Ochnaceae

Ochnaceae is a family of flowering plants in the order Malpighiales.Vernon H. Heywood, Richard K. Brummitt, Ole Seberg, and Alastair Culham. ''Flowering Plant Families of the World''. Firefly Books: Ontario, Canada. (2007). . In the APG III system of classification of flowering plants, Ochnaceae is defined broadly, to include about 550 species,Maria do Carmo E. Amaral, and Volker Bittrich. 2014. "Ochnaceae". pages 253-268. In: Klaus Kubitzki (editor). 2014. ''The Families and Genera of Vascular Plants'' volume XI. Springer-Verlag: Berlin, Heidelberg, Germany. (print). (eBook). and encompasses what some taxonomists have treated as the separate families Medusagynaceae and Quiinaceae. In a phylogenetic study that was published in 2014, Ochnaceae was recognized in the broad sense,Julio V. Schneider, Pulcherie Bissiengou, Maria do Carmo E. Amaral, Ali Tahir, Michael F. Fay, Marco Thines, Marc S.M. Sosef, Georg Zizka, and Lars W. Chatrou. 2014. "Phylogenetics, ancestral state reconstruction, and a new infrafamilial classification of the pantropical Ochnaceae (Medusagynaceae, Ochnaceae s.str., Quiinaceae) based on five DNA regions". ''Molecular Phylogenetics and Evolution'' 78:199-214. . but two works published after APG III have accepted the small families Medusagynaceae and Quiinaceae.Zhenxiang Xi, Brad R. Ruhfel, Hanno Schaefer, André M. Amorim, Manickam Sugumaran, Kenneth J. Wurdack, Peter K. Endress, Merran L. Matthews, Peter F. Stevens, Sarah Mathews, and Charles C. Davis. 2012. "Phylogenomics and a posteriori data partitioning resolve the Cretaceous angiosperm radiation Malpighiales". ''PNAS'' (''Proceedings of the National Academy of Sciences of the United States of America'') 109(43):17519-17524. . (See ''External links'' below). These have not been accepted by APG IV (2016).

In this article, "Ochnaceae" will refer to the larger circumscription of the family, which is otherwise known as Ochnaceae sensu lato or as the ochnoids. In this sense the family includes 32 genera with about 550 species.

Ochnaceae, defined broadly or narrowly, is pan tropical in distribution, with a few species cultivated outside of this range. Ochnaceae is most diverse in the neotropics, with a second center of diversity in tropical Africa. It consists mostly of shrubs and small trees, and, in ''Sauvagesia'', a few herbaceous species. Many are treelets, with a single, erect trunk, but low in height. The Ochnaceae are notable for their unusual leaves. These are usually shiny, with closely spaced, parallel veins, toothed margins, and conspicuous stipules. Most of the species are buzz pollinated.Paul A. De Luca and Mario Vallejo-Marin. 2013. "What's the buzz about? The ecology and evolutionary significance of buzz pollination". ''Current Opinion in Plant Biology'' 16(4):429-435. . In eight of the genera in tribe Sauvagesieae, the flower changes form after opening, by continued growth of tissue within the flower.

A few species of ''Ochna'' are cultivated as ornamentals. Anthony Huxley, Mark Griffiths, and Margot Levy (1992). ''The New Royal Horticultural Society Dictionary of Gardening''. The Macmillan Press,Limited: London. The Stockton Press: New York. (set). ''Ochna thomasiana'' is probably the most commonly planted, but it is often misidentified in the horticultural literature.Warren L. Wagner, Derral R. Herbst, and Sy H. Sohmer. ''Manual of the Flowering Plants of Hawaii'', Revised Edition, 1999. Bishop Museum Press: Hololulu

The leaves of ''Cespedesia'' are sometimes to in length and are used for roofing.David J. Mabberley. 2008. ''Mabberley's Plant-Book'' third edition (2008). Cambridge University Press: UK. (See ''External links'' below). An herbal tea is made from the pantropical weed ''Sauvagesia erecta''.

In its evolution, Ochnaceae has been unusual, in " reverting" to character states that are regarded as ancestral or primitive. For example, an actinomorphic floral symmetry has appeared twice in the subfamily Ochnoideae. Also, two clades of Ochnaceae, one in Ochnoideae and another in Quiinoideae have a derived condition very close to apocarpy. The complete separation of the carpels ( apocarpy) is thought to be the ancestral state for angiosperms.Peter K. Endress. 2011. "Evolutionary diversification of the flowers in angiosperms". ''American Journal of Botany'' 98(3):370-396. . (See ''External links'' below).

