taxonomic rank is the relative level of a group of organisms (a taxon) in an ancestral or hereditary hierarchy. A common system consists of species, genus, family (biology), family, order (biology), order, class (b ...

of family as Icacinaceae.John Miers. 1852. page 221. In: "Observations on the Affinities of the Icacinaceae". ''Annals and Magazine of Natural History, iccluding Zoology, Botany, and Geology'', series 2. 9:218-226. (see ''External links'' below). Philippe van Tieghem realized that the family Icacinaceae, as

by Miers, consisted of groups that were only distantly related, and in 1897, he divided it into seven families.Philippe E.L. van Tieghem. 1897. page 842. In: "Sur les inséminées à nucelle pourvu d'un seul tégument formant la subdivisions des Unitegminées ou Icacinées". Séance du Mardi 20 Avril 1897. ''Comptes Rendus Hebdomadaires des Séances de l'Académie des Sciences'' 124:839-844.Philippe E.L. van Tieghem. 1897. "Sur les phanerogams sans graines, formant la divisions des inséminées". ''Bulletin de la Société Botanique de France'' 44:99-139. (see ''External links'' below). Van Tieghem's treatment in some ways anticipated the results of 21st century

phylogenetic In biology, phylogenetics (; from Greek φυλή/ φῦλον [] "tribe, clan, race", and wikt:γενετικός, γενετικός [] "origin, source, birth") is the study of the evolutionary history and relationships among or within groups o ...

studies Study or studies may refer to: General * Education **Higher education * Clinical trial * Experiment * Observational study * Research * Study skills, abilities and approaches applied to learning Other * Study (art), a drawing or series of drawin ...

, in particular, by his establishment of the families Emmotaceae and Leptaulaceae. His division of Icacinaceae into smaller families was not accepted and other authors continued to define Icacinaceae in the broad sense, known as Icacinaceae

sensu lato ''Sensu'' is a Latin word meaning "in the sense of". It is used in a number of fields including biology, geology, linguistics, semiotics, and law. Commonly it refers to how strictly or loosely an expression is used in describing any particular co ...

. In 1942,

Hermann Sleumer Hermann Otto Sleumer (February 21, 1906 in Saarbrücken – October 1, 1993 in Oegstgeest) was a Dutch botanist of German birth. The plant genera ''Sleumerodendron'' Virot (Proteaceae The Proteaceae form a family of flowering plants predomina ...

defined Icacinaceae broadly in his coverage of the family for

Die Natürlichen Pflanzenfamilien ''Die Natürlichen Pflanzenfamilien'' (1887–1915) by Adolf Engler (1844–1930) and Karl Anton Prantl is a complete revision of plant families down to generic level and often even further. As such it forms part of the Engler system of plant tax ...

.Hermann Sleumer. 1942. "Icacinaceae" pages 322-396. In: H.G. Adolf Engler and Karl A.E. Prantl, with Hermann Harms and Johannes Mattfeld (editors). ''Die Natürlichen Pflanzenfamilien'' volume 20b. Duncker and Humblot: Berlin, Germany. 1960 reprint of 1942 publication. Later authors did likewise. In the 1940s, Richard A. Howard wrote a series of papers on several of the genera. Sleumer wrote about the

Asia Asia (, ) is one of the world's most notable geographical regions, which is either considered a continent in its own right or a subcontinent of Eurasia, which shares the continental landmass of Afro-Eurasia with Africa. Asia covers an area ...

n genera in 1969,Hermann Sleumer. 1969. "Materials toward the knowledge of the Icacinaceae of Asia, Malesia, and adjacent areas". ''Blumea'' 17(1):181-264. and the

Malesia Malesia is a biogeographical region straddling the Equator and the boundaries of the Indomalayan and Australasian realms, and also a phytogeographical floristic region in the Paleotropical Kingdom. It has been given different definitions. The ...

n genera in 1971.Hermann Sleumer. 1971. "Icacinaceae" pages 1-87. In: Cornelis G.G.J. van Steenis (editor). ''Flora Malesiana'' series 1, volume 7. Noordhoff International Publishing: Leyden, Holland. Much of what is known about the family comes from regional

floras Flora is all the plant life present in a particular region or time, generally the naturally occurring (indigenous) native plants. Sometimes bacteria and fungi are also referred to as flora, as in the terms '' gut flora'' or '' skin flora''. E ...

such as

Flora of Australia The flora of Australia comprises a vast assemblage of plant species estimated to over 30,000 vascular and 14,000 non-vascular plants, 250,000 species of fungi and over 3,000 lichens. The flora has strong affinities with the flora of Gondwana, ...

Gordon P. Guymer. 1984. "Icacinaceae" pages 204-211. In: Alexander S. George (executive editor). ''Flora of Australia'' volume 22. Australian Government Publishing Service: Canberra, Australia. and

Flora of China The flora of China consists of a diverse range of plant species including over 39,000 vascular plants, 27,000 species of fungi and 3000 species of bryophytes.Wu, Z. Y., P. H. Raven & D. Y. Hong, eds. 2006. Flora of China. Vol. 22 (Poaceae). Sc ...

