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== Taxonomy == Saxifragales is a relatively small [[angiosperm]] [[Order (biology)|order]], having only 15 [[Families (biology)|families]], about 100 [[genera]] and about 2,470 [[species]].{{sfn|Soltis et al|2013}} === History === [[File:Lindley Saxifragaceae.png|thumb|[[John Lindley]]'s description of Saxifragales 1853|alt=First page of Lindley's Saxifragales from 1853]] Saxifragales was first described in 1820 by [[Berchtold and Presl]] as a group of plants, Saxifrageae, with five genera, including ''[[Saxifraga]]'', lending their names as the [[botanical authority]] (Bercht. & J.Presl).{{sfn|Berchtold|Presl|1820|loc=pp. 259–60}} At times, that authority has also been given to [[Barthélemy Charles Joseph Dumortier|Dumortier]], due to a later publication (1829). Dumortier first used the word Saxifragaceae.{{sfn|Dumortier|1829|loc=p. 38}} By the time of [[John Lindley]]'s ''The Vegetable Kingdom'' (1853), the term Saxifragales was in use, which Lindley called an Alliance, containing five families.{{sfn|Lindley|1853|loc=pp. 566–575}} Later, the Saxifragales were placed in the [[angiosperm]] [[Class (biology)|class]] [[Dicotyledons]], also called [[Magnoliopsida]].{{sfn|Singh|2004}} === Phylogeny === The [[Order (biology)|order]] Saxifragales has undergone considerable revision in both placement and composition, since the use of [[molecular phylogenetics]], and the use of the modern [[Angiosperm Phylogeny Group]] (APG) classification.{{sfn|APG IV|2016}}{{sfn|Cole et al|2019}} They are identified as a strongly [[monophyletic]] group.{{sfn|Kubitzki|2007a}} In the initial APG publication (1998), the Saxifragales were identified within the [[core eudicots]] [[clade]] but its relationship to other clades was uncertain. The core eudicots consist of the order [[Gunnerales]] and a large clade of [[Pentapetalae]] (so named for having a [[synapomorphy]] of [[pentamerous]] (5 part) perianths), the latter representing about 70% of all angiosperms, with eight major lineages.{{sfn|Moore et al|2010}}{{sfn|Zeng et al|2017}} Later (2003), the order was described as "one of the major surprises of molecular phylogenetic analyses of the angiosperms", having elements previously placed in three or four separate subclasses based on morphology.{{sfn|APG II|2003}}{{sfn|Soltis et al|2013}} This was eventually resolved in the third APG system (2009), placing Saxifragales as a [[sister group]] to the [[rosids]] (Rosidae), within the Pentapetalae clade.{{sfn|Burleigh et al|2009}}{{sfn|Wang et al|2009}}{{sfn|APG III|2009}} This large combination has subsequently been given the name [[superrosids]] (Superrosidae), representing part of an early diversification of the [[angiosperms]].{{sfn|Soltis et al|2013}}{{sfn|APG IV|2016}}{{sfn|Soltis et al|2018}} Among the rosids, they share a number of similarities with the [[Rosales]], particularly [[Rosaceae]], including a hypanthium, five part flowers and free floral parts.{{sfn|Byng|2014}} As circumscribed, Saxifragales account for 1.3% of eudicot diversity.{{sfn|Stevens|2019}} {{cladogram | title=Cladogram of Saxifragales relationships among core eudicots{{sfn|APG IV|2016}}{{sfn|Cole et al|2019}}{{sfn|Tank et al|2015}} | caption=Numbers indicate [[Divergence time estimation|divergence times]] in Myr | align=left | cladogram= {{clade |style="text-align:left; padding:2.5px; background:#eef"; width:500px; | sublabel1=130 | label1=[[Core eudicots]] | 1={{clade |sublabel1=126 |1=[[Gunnerales]] |sublabel2=128 |label2=[[Pentapetalae]] |2={{clade |label1= |1=[[superasterids]] |2={{clade |sublabel1=125 |1=[[Dilleniales]] |sublabel2=124 |label2=[[superrosids]] |2={{clade |sublabel1=116 |1=[[rosids]] |sublabel2=108 |2='''Saxifragales''' }} }} }} }} }} }} {{Clear}} === Biogeography and evolution === [[Genetic divergence|Diversification]] among Saxifragales was rapid, with the extensive [[fossil]] record{{sfn|Hermsen et al|2006}}{{sfn|Hermsen et al|2003}}{{sfn|Pigg et al|2004}}{{sfn|Hernández-Castillo|Cevallos-Ferriz|1999}}{{sfn|Crane|1989}}{{sfn|Endress|1989}} indicating that the order was more [[Biodiversity|diverse]] and more [[Cosmopolitan distribution|widespread]] than an examination of the [[Extant taxon|extant]] members suggests, with considerable [[phenotypic]] diversity occurring early.