Fossils attributed to Ochnaceae are known from the early Eocene of Mississippi.Daniel Danehy, Peter Wilf, and Stefan A. Little. 2007. "Early Eocene macroflora from the red hot truck stop locality (Meridian, Mississippi, USA)". ''Palaeontologia Electronica'' 10(3):17A:31pages. (See ''External links'' below). The age of the family is very roughly estimated at 100 million years.Susana Magallon, Khidir W. Hilu, and Dietmar Quandt. 2013. "Land plant evolutionary timeline: Gene effects are secondary to fossil constraints in relaxed clock estimation of age and substitution rates". ''American Journal of Botany'' 100(3):556-573. . (See ''External links'' below).

A great many genus names have been published in Ochnaceae.The Plant List: Ochnaceae. (See ''External links'' below). In a taxonomic revision of Ochnaceae, as three families, in 2014, only 32 of these genera were accepted; one in Medusagynaceae, four in Quiinaceae, and 27 in Ochnaceae s.s. In that same year, a 33rd genus, ''Neckia'', was reestablished in order to preserve the monophyly of another genus, ''Sauvagesia''.

The largest genera in Ochnaceae are: ''Ouratea'' (200 species), ''Ochna'' (85), ''Campylospermum'' (65), ''Sauvagesia'' (39), and ''Quiina'' (34). None of the larger genera has been the subject of a phylogenetic analysis of DNA sequences of selected genes. In one study of the subfamily Quiinoideae, based on the trn L-F intergenic spacer, only nine species were sampled from this subfamily.Julio V. Schneider, Ulf Swenson, Rosabelle Samuel, Tod Stuessy, and Georg Zizka. 2006. "Phylogenetics of Quiinaceae (Malpighiales): evidence from trnL-trnF sequence data and morphology". ''Plant Systematics and Evolution'' 257(3-4):189-203. .

Genera

The following list of 33 genera consists of ''Neckia'', which was resurrected in 2014, plus the 32 genera that were described in the most recent revision of Ochnaceae.Klaus Kubitzki. 2014. "Quiinaceae". pages 277-281. . In: Klaus Kubitzki (editor). 2014. ''The Families and Genera of Vascular Plants'' volume XI. Springer-Verlag: Berlin, Heidelberg, Germany. (print). (eBook). William C. Dickison and Klaus Kubitzki. 2014. "Medusagynaceae". pages 249-251. . In: Klaus Kubitzki (editor). 2014. ''The Families and Genera of Vascular Plants'' volume XI. Springer-Verlag: Berlin, Heidelberg, Germany. (print). (eBook). The classification is from Schneider et alii (2014).

Description

The following description is excerpted from the descriptions of Medusagynaceae, Quiinaceae, and Ochnaceae s.s. in ''The Families and Genera of Vascular Plants'', with some information from other sources, as indicated.

Mostly shrubs and small trees, plus a few trees of moderate size, and in ''Sauvagesia'', a few herbs.

Leaves simple, except pinnately compound in ''Krukoviella'', and in Quiinoideae, often pinnately lobed or compound on juvenile plants. Pinnate leaves are typical of ''Rhytidanthera''. The leaves are often coriaceous and conspicuously serrate. Stipules present, except ''Medusagyne''.

Venation often scalariform (ladder-like) in appearance, with parallel and closely spaced secondary and tertiary veins. Petioles absent or short, sometimes resembling a pulvinus.

Unisexual flowers are common in ''Medusagyne'' and in Quiinoideae (except ''Froesia''), but restricted to a clade of three genera in Ochnoideae. Unisexual flowers are found in ''Schuurmansia'', ''Schuurmansiella'', and ''Euthemis''. The flowers are always unisexual in ''Schuurmansiella''.

In polygamous species, flowers have been assessed as bisexual on the basis of morphology only. Pollen produced by apparently hermaphroditic flowers has, in a few cases, been found to be inaperturate, rendering the flower functionally female.

Sepals 3 to 5, often unequal, sometimes accrescent.

Petals 4 or 5, or rarely 3, 6, 7, or 8, often contort, free or fuzed at base only, sometimes reflexed over the sepals.

Fertile stamens 5 to 10 or numerous, rarely one. Filaments sometimes persistent, sometimes narrowed near the anthers.

Anthers basifixed or slightly dorsifixed, usually dehiscing by one or two apical or subapical pores, sometimes latrorsely by longitudinal slits. In ''Medusagyne'' and Quiinoideae, an enlarged septum separates the thecae.

Staminodes often present, free or connate, sometimes petaloid, sometimes enveloping the fertile stamens.

Nectar not produced. Flowers usually buzz-pollinated.

Ovary superior, longitudinally ribbed in ''Medusagyne'' and Quiinoideae; unribbed in Ochnoideae. Carpels completely fused or nearly separate; 2-15, or up to 25 in ''Medusagyne''. Style apical or gynobasic.