.Hua Peng and Richard A. Howard. 2008. "Icacinaceae" pages 505-514. In: Zhengyi Wu, Peter H. Raven, and Deyuan Hong (editors). ''Flora of China'' volume 11. Science Press: Beijing, China; Missouri Botanical Garden Press: St. Louis, Missouri, USA. In 2001, Jesper Kårehed, using a combination of morphological and

DNA sequence DNA sequencing is the process of determining the nucleic acid sequence – the order of nucleotides in DNA. It includes any method or technology that is used to determine the order of the four bases: adenine, guanine, cytosine, and thymine. Th ...

data, showed that Icacinaceae sensu lato was distantly

and was, at least arguably, the worst of the plant families. It is now known to have rivaled

Flacourtiaceae The Flacourtiaceae is a defunct family of flowering plants whose former members have been scattered to various families, mostly to the Achariaceae and Salicaceae. It was so vaguely defined that hardly anything seemed out of place there and it beca ...

as an unnatural assemblage of disparate groups. Kårehed divided it into four families:

, and Icacinaceae

. Pennantiaceae consists of the single genus ''

Pennantia ''Pennantia'' is the sole genus in the plant family Pennantiaceae. In older classifications, it was placed in the family Icacinaceae. Most authorities have recognised three or four species, depending on whether they recognised '' Pennantia bayli ...

'' and is the most basal

clade A clade (), also known as a monophyletic group or natural group, is a group of organisms that are monophyletic – that is, composed of a common ancestor and all its lineal descendants – on a phylogenetic tree. Rather than the English term, ...

in the campanulid order

.Jesper Kårehed. 2003. "The family Pennantiaceae and its relationships to Apiales". ''Botanical Journal of the Linnean Society'' 141(1):1-24.Gregory M. Plunkett, Gregory T. Chandler, Porter P. Lowry, Steven M. Pinney, and Taylor S. Sprenkle. 2004. "Recent advances in understanding Apiales and a revised classification". ''South African Journal of Botany'' 70(3):371-381. Stemonuraceae is a family of 12 genera in the campanulid order Aquifoliales. It is

sister A sister is a woman or a girl who shares one or more parents with another individual; a female sibling. The male counterpart is a brother. Although the term typically refers to a familial relationship, it is sometimes used endearingly to refer to ...

to Cardiopteridaceae. Before the

phylogeny A phylogenetic tree (also phylogeny or evolutionary tree Felsenstein J. (2004). ''Inferring Phylogenies'' Sinauer Associates: Sunderland, MA.) is a branching diagram or a tree showing the evolutionary relationships among various biological spec ...

produced by Kårehed, Cardiopteridaceae contained only '' Cardiopteris''. Kårehed transferred '' Citronella'', '' Gonocaryum'', and '' Leptaulus'' from Icacinaceae to this family, and provisionally placed ''

Metteniusa ''Metteniusa'' is a genus of flowering plants in the Family (biology), family Metteniusaceae. It was named by Gustav Karl Wilhelm Hermann Karsten, Hermann Karsten in 1860 for the Germans, German botanist Georg Heinrich Mettenius. It has seven sp ...

'', ''

Dendrobangia ''Dendrobangia'' is a genus of flowering plants in the Family (biology), family Metteniusaceae.Stevens, P.F. (2001 onwards)"Metteniusaceae" ''Angiosperm Phylogeny Website''. Retrieved 2016-06-09. It was formerly placed in the family Cardiopterida ...

'', and ''

Pseudobotrys ''Pseudobotrys'' is a genus of trees and shrubs in the Family (biology), family Cardiopteridaceae described as a genus in 1912. The entire genus is endemic to New Guinea. ;Species # ''Pseudobotrys cauliflora'' (Pulle) Sleumer 1940 # ''Pseudobot ...

'' there as well. ''Metteniusa'' was shown to be a

lamiid In the APG IV system (2016) for the classification of flowering plants, the name asterids denotes a clade (a monophyletic group). Asterids is the largest group of flowering plants, with more than 80,000 species, about a third of the total flowe ...

in 2007, and was placed in a family by itself.Favio González, Julio Betancur, Olivier Maurin, John V. Freudenstein, and Mark W. Chase. 2007. "Metteniusaceae, an early-diverging family in the lamiid clade". ''Taxon'' 56(3):795-800. The affinities of ''Dendrobangia'' and ''Pseudobotrys'' remain obscure. Some authors have continued to maintain Cardiopteridaceae as a

monogeneric In biology, a monotypic taxon is a taxonomic group (taxon) that contains only one immediately subordinate taxon. A monotypic species is one that does not include subspecies or smaller, infraspecific taxa. In the case of genera, the term "unispec ...

family, placing ''Citronella'', ''Gonocaryum'', ''Leptaulus'', ''Dendrobangia'', and ''Pseudobotrys'' in Leptaulaceae.Timothy M.A. Utteridge and Richard K. Brummitt. 2007. "Leptaulaceae" pages 191-192. In: Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada. (2007). The study by Kårehed showed ''Cardiopteris'' to be embedded in Leptaulaceae, but statistical support for this position was not strong. Some genera have later been moved to

( Metteniusales).

References

External links

Icacinaceae
At

At
Missouri Botanical Garden Website

Transactions of the Linnean Society of London 18:671 (1841)

Annals and Magazine of Natural History, (1852)

Bulletin de la Société Botanique de France. 1897
{{Taxonbar, from=Q133316 Asterid families