{{sfn|Casas et al|2016}} The earliest fossil evidence is found in the [[Turonian]]-[[Campanian]] (late Cretaceous), suggesting a minimum age of 89.5 [[Myr]]. However, molecular [[divergence time estimation]] suggest an earlier time of 102–108 Myr, into the early Cretaceous, for the [[Crown group|crown]] and [[stem group]]s respectively. Within the order Saxifragales, the molecular data imply a very rapid initial diversification time of about 6–8 Myr, between 112 and 120 Myr, with major lineages appearing within 3–6 Myr.{{sfn|Jian et al|2008}}{{sfn|Soltis et al|2013}}{{sfn|Li et al|2019}} The ancestral state appears to be woody, as in Peridiscaceae and the woody clade, but is also ancestral to Grossulariaceae. A number of independent transitions to a herbaceous habit occurred in the ancestors of Crassulaceae, Saxifragaceae and the base of the Haloragaceae-Penthoraceae clade (the other two families in Haloragaceae ''s.l.'' remaining woody), while other taxa reverted to a woody habit, especially Crassulaceae. Most of Saxifragales have a superior [[Ovary (botany)|ovary]], but some families show frequent transition with inferior or subinferior position, particularly Saxifragaceae and to a lesser extent Hamamelidaceae. Almost all Grossulariaceae have an inferior ovary. The ancestral [[carpel]] number is two, with transition to higher numbers, such as four in Haloragaceae ''s.l.'' and Peridiscaceae with five in Penthoraceae. The ancestral carpel number for Crassulaceae is five, decreasing to four in ''[[Kalanchoe]]'', where it is synapomorphic for the genus, though the most frequent transition in this family is 6–10, but only where [[stamen]] number is increased above five. Some [[Macaronesian]] taxa (Aeonieae) have 8–12, with up to 32 carpels for ''[[Aeonium]]''.{{sfn|Soltis et al|2013}} The ancestral petal number is five, with three major transitions; 5 to 0, 5 to 4, 5 to 6–10. Increased petal number is seen in Paeoniaceae and Crassulaceae, particularly where stamen number is also increased. Cercidiphyllum + Daphniphyllum, Chrysosplenium and ''[[Altingia]]'' are examples of the complete loss of petals. The ancestral stamen:petal ratio is 1, with transitions characterising several clades, e.g. Paeonicaceae+woody clade >2, Crassulaceae 2 (but ''[[Crassula]]'' 1). Overall there has been a decrease over evolution, but independent of a decrease in petal number, so that it is the stamen number that has decreased.{{sfn|Soltis et al|2013}} The ancestral habitat appears to be forests, followed by early diversification into desert and aquatic habitats, with shrubland the most recent colonization.{{sfn|Casas et al|2016}} Species diversification was rapid following a transition from a warmer, wetter Earth in the [[Eocene]] (56–40 Myr) to early [[Miocene]] (23–16 Myr), to the cooler drier conditions of the mid-Miocene (16–12 Myr). However, this appears to not have coincided with ecological and phenotypic evolution, which are themselves correlated. There is a clear lag, whereby increase in species diversification was followed later by increases in niche and phenotypic lability.{{sfn|Folk et al|2019}} === Subdivision === The first APG classification (1998) placed 13 families within the order Saxifragales:{{sfn|APG I|1998}} {{div col|colwidth=160px}} * [[Altingiaceae]] * [[Cercidiphyllaceae]] * [[Crassulaceae]] * [[Daphniphyllaceae]] * [[Grossulariaceae]] * [[Haloragaceae]] * [[Hamamelidaceae]] * [[Iteaceae]] * [[Paeoniaceae]] * [[Penthoraceae]] * [[Pterostemonaceae]] * [[Saxifragaceae]] * [[Tetracarpaeaceae]] {{div col end}} This was subsequently revised to 15, in the fourth version (2016).