Fruit sometimes winged; rarely a nut or drupe, often berry-like; usually a septicidal capsule, or else the ovary separating to form blackish drupelets on a usually reddish, accrescent receptacle.

Seeds albuminous or exalbuminous, winged or not. The seed coat often includes a layer of cristarque cells. These are sclereids, each containing calcium oxalate crystals in the form of a druse.

Classification

Until near the end of the 20th century, Ochnaceae was regarded as a rather odd family, difficult to place with a high degree of certainty. Even into the 21st century, some authors treated the genus '' Strasburgeria'' as the closest relative of Ochnaceae, and some even placed it within the family.Armen L. Takhtajan (Takhtadzhian). ''Flowering Plants'' second edition (2009). Springer Science+Business Media. . . (See ''External links'' below). In the APG III system, ''Strasburgeria'' is grouped with '' Ixerba'' to form the family Strasburgeriaceae in the rosid order Crossosomatales.Peter F. Stevens (2001 onwards). "Ochnaceae" At: Angiosperm Phylogeny Website. At: Missouri Botanical Garden Website. (see ''External links'' below)

More rarely, the genus ''Diegodendron'' was thought to be close to ''Strasburgeria'' and Ochnaceae. Molecular phylogenetic studies have strongly supported the inclusion of ''Diegodendron'' in the rosid order Malvales, and it is sometimes treated as a monospecific family therein.Clemens Bayer. 2003. ''Diegodendron''. pages 175-177. In: Klaus Kubitzki (editor). 2003. ''The Families and Genera of Vascular Plants'' volume V. (volume editors: K. Kubitzki and C. Bayer). Springer-Verlag: Berlin, Heidelberg, Germany. (print) (eBook). . Otherwise, it has been placed in Bixaceae, though there is reason to suspect that it might be closer to Sphaerosepalaceae.

All of the genera mentioned above, as well as Ochnaceae, were long regarded as anamalous taxa of uncertain affinity. All had been placed, at one time or another, with Ochnaceae, near Theaceae, a family now included in the basal asterid order Ericales.

In 2012, an analysis of chloroplast DNA resolved Ochnaceae as sister to a group of five families known as the clusioids. This result had only weak bootstrap support. The clusioids were once regarded as a group of four families,Peter F. Stevens. 2007. "Clusiaceae – Guttiferae". pages 48-66. In: Klaus Kubitzki (editor). 2007. ''The Families and Genera of Vascular Plants'' volume IX. (volume editors: K. Kubitzki in collaboration with C. Bayer and P.F. Stevens). Springer-Verlag: Berlin, Heidelberg, Germany. () () . but Clusiaceae was divided in 2009 and the name Calophyllaceae was resurrected for one of the resulting segregates.Brad R. Ruhfel, Peter F. Stevens, and Charles C. Davis. 2013. "Combined morphological and molecular phylogeny of the clusioid clade (Malpighiales) and the placement of the ancient rosid macrofossil ''Paleoclusia''". ''International Journal of Plant Sciences'' 174(6):910–936. .

There are only a few morphological characters that unite the clusioids with Ochnaceae. The petal aestivation is often contort in the clusioids, and usually so in Ochnaceae. In both groups, the flowers usually bear numerous stamens, and in the ovary, the placentation is mostly axile. In the ovules, the nucellus is often thin, and the outer integument is usually thicker than the inner.Peter K. Endress, Charles C. Davis, and Merran L. Matthews. 2013. "Advances in the floral structural characterization of the major subclades
of Malpighiales, one of the largest orders of flowering plants". ''Annals of Botany'' 111(5): 969–985. . (See ''External links'' below).

Taxonomy

Ochnaceae is divided into three subfamilies: Medusagynoideae, Quiinoideae, and Ochnoideae.

A molecular phylogenetic study resolved Medusagynoideae and Quiinoideae as sister subfamilies, but this result had only weak statistical support. In both subfamilies, the flowers are polystemonous. Except for the genus ''Froesia'', many or all of the flowers are unisexual. The anthers contain a massive septum between the thecae that persists after anther dehiscence. The styles radiate outward from the ovary. At anthesis, the ovary is sculpted with longitudinal ribs.Merran L. Matthews, Maria do Carmo E. Amaral, and Peter K. Endress. 2012. "Comparative floral structure and systematics in Ochnaceae s.l. (Ochnaceae, Quiinaceae and Medusagynaceae; Malpighiales)". ''Botanical Journal of the Linnean Society '' 170(3):299-392. .

Medusagynoideae consists of a single species, ''Medusagyne oppositifolia''. It is endemic to the island of Mahé in the Seychelles.William C. Dickison. 1990. "The morphology and relationships of ''Medusagyne'' (Medusagynaceae)". ''Plant Systematics and Evolution'' 171(1-4):27-55.

