Saxifragaceae


Saxifragaceae is a family of herbaceous perennial flowering plants, within the core eudicot order Saxifragales. The taxonomy has been greatly revised and the scope of the family much reduced in the era of molecular phylogenetic analysis. The family is divided into ten clades, with about 640 known species in about 35 accepted genera. About half of these consist of a single species, but about 400 of the species are in the type genus Saxifraga. The family is predominantly distributed in the northern hemisphere, but also in the Andes in South America.

Description

, sometimes succulent or xerophytic, often with perennating rhizomes. Leaves usually basally aggregated in alternate rosettes, sometimes on inflorescence stems. They are usually simple, rarely pinnately or palmately compound. Their margins may be entire, deeply lobed, cleft, crenate or dentate and petiolate with stipules.
Inflorescences are bracteate racemes or cymes. Flowers are hermaphroditic, rarely unisexual, actinomorphic. The perianth is placed on a hypanthium that may be free or may be partly fused with the ovary. Sepals usually 5 but may be 3–10 and fused with hypanthium, occasionally petaloid. Petals are clawed, sometimes cleft at the tip or finely dissected. Stamens 5–10, free and opposite petals, anthers usually basifixed and opening by lengthwise slits. Ovary inferior to semi-inferior with two carpels usually fused at the base, sometimes free, each topped with stylodium and capitate stigma. Ovules few to many, placentation axile or parietal, styles 2–3. Fruit usually septicidal capsule or follicle, with numerous small seeds.

Taxonomy

Saxifragaceae has long been considered in a pivotal position in the evolution of angiosperm lineage, located in an ancestral "Saxifragalean stock". Historically the Saxifragaceae have included many very morphologically disparate taxa in systems based on morphology alone, and has been very difficult to classify and characterize phenotypically. Consequently many different classifications have been published, differing considerably in both the relationship of the family to other angiosperms, and its internal structure, for instance the systems of Cronquist, Dahlgren, Engler, Hutchinson, Schulze-Menz, Takhtajan and Thorne . Within those subfamilies, the large majority of genera were located within subfamily Saxifragoideae, the core group, with all but two of the remainder only having 1-3 genera.
The circumscription of the family has changed considerably in recent years, in large part due to molecular phylogenetic work showing the family s.l. to be polyphyletic, and probably represents the most extreme example, with at least 10 evolutionary lines. Consequently the circumscription has been considerably reduced, with many of the subfamilies being either elevated to separate families, or placed as components of other families, often quite distant. For instance subfamily Parnassioideae was raised to the level of family Parnassiaceae, and eventually a subfamily of Celastraceae. Similarly the Hydrangoideae is now the family Hydrangeaceae.
The reduced Saxifragaceae is distinguished by being referred to as Saxifragaceae sensu stricto, corresponding to the Saxifragoideae a natural monophyletic group. Historically, the Saxifragaceae has been placed in either the orders Rosales or Saxifragales. This reduced