{{sfn|APG IV|2016}} The Saxifragales families have been grouped into a number of informally named suprafamilial subclades, with the exception of the [[basal split]] of Peridiscaceae, which thus forms a [[sister group]] with the rest of Saxifragales. The two major ones are (Paeoniaceae + the woody clade of primarily woody families) and the "core" Saxifragales (i.e. the primarily herbaceous families), with the latter subdivided into two further subclades, (Haloragaceae ''sensu lato'' + Crassulaceae) and the Saxifragaceae alliance.{{sfn|Soltis et al|2013}} In the clade Haloragaceae ''sensu lato'' ''(s.l.)'' + Crassulaceae the genera constituting Haloragaceae ''s.l.'' are all small, and APG II (2003) proposed merging them into a single larger Haloragaceae ''s.l.'', but transferred ''[[Aphanopetalum]]'' from [[Cunoniaceae]] to this group.{{sfn|APG II|2003}} The Saxifragaceae alliance represents Saxifragaceae together with a number of woody members of the traditional Saxifragaceae ''sensu'' Engler (1930).{{sfn|Engler|1930}} Within this, APG II (2003) proposed placing the two species of ''[[Pterostemon]]'' that constitute Pterostemonaceae within [[Iteaceae]], and all subsequent versions have maintained this practice.{{sfn|APG II|2003}} Thus Saxifragales ''sensu'' APG II consisted of only 10 families. The third version (2009) added [[Peridiscaceae]] (from [[Malpighiales]]), as sister to all other families, but re-expanded Haloragaceae to provide for a narrower circumscription, Haloragaceae ''sensu stricto'' (''s.s.''), to give a total of 14 families. APG IV (2016) added the parasitic family Cynomoriaceae to provide a total of 15 families, although its placement within the order remained unclear.{{sfn|Bellot et al|2016}}{{sfn|APG IV|2016}} Of the 15 families included in APG IV, the basal divergence Peridiscaceae underwent radical shifting and recircumscription from 2003 to 2009. Originally, it consisted of two closely related genera, ''Peridiscus'' and ''Whittonia''. The [[APG II]] system placed the family in [[Malpighiales]], based on a [[DNA sequence]] for the [[RuBisCO|''rbcL'']] [[gene]] from ''Whittonia''. This sequence turned out to be not from ''Whittonia'', but from other plants whose [[DNA]] had contaminated the sample.{{sfn|Davis|Chase|2004}} After placement in Saxifragales, it was expanded to include ''[[Soyauxia]]'' in 2007,{{sfn|Soltis et al|2007}} and ''[[Medusandra]]'' in 2009.{{sfn|Wurdack|Davis|2009}} In the first of the subclades of the remaining Saxifragales, Paeoniaceae possesses many [[autapomorphy|unique]] features and its taxonomic position was controversial for a long time,{{sfn|Tamura|2007}} and ''[[Peony|Paeonia]]'' was placed in [[Ranunculales]], close to ''[[Glaucidium (plant)|Glaucidium]]'',{{sfn|Mabberley|2008}}{{sfn|Halda|Waddick|2010}} prior to transfer to Saxifragales as sister to the woody clade.{{sfn|Jian et al|2008}}{{sfn|Wang et al |2009a}} In the woody clade, the genus ''[[Liquidambar]]'' was included in Hamamelidaceae until [[molecular phylogenetic]] studies showed that its inclusion might make Hamamelidaceae [[Paraphyly|paraphyletic]], and was segregated as a separate monotypic family, Altingiaceae in 2008.{{sfn|Jian et al|2008}} Cercidiphyllaceae was for a long time associated with Hamamelidaceae and [[Trochodendraceae]] and was often thought to be closer to the latter,{{sfn|Endress|1986}} which is now in the basal [[eudicot]] order [[Trochodendrales]].{{sfn|Worberg et al|2007}} ''Daphniphyllum'' was always thought to have an anomalous combination of characters{{sfn|Tseng-Chieng|1965}}{{sfn|Tseng-Chieng|1965}} and was placed in several different orders before molecular phylogenetic analysis showed it to belong to Saxifragales.{{sfn|Kubitzki|2007b}} In the core Saxifragales, Crassulaceae{{sfn|Thiede |Eggli|2007}} and Tetracarpaeaceae{{sfn|Hils et al|1988}} have been associated with Saxifragaceae, while ''Penthorum'' has been associated both with Crassulaceae and Saxifragaceae,{{sfn|Thiede|2007}} before being placed here. ''[[Aphanopetalum]]'' was often placed in [[Cunoniaceae]], a family in [[Oxalidales]], even though there were good reasons to put it in Saxifragales,{{sfn|Dickison et al|1994}} and it was subsequently transferred.