Quiinoideae comprises about 48 species in four genera: ''Froesia'', ''Quiina'', ''Touroulia'', and ''Lacunaria''. It is restricted to tropical America.Julio V. Schneider and Georg Zizka. in press. "Quiinaceae". ''Flora Neotropica'' monograph ??. ''Froesia'' is distinct from the other three genera. Its flowers are always bisexual, and the fruit consists of three structures that resemble follicles, except that they are not quite entirely separate from each other.

The subfamily Ochnoideae was revised in 2014 as Ochnaceae sensu stricto. In that treatment, 27 genera were described. An additional genus, ''Neckia'', was resurrected in that same year, based on the results of a molecular phylogenetic study. Amaral and Bittrich (2014) divided Ochnoideae into three tribes: Luxemburgieae, Sauvagesieae, and Ochneae. The genus ''Testulea'' was included in the tribe Sauvagesieae. No subtribes were recognized.

In 2014, a second reclassification of Ochnoideae was published, based on cladistic analysis of DNA sequences. In that paper, Schneider et alii divided Ochnoideae into four tribes: Testuleeae, Luxemburgieae, Sauvagesieae, and Ochneae. Their circumscription of the tribes was the same as that of Amaral and Bittrich (2014), except that ''Testulea'' was removed from Sauvagesieae into its own tribe, Testuleeae. Inclusion of ''Testulea'' in Sauvagesieae renders that tribe paraphyletic over Luxemburgieae.

''Testulea'' consists of a single species, ''Testulea gabonensis'', that is endemic to Gabon. It is unique in Ochnoideae in that its leaves have a brochidodromous pattern of venation and its flowers are tetramerous. In addition, only one of the stamens is fertile. The others are modified into staminodes and united into a column for up to ²⁄₃ of their length.

The tribe Luxemburgieae consists of two genera: ''Philacra'' and ''Luxemburgia''. ''Philacra'' is native to Venezuela and northern Brazil. ''Luxemburgia'' is from Brazil.

The tribe Sauvagesieae is pantropical in distribution, and consists of 16 genera, most of them small. The largest, by far, is ''Sauvagesia'', with 38 species, 35 of which are restricted to the neotropics. ''Sauvagesia'' is heterogeneous, and might be paraphyletic, even with ''Neckia'' removed from it. Relationships in the tribe Sauvagesieae are not well understood, and for this reason, it has not been divided into subtribes.

The tribe Ochneae is found in most of the tropics, but is most abundant in Africa and tropical America. It is distinguished from the rest of Ochnoideae by the absorption of the endosperm before the seed reaches maturity. Its nine genera belong to three subtribes: Lophirinae, Elvasiinae, and Ochninae.

The subtribe Lophirinae consists of a single genus, ''Lophira''. It has two species, both confined to tropical Africa. It produces an unusual fruit, in which two of the sepals become greatly enlarged and form wings that facilitate seed distribution by wind.

The subtribe Elvasiinae consists of two genera, ''Perissocarpa'' and ''Elvasia'', both confined to the American tropics. ''Perissocarpa'' has never been sampled for a molecular phylogenetic study.

The subtribe Ochninae consists of six genera: ''Campylospermum, Ouratea, Idertia, Brackenridgea, Rhabdophyllum,'' and ''Ochna''. The largest of these, ''Ouratea'', is confined to the New World and contains all of the New World species in Ochninae. All of the genera in Ochneae appear to be monophyletic, as defined by Amaral and Bittrich (2014), but in one molecular phylogenetic study, ''Ouratea'' and ''Ochna'' received only weak bootstrap support in the maximum likelihood analysis. ''Idertia'' and ''Brackenridgea'' are probably sister genera, but no other relationships have been resolved among the genera of Ochninae.

Phylogeny

The phylogenetic tree below is adapted from the one that was published in 2014. Weakly supported nodes are collapsed to form polytomies. Maximum likelihood bootstrap support is > 75%, except where indicated. ''Perissocarpa'' and ''Indosinia'' have not been sampled for DNA. Their placement on the phylogenetic tree is based on anatomy and morphology only.

Evolution

The evolution of Ochnaceae has been unusual in that it has included two complete reversions to an actinomorphic floral symmetry and two nearly complete reversions to apocarpy, a condition in which the carpels are entirely separate. Actinomorphy and apocarpy are believed to be " primitive" character states in angiosperms.Douglas E. Soltis, Pamela S. Soltis, Peter K. Endress, and Mark W. Chase. 2005. ''Phylogeny and Evolution of Angiosperms''. Sinauer Associates, Inc.: Sunderland, MA, USA. . Secondary apocarpy is especially rare and has arisen most notably in Rosaceae, Apocynaceae, Sapindales, and Malvales.

The flowers are actinomorphic in ''Medusagyne'' and Quiinoideae, but in Ochnoideae, zygomorphy is the ancestral condition. In ''Testulea'', ''Philacra'', and ''Luxemburgia'', the flowers develop zygomorphically in the bud. But in the four basal clades of Sauvagesieae, comprising the genera ''Blastemanthus, Godoya, Rhytidanthera, Krukoviella, Cespedesia, Fleurydora, Poecilandra,'' and ''Wallacea'', the flowers develop actinomorphically in the bud, then become zygomorphic after opening by growth of certain parts of the flower. Such late zygomorphy is very rare in flowering plants. In the remaining, fifth clade of Sauvagesieae, comprising the genera ''Neckia, Schuurmansia, Schuurmansiella, Euthemis, Tyleria, Adenarake, Indosinia,'' and ''Sauvagesia'', the flowers remain actinomorphic after anthesis. In the tribe Ochneae, all of the species have actinomorphic flowers.

In the subtribe Ochninae, and in the genus ''Froesia'', the components of the ovary (carpels) are very shortly united at the base. Otherwise, the ovary in Ochnaceae is syncarpous, consisting of carpels that are completely fuzed.

In ''Medusagyne'' and Quiinoideae, many of the flowers are unisexual, except in ''Froesia'', where they are strictly hermaphrodite. In Ochnoideae, unisexual flowers are limited to a clade consisting of ''Schuurmansia'', ''Schuurmansiella'', and ''Euthemis''.

In ''Medusagyne'' and Quiinoideae, as in most angiosperms, the anthers open by longitudinal slits. In Ochnoideeae, the anther dehiscence is ancestrally poricidal, with several reversions to longitudinal slits. ''Testulea'', ''Philacra'', and ''Luxemburgia'' have anthers that open by apical pores. So do the three most basal clades of Sauvagesieae, namely ''Blastemanthus'', ''Fleurydora'', and a clade of four genera that have five carpels and many ovules per carpel (''Godoya, Rhytidanthera, Krukoviella'', and ''Cespedesia''). ''Poecilandra'' has poricidal anther dehiscence, but in its sister genus, ''Wallacea'', the anthers open by longitudinal slits.

In the rest of Sauvagesieae, anther dehiscence is various. In ''Schuurmansia'', ''Schuurmansiella'', and ''Adenarake'', the anther dehiscence is apically longicidal. This means that the longitudinal slit is short, and does not extend far from the apical end of the anther. In some species of ''Sauvagesia'', the anthers split lengthwise, but the entire androecium is wrapped in petaloid staminodes, so that the pollen can escape from only the apex of the anther. This is known as a poricidal system because it functions as if the anthers were truly poricidal.

In the tribe Ochneae, anther dehiscence by longitudinal slits is restricted to ''Brackenridgea'' and a few species of ''Ochna''.

''Testulea'' is peculiar in having only one fertile stamen. Its anther opens by an apical pore. The other stamens are modified into sterile staminodes that are fuzed into a column for up to ²⁄₃ of their length.

In the genera ''Froesia'' and ''Quiina'', and in the tribe Ochneae, the endosperm is completely absorbed early in seed development. It is not clear whether the presence or absence of endosperm is the ancestral state in Ochnaceae. For a long time, the subfamily Ochnoideae was divided into two groups based on this character alone. In such a classification, the group containing endosperm would be paraphyletic over Ochneae because it would contain ''Testulea'', ''Philacra'', and ''Luxemburgia''.

The number of ovules per carpel varies widely in Ochnaceae. ''Medusagyne'' and Quiinoideae have two ovules per carpel. In ''Testulea'' and in a clade of four genera in Sauvagesieae (''Godoya, Rhytidanthera, Krukoviella'', and ''Cespedesia''), the number of ovules is 100 to 200 per carpel. For the remainder of Sauvagesieae, except ''Euthemis'', and for ''Philacra'' and ''Luxemburgia'', the number of ovules per carpel ranges from four to 50. ''Euthemis'' has two ovules per carpel.

In Ochneae, ''Lophira'' has 4 to 50 ovules per carpel. In the subtribes Elvasiinae and Ochninae, the number of ovules per carpel is one.