{{sfn|Bradford|Fortune-Hopkins|Barnes|2004}} Haloragaceae was included in [[Myrtales]],{{sfn|Kubitzki|2007c}} before being placed in Saxifragales.{{sfn|Moody | Les |2007}} The other "core" group, the Saxifragaceae alliance comprises four families: Pterostemonaceae, Iteaceae, Grossulariaceae, and Saxifragaceae,{{sfn|Jian et al|2008}} which have long been known to be related to each other, but the [[Circumscription (taxonomy)|circumscription]] of Saxifragaceae has been much reduced and Pterostemonaceae submerged as ''[[Pterostemon]]'' in Iteaceae.{{sfn|Soltis et al|2001}} Most of the families are [[monogeneric]]. ''Choristylis'' is now considered a synonym of [[Itea (plant)|''Itea'']], but the addition of ''[[Pterostemon]]'', gives Iteaceae two genera.{{sfn|Kubitzki|2007d}} ''[[Liquidambar]]'' and ''[[Semiliquidambar]]'' are also submerged into ''[[Altingia]]'', making Altingiaceae monogeneric.{{sfn|Ickert-Bond|Wen|2006}}{{sfn|Ickert-Bond |Wen|2013}} About 95% of the species are in five families: [[Crassulaceae]] (1400), [[Saxifragaceae]] (500), [[Grossulariaceae]] (150–200), [[Haloragaceae]] (150), and [[Hamamelidaceae]] (100).{{sfn|Stevens|2019}}{{sfn|Jian et al|2008}}{{sfn|Kubitzki|2007a|pp=15–18}} The relationships of the Saxifragales families to each other is shown in the following [[cladogram]]. The [[phylogeny]] in this cladogram still has some uncertainty as to the exact relationships, and the phylogenetic tree is subject to further revision.{{sfn|Dong et al|2018}}{{sfn|Ding et al|2019}} [[Cynomoriaceae]], previously placed in [[Santales]] or [[Rosales]] is included in Saxifragales, but unplaced within it. Li et al. (2019) have slightly different relationships, and also place Cynomoriaceae as the first branch in the Crassulaceae+Haloragaceae ''s.l.'' tree, i.e. as sister to those two families.{{sfn|Li et al|2019}} The number of genera in each family is shown in parentheses: {{cladogram |title= Cladogram of Saxifragales families{{sfn|APG IV|2016}}{{sfn|Stevens|2019}}{{sfn|Jian et al|2008}} |caption=100% [[maximum likelihood]] [[Bootstrapping (statistics)|bootstrap support]] except where labeled with bootstrap percentage<br/>Monogeneric families are represented by genus names, otherwise the number of genera is in (parentheses)<br/>[[Cynomorium]] (Cynomoriaceae) remains unplaced within this tree |align=left |cladogram= {{clade | style=font-size:100%;line-height:120%;width:700px; |label1='''Saxifragales''' |1={{clade |1=[[Peridiscaceae]] (4) |label2=97 |2={{clade |label1= |1={{clade |1=''[[Peony|Paeonia]]'' (Paeoniaceae) |label2=woody clade |2={{clade |1=''[[Liquidambar]]'' (Altingiaceae) |label2=69 |2={{clade |label1=98 |1=[[Hamamelidaceae]] (27) |label2=95 |2={{clade |1=''[[Cercidiphyllum]]'' (Cercidiphyllaceae) |2=''[[Daphniphyllum]]'' (Daphniphyllaceae) }} }} }} }} |label2=core Saxifragales |2={{clade |1={{clade |1=[[Crassulaceae]] (34) |label2=Haloragaceae''s.l.'' |2={{clade |1=''[[Aphanopetalum]]'' (Aphanopetalaceae) |2={{clade |1=''[[Tetracarpaea]]'' (Tetracarpaeaceae) |2={{clade |1=''[[Penthorum]]'' (Penthoraceae) |2=[[Haloragaceae]] ''s.s.'' (8) }} }} }} }} |label2=Saxifragaceae alliance |2={{clade |1={{clade |1=[[Iteaceae]] (including [[Pterostemonaceae]]) (2) }} |2={{clade |1=''[[Ribes]]'' (Grossulariaceae) |2=[[Saxifragaceae]] (33) }} }} }} }} }} }} }} {{Clear}} === Families === {| |[[File:Peridiscus lucidus (Hooker).png|upright|thumb|130px|left|''[[Peridiscus|Peridiscus lucidus]]''|alt=Botanical illustration of Peridiscus lucidus]] ==== Peridiscaceae ==== {{Main|Peridiscaceae}} The Peridiscaceae (Ringflower family) are a small tropical family of 4 genera and 11–12 species of small trees and shrubs found in the [[Guiana Shield]] of S America (2 genera, one of which, ''Whittonia'', is thought to be [[extinct]]) and West and Central Africa (2 genera). The majority of species occur in the African genus ''[[Soyauxia]]''. The name comes from the [[Ancient greek language|Greek]], ''peri'' (around) ''discos'' (ring).