History

The family Ochnaceae was erected by Augustin Pyramus de Candolle in 1811.Ochnaceae in International Plant Names Index. (see ''External links'' below).James L. Reveal. 2008 onward. "A Checklist of Family and Suprafamilial Names for Extant Vascular Plants." At: Home page of James L. Reveal and C. Rose Broome. (see ''External links'' below). At that time, he described ''Elvasia'', a new genus in the family, and he included three others: ''Ochna'', ''Walkera'', and ''Gomphia''.Augustin Pyramus de Candolle. 1811. ''Nouveau bulletin des sciences par la Société philomathique de Paris'' 2(40):208. (See ''External links'' below). ''Walkera'' was described by Johann Christian Daniel von Schreber in 1789, but is no longer recognized. Its type species was described as ''Gomphia serrata'' by Andrias Kanis in 1968,Andrias Kanis. 1968. "A revision of the Ochnaceae of the Indo-Pacific area". ''Blumea'' 16(1):1-83. but it is now placed in the genus ''Campylospermum''. ''Gomphia'' has long been a source of confusionWerner Greuter and Rosa Rankin Rodríguez. 2014. "A type for ''Gomphia'' (Ochnaceae) – once again". ''Taxon'' 63(5):1122-1123. and it was not recognized in the most recent revision of Ochnaceae.

''Godoya'' and ''Sauvagesia'' were known in 1811, when de Candolle erected the family Ochnaceae, but he placed them in other families. In his ''Prodromus'', he placed ''Godoya'' in the family that would later be known as Clusiaceae.Augustin Pyramus de Candolle. 1824. "Violaceae" (as Violarieae), pages 287-316; Clusiaceae (as Guttiferae), pages 557-564; "Ochnaceae", pages 735-738. In: ''Prodromus Systematis Naturalis Regni Vegetabilis'' volume 1. (See ''External links'' below). He regarded ''Lauradia'' (''Lavradia'') as separate from ''Sauvagesia'', and placed both of them in Violaceae. He added the genus ''Castela'' to Ochnaceae, but it is now part of Simaroubaceae.Joshua W. Clayton. 2011. "Simaroubaceae", pages 408-423. In: Klaus Kubitzki (editor). 2011. ''The Families and Genera of Vascular Plants'' volume X. Springer-Verlag: Berlin, Heidelberg, Germany. (print). (eBook). De Candolle believed that Simaroubaceae was closely related to Ochnaceae, but it is now placed in the order Sapindales. A few authors placed ''Godoya'', ''Sauvagesia'', and others in the family Sauvagesiaceae, until the beginning of the 21st century. Other authors, such as Adolf Engler, included them in Ochnaceae.

In 1874, Engler divided Ochnaceae into two groups, based on the absence or presence of endosperm in the mature seed.H.G. Adolf Engler. 1874. "Ueber Begrenzung und systematische Stellung der natürlichen Familie der Ochnaceae". ''Nova Acta Academieae Caesarieae Leopoldino - Carolinae Germanicae Naturae Curiosorum'' 37(2):1-28. (See ''External links'' below). The group without endosperm corresponds to de Candolle's concept of Ochnaceae, and to the modern tribe Ochneae. The group with endosperm is now known to be paraphyletic and consists of the tribes Testuleeae, Luxemburgieae, and Sauvagesieae. In 1876, in a flora of Brazil, Engler described many new species in Ochnaceae, especially in its largest genus, ''Ouratea''.H.G. Adolf Engler. 1876. "Ochnaceae" In: Carl F.P. von Martius and August Wilhelm Eichler (editors). ''Flora Brasiliensis'' 12(2):297-366. (See ''External links'' below). He described 85 species in ''Ouratea'', 17 of which he named as new species at that time. He also transferred 63 species to ''Ouratea'' from other genera.

The genera ''Quiina'' and ''Touroulia'' had been known since 1775, when they were described by Jean Baptiste Aublet,Jean Baptiste C.F. Aublet. 1775. ''Histoire des Plantes de la Guiane Françoise'': · · · . ''Quiina'': Supplement to volumes 1 and 2, page 19; ''Touroulia'': 1:492; ''Ouratea'': 1:397. (See ''External links'' below). and they were variously classified by 19th century taxonomists. Jacques Denys Choisy erected the family Quiinaceae (as Quiinacées) for them in 1849,Jacques Denys Choisy. 1849. ''Description des Guttiferes de l'Indie'', · · · :12. but he did not meet the requirements for valid publication of a botanical name. The name Quiinaceae was validated by Engler in Flora Brasiliensis in 1888.H.G. Adolf Engler. 1888. "Quiinaceae". pages 475-486 and tables 109-110. In: Carl F.P. von Martius and August Wilhelm Eichler (editors). ''Flora Brasiliensis'' 12(1). (See ''External links'' below).

The genus ''Medusagyne'' had been described by John Gilbert Baker in 1877, in a flora of Mauritius and the Seychelles,John Gilbert Baker. 1877. ''Flora of Mauritius and the Seychelles'': 16. L. Reeve & Co.: London, UK. (See ''External links'' below). but it was not until 1924 that it was segregated into its own monogeneric family.H.G.Adolf Engler and Ernest F. Gilg. 1924. ''Syllabus der Pflanzenfamilien: eine Übersicht · · ·'' neunte und zehnte auflage: 280. (Syllabus of the Plant Families: an overview · · · editions 9 and 10, page 280.)

In 1893, Ernest Friedrich Gilg covered Ochnaceae, and Adolf Engler covered Quiinaceae for the first edition of Die Natürlichen Pflanzenfamilien.Ernest F. Gilg. 1893. "Ochnaceae". pages 131-153. In: H.G. Adolf Engler and Karl A.E. Prantl (editors). 1895. ''Die Natürlichen Pflanzenfamilien'' 1st edition: Teil III, Abteilung 6 (volume 3, part 6). Verlag von Wilhelm Engelmann: Leipzig, Germany. (See ''External links'' below).H.G. Adolf Engler. 1893. "Quiinaceae". pages 165-167. In: H.G. Adolf Engler and Karl A.E. Prantl (editors). 1895. ''Die Natürlichen Pflanzenfamilien'' 1st edition: Teil III, Abteilung 6 (volume 3, part 6). Verlag von Wilhelm Engelmann: Leipzig, Germany. (See ''External links'' below). Engler wrote a description of ''Medusagyne'' in a supplement to the first edition of '' DNP'' in 1897.H.G. Adolf Engler. 1897. ''Medusagyne''. page 250. In: H.G. Adolf Engler. 1897. "Guttiferae". pages 247-250. In: H.G. Adolf Engler and Karl A.E. Prantl. 1897. ''Die Natürlichen Pflanzenfamilien: Nachträge zum II-IV Teil''. (The Natural Plant Families: First Supplement for volumes 2-4). (See ''External links'' below). He placed ''Medusagyne'' under the heading "Zweifelhafte, möglicherweise zu den Guttiferae gehörige Gattung" (Doubtful, possibly a genus belonging to Guttiferae). Guttiferae is an obsolete name for Clusiaceae.

In 1902, Philippe van Tieghem recognized six families in what is now the subfamily Ochnoideae.Philippe E.L. van Tieghem. 1902. "Sur les Ochnacées". ''Annales des Sciences Naturelles – botanique'', séries 8 16:161-416. These were Luxemburgiaceae, Sauvagesiaceae, Wallaceaceae, Euthemidaceae, Lophiraceae, and Ochnaceae. Three of these, (Wallaceaceae, Euthemidaceae, and Lophiraceae) were monogeneric and were erected by van Tieghem at that time. His Luxemburgiaceae included the basal clades of what is now the tribe Sauvagesieae. Van Tieghem named a great many genera in 1902, circumscribing them very narrowly. In what is now the subtribe Ochninae, he delineated 53 genera. The most recent revision of that group divides it into six genera.

In 1925, for the second edition of '' DNP'', Engler and Gilg expanded on their treatment of Quiinaceae and Ochnaceae, respectively, compared to what they had written in 1893.H.G. Adolf Engler. 1925. "Quiinaceae". pages 106-108. In: H.G. Adolf Engler and Karl A.E. Prantl (editors). 1925. ''Die Natürlichen Pflanzenfamilien: zweite auflage'' 21 Band. (The Natural Plant Families: 2nd edition, volume 21). Verlag von Wilhelm Engelmann: Leipzig, Germany.Ernest F. Gilg. 1925. "Ochnaceae". pages 53-87. In: H.G. Adolf Engler and Karl A.E. Prantl (editors). 1925. ''Die Natürlichen Pflanzenfamilien: zweite auflage'' 21 Band. (The Natural Plant Families: 2nd edition, volume 21). Verlag von Wilhelm Engelmann: Leipzig, Germany. Medusagynaceae was covered in the same volume of ''DNP'' by Adolf Engler and Hans Melchior.Hans Melchior. 1925. "Medusagynaceae". pages 50-52. In: H.G. Adolf Engler and Karl A.E. Prantl (editors). 1925. ''Die Natürlichen Pflanzenfamilien: zweite auflage'' 21 Band. (The Natural Plant Families: 2nd edition, volume 21). Verlag von Wilhelm Engelmann: Leipzig, Germany.

For the second edition of ''DNP'', Engler recognized two genera, ''Quiina'' and ''Touroulia'', in Quiinaceae. ''Lacunaria'' and ''Froesia'' were discovered later and named in 1925 and 1948, respectively.

In that same volume, Gilg divided his Ochnaceae (equivalent to modern Ochnoideae) into 21 genera, including ''Indovethia, Leitgebia, Vausagesia,'' and ''Lauradia'' (as ''Lavradia''), which are now regarded as synonyms of ''Sauvagesia''. Eight of the modern genera, (''Philacra, Krukoviella, Fleurydora, Tyleria, Adenarake, Indosinia, Perissocarpa,'' and ''Idertia'') consist of plants that had not been uncovered by botanical exploration at that time. Gilg placed ''Rhytidanthera'' in synonymy under ''Godoya'', but ''Rhytidanthera'' is accepted today. He included ''Campylospermum'' and ''Rhabdophyllum'' in ''Gomphia'' and placed ''Gomphia'' in synonymy under ''Ouratea''. In his revision of Ochnaceae, Gilg provided a summary of van Tieghem's classification, as well as his own. Three of van Tieghem's genera, (''Campylospermum'', ''Rhabdophyllum'', and ''Rhytidanthera'') are still recognized today.