{{sfn|Berry|2017}}{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Bánáti bazsarózsák a Somos-hegyen.jpg|thumb|130px|left|''[[Paeonia officinalis]]''|alt=Paeonia officinalis growing in Hungary]] ==== Paeoniaceae ==== {{Main|Paeoniaceae}} The Paeoniaceae (Peony family) consist of a single genus (''Paeonia'') with about 33 species of perennial herbs and small shrubs with showy flowers, found from the Mediterranean to Japan, but two species occur in western N America. They are commercially important as popular garden ornamentals, cultivated since antiquity, and have been used medicinally. The herbaceous varieties are derived from ''[[Paeonia lactiflora|P. lactiflora]]'', while the shrubs are derived from ''[[Paeonia suffruticosa|P. suffruticosa]]'' (tree peony), both Asian species. The botanical name comes from its Greek name, ''paionia'', named in turn for the God [[Pan (god)|Pan]].{{sfn|Berry|2017}}{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Liquidambar. Western Way, Exeter - geograph.org.uk - 1058382.jpg|thumb|130px|left|''[[Liquidambar styraciflua]]''|alt=Liquidambar styraciflua tree]] ==== Altingiaceae ==== {{Main|Altingiaceae}} The Altingiaceae (Sweetgum family) consist of a single genus (''[[Liquidambar]]'') with 15 species of trees with unisexual flowers found in Eurasia, but with one species in North and Central America, ''[[Liquidambar styraciflua]]'' (American sweetgum). ''Liquidambar'' is used for its [[resin]] and timber, as well being ornamental trees. The nominative genus and family are named after [[Willem Alting]], and Liquidambar for liquid ''ambar'', Arabic for the resin.{{sfn|Berry|2017}}{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Hamamelis virginiana FlowersLeaves BotGardBln0906.JPG|thumb|130px|left|''[[Hamamelis virginiana]]''|alt=Branch of Hamamelis virginiana showing flowers and leaves]] ==== Hamamelidaceae ==== {{Main|Hamamelidaceae}} The Hamamelidaceae (Witch-hazel family) consists of trees and shrubs with a widespread distribution, but main centres in East Asia and Malaysia. They are found in wet woodlands and forested slopes. The family has 26 genera and about 80–100 species, in five subfamilies, of which the nominative, Hamamelidoideae, contains over 75% of the genera. The species have uses as medicaments, timber and ornamental plants for their flowers, such as ''[[Hamamelis]]'' (witch hazel) or leaves, such as ''[[Parrotia persica]]'' (Persian ironwood). The family and nominative genus is named for the Greek ''hamamelis'', the wych elm.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Cercidiphyllum japonicum (arboretum Aubonne).JPG|thumb|130px|left|''[[Cercidiphyllum japonicum]]''|alt=Cercidiphyllum japonicum]] ==== Cercidiphyllaceae ==== {{Main|Cercidiphyllaceae}} The Cercidiphyllaceae (Caramel-tree family) are a small family of deciduous trees found in China and Japan, with a single genus, ''Cercidiphyllum'' and two species, ''[[Cercidiphyllum japonicum|C. japonicum]]'' and ''[[Cercidiphyllum magnificum|C. magnificum]]''. The trees are valued for their wood (''katsura'') and as ornamentals. ''[[Cercidiphyllum japonicum|C. japonicum]]'' is the largest deciduous tree in Japan. The name is derived from the Greek words ''kerkis'' ([[Poplar (botany)|poplar]]) and ''fyllon'' (leaf), from a supposed similarity in leaves.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Daphniphyllum macropodum 120502-1.jpg|thumb|130px|left|''[[Daphniphyllum macropodum]]''|alt=Daphniphyllum macropodum]] ==== Daphniphyllaceae ==== {{Main|Daphniphyllaceae}} The Daphniphyllaceae (Laurel-leaf family) consist of a single genus, ''Daphniphyllum'', with about 30 species. They are evergreen unisexual trees and shrubs distributed in SE Asia and the Solomon Islands. The dried leaves of ''[[Daphniphyllum macropodum]]''{{efn|''D. humile'' is a synonym of the accepted ''D. macropodum''}} have been used for smoking in Japan and Siberia. The name is derived from the Greek words ''dafne'' (laurel) and ''fyllon'' (leaf), from a supposed resemblance to the leaves of the former (''[[Laurus nobilis]]'').