In 1968, Andrias Kanis published a paper that greatly influenced subsequent work in Ochnaceae, right up to the 2014 revision. Claude H.L. Sastre named many new species in Ochnaceae, in several papers from 1970 to 2003.

In 1991, a cladistic analysis was published for Ochnaceae.Maria do Carmo E. Amaral. 1991. "Phylogenetische Systematik der Ochnaceae". ''Botanische Jahrbücher für Systematik, Pflanzengeschichte, und Pflanzengeographie'' 113(1):105-196. In that same year, ''Neckia'', a 28th genus for Ochnoideae, was resurrected in a molecular phylogenetic study based on four chloroplast DNA loci, and nuclear ribosomal ITS. Seventy-nine species of Ochnaceae were sampled and a new classification was presented. Also, ''Testulea'' was removed from the tribe Sauvagesieae, and placed in the monotypic tribe Testuleeae.

References

External links

Xi2012Pubmed CentralNCBI (National Center for Biotechnology Information)

''Cespedesia''''Ochna''Sauvagesia

Endress (2011)land plant evolutionary timelineArchive''American Journal of Botany''


Eocene macroflora from Mississippi (print)back issues by coverBrowse Articles''Palaeontologia Electronica''/content

OchnaceaeangiospermsBrowseThe Plant List

Ochnaceae

Ochnoideae distribution mapMissouri Botanical Garden

the clusioid clade and ''Paleoclusia''volume 174, issue 6IJPS

floral structure in MalpighialesArchive''Annals of Botany''

OchnaceaePlant NamesIPNI


ttp://www.plantsystematics.org/reveal/ Home page of James L. Reveal and C. Rose Broome
''Gomphia''''Taxon''

''Flora of Mauritius and the Seychelles''Archive.org
*
BHL
(Biodiversity Heritage Library):
:
Ochnaceae, page 208Issue 40View BookNouveau bulletin des sciences par la Société philomathique de Paris, tome 2
:
''Lavradia''''Sauvagesia''''Godoya''''Castela''View Book''Prodromus'' v.1 (1824)
:
Ochnaceae in volume 37, number 2volume 37Nova acta Academiae Caesareae Leopoldino-Carolinae Germanicae Naturae Curiosorum
:
Ochnaceaevolume12 part 2''Flora Brasiliensis''
:
''Quiina''Supplement to volume 2volume 2''Touroulia''''Ouratea''volume 1''Histoire des plantes de la Guiane Françoise''
:
Quiinaceaevolume12 part 1

:
OchnaceaeQuiinaceaevolume 3 part 6Die Natürlichen Pflanzenfamilien
:
''Medusagyne''Nachträge (Supplement) Teil 2-4
Die Natürlichen Pflanzenfamilien
:
Sur les OchnacéesSeries 8, volume 16Annales des sciences naturelles, huitieme série – Botanique.

Ochnaceae (Search Exact)
At:Names
At:Tropicos
At:Science and Conservation
At:Missouri Botanical Garden


i
L. Watson and M.J. Dallwitz (1992 onwards). The families of flowering plants: descriptions, illustrations, identification, information retrieval.
http://delta-intkey.com

List of genera in OchnaceaeAbout the ChecklistWorld Checklist of Selected Plant Families





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OchnaceaeMalpighialesMagnoliopsidaTracheophytaPlantsGlobal Species

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Malpighialesrosidscore eudicots (Gunneridae)eudicotsangiospermsSpermatopsidaembryophytesgreen plantseukaryoteslife on earth (root of the tree) Tree of Life Web Project (ToL)

Complete list of generaOchnaceae
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At:Families and Genera in GRIN
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OchnaceaeMalpighialesfabidsrosidsPentapetalaeGunneridaeeudicotyledonsMesangiospermaeMagnoliophyta (flowering plants)SpermatophytaEuphyllophytaTracheophytaEmbryophytaStreptophytinaStreptophytaViridiplantaeEukaryotaTaxonomyUniProt

OchnaceaeMalpighialesfabidsrosidsPentapetalaeGunneridaeeudicotyledonsMesangiospermaeMagnoliophyta (flowering plantsEukaryotaTaxonomy BrowserTaxonomy DatabaseTaxonomyNCBI (National Center for Biotechnology Information)

{{Taxonbar, from=Q133319

Malpighiales families