{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Crassula perfoliata minor 1c.JPG|thumb|130px|left|''[[Crassula perfoliata]]''|alt=Crassula perfoliata]] ==== Crassulaceae ==== {{Main|Crassulaceae}} The Crassulaceae (Orpine and Stonecrop family) are a medium size diverse and cosmopolitan family, that form the largest family within Saxifragales. They are mainly [[succulent]], rarely aquatic, with a specialised form of [[photosynthesis]] ([[Crassulacean Acid Metabolism]]). Genera vary from 7 to 35, depending on the circumscription of the large genus ''[[Sedum]]'', and there are about 1,400 species. Uses are diverse, including spices, medicaments and roof coverings as well as ornamental [[rock garden]] and household plants such as the S African ''[[Crassula ovata]]'', the jade or money plant. The name is derived from the Latin, ''crassus'' (thick), referring to the fleshy leaves.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Aphanopetalumresinosumgdn.jpg|thumb|130px|left|''[[Aphanopetalum resinosum]]''|alt= Aphanopetalum resinosum vine]] ==== Aphanopetalaceae ==== {{Main|Aphanopetalaceae}} The Aphanopetalaceae (Gum-vine family) consists of a single genus of Australian climbing shrubs, ''Aphanopetalum'', which has two species, ''[[Aphanopetalum clematideum|A. clematidium]]'' (SW Australia) and ''[[Aphanopetalum resinosum|A. resinosum]]'' (Queensland, NSW). The name is derived from the Greek words ''afanos'' (inconspicuous) and ''petalon'' (petal).{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}} {| |[[File:Tetracarpaea.jpg|thumb|130px|left|''[[Tetracarpaea tasmannica]]''|alt= Flowers of Tetracarpaea tasmannica]] ==== Tetracarpaeaceae ==== {{Main|Tetracarpaeaceae}} The Tetracarpaeaceae (Delicate-laurel family) is a very small evergreen Australian shrub family with a single genus, ''Tetracarpaea'' and a single species, ''T. tasmannica'', confined to subalpine Tasmania. The name is derived from the Greek words ''tetra'' (four) and ''carpos'' (fruit), referring to the ovaries which have four carpels.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Penthorum sedoides 002.JPG|thumb|130px|left|''[[Penthorum sedoides]]''|alt= Flowers of Penthorum sedoides]] ==== Penthoraceae ==== {{Main|Penthoraceae}} The Penthoraceae (Ditch-stonecrop family) is a very small family of rhizomatous perennial herbs found in eastern N America and E Asia, in mainly wet environments. It consists of a single genus, ''Penthorum'' with two species, ''[[Penthorum sedoides|P. sedoides]]'' in N America and ''[[Penthorum chinense|P. chinense]]'' from Siberia to Thailand. ''P. sedoides'' is used in aquaria and water gardens.{{sfn|Les|2017}} The name is derived from the Greek word ''pente'' (five) referring to the five-part fruit.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Haloragis erecta 2007-06-02 (flower).jpg|thumb|130px|left|''[[Haloragis erecta ]]''|alt= Flowers and leaves of Haloragis erecta ]] ==== Haloragaceae ==== {{Main|Haloragaceae }} The Haloragaceae (Water-milfoil family) is a small family of trees, shrubs, perennial, annual terrestrial, marsh and aquatic herbs with global distribution, but especially Australia. It consists of 9–11 genera and about 145 species. The largest genus is ''[[Gonocarpus]]'' with about 40 species. The major horticultural genus is ''[[Myriophyllum]]'' (watermilfoil) whose species are valued as aquaria and pond plants but may escape and naturalise, becoming invasive. Some cultivars of ''[[Haloragis]]'' are valued as ornamentals.{{sfn|Les|2017}} Only one genus, ''[[Haloragodendron]]'', is a shrub and is confined to S Australia. The family and nominative genus, ''Haloragis'' are named from the Greek words ''halas'' (salt) and ''rhoges'' (berries).{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Itea virginica 2zz.jpg|thumb|130px|left|''[[Itea virginica]]''|alt= Itea virginica plant ]] ==== Iteaceae ==== {{Main|Iteaceae }} The Iteaceae (Sweetspire family) is a widespread small family of trees and shrubs, with 2 genera, and 18–21 species, found in tropical to northern temperate regions. The larger genus, ''[[Itea (plant)|Itea]]'' (c. 16 spp.) is more widespread, from the Himalayas to Japan and western [[Malesia]] and one species in eastern N America (''[[Itea virginica|I. virginica]]'') whereas ''[[Pterostemon]]'' (c. 2 spp) is confined to [[Oaxaca]], Mexico. ''I. virginica'' and ''[[Itea ilicifolia|I. ilicifolia]]'', from China, are valued as ornamental shrubs. The name is derived from the Greek word ''itea'' (willow) for its rapid growth and similar leaf form.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Ribes_rubrum.jpg|thumb|130px|left|''[[Ribes rubrum]]''|alt= Fruit and leaves of Ribes rubrum ]] ==== Grossulariaceae ==== {{Main| Grossulariaceae }} The Grossulariaceae (Gooseberry family) are shrubs that are usually deciduous. The single genus, ''[[Ribes]]'', has about 150 species that are commercially important and widely cultivated for their fruit and also grown as ornamentals, such as ''[[Ribes uva-crispa|R. uva-crispa]]'' (gooseberry) and ''[[Ribes nigrum|R. nigrum]]'' (blackcurrant). They are found in temperate northern hemisphere regions but extending through the Andes into S America. The family name is derived from the Latin word ''grossulus'' (an unripe fig), and ''Ribes'' is Latinised from the [[Semitic languages|semitic]] word ''ribas'' (acid taste).{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Saxifraga granulata 140505.jpg|thumb|130px|left|''[[Saxifraga granulata]]''|alt= Flowers of Saxifraga granulata]] ==== Saxifragaceae ==== {{Main| Saxifragaceae }} The Saxifragaceae (Saxifrage family) are mainly perennial herbs distributed throughout the Northern Hemisphere and Andes, and New Guinea, in damp woodlands and cooler northern regions, rarely aquatic, but are adapted to a wide range of moisture conditions. The family, greatly reduced, includes 35 genera and about 640 species, in two lineages, saxifragoids (e.g. ''[[Saxifraga]]'', rockfoil) and heucheroids (e.g. ''[[Heuchera]]'', coral bells). The largest genus is ''Saxifraga'', the type genus (370 species), though several genera are monotypic. Saxifragaceae are the most horticulturally important of the herbaceous Saxifragales. They provide foodstuffs and medicaments and include many ornamentals, particularly of border, rock and woodland gardens, such as ''[[Astilbe]]'', though the largest number of cultivated species belong to ''Saxifraga''. The family and type genus name are derived from the two Latin words ''saxum'' (rock), and ''frango'' (to break), but the exact origin is unknown, although surmised to be either because of the ability of ''Saxifraga'' to grow in crevices in rocks or medicinal use for [[kidney stones]].{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}}{{sfn|Berry|2017}} |} {{Clear}} {| |[[File:Cynomorium coccineum 2.jpg|thumb|130px|left|''[[Cynomorium coccineum]]''|alt= Inflorescence of Cynomorium coccineum]] ==== Cynomoriaceae ==== {{Main| Cynomoriaceae }} The Cynomoriaceae (Tarthuth or Maltese Mushroom family) consists of a single genus, ''Cynomorium'' with one or two species, ''[[Cynomorium coccineum|C. coccineum]]'' (Mediterranean basin) and ''[[Cynomorium songaricum|C. songaricum]]'' (central Asia and China; sometimes treated as a variety of ''C. coccineum''). They are perennial bisexual herbaceous parasitic plants lacking chlorophyll, from deserts and arid regions. They have been harvested for food, as a dye and in traditional medicine. The name is derived from two Greek words ''kynos'' (dog), and ''morion'' (penis), for its shape.{{sfn|Christenhusz et al|2017}}{{sfn|Byng|2014}} |} {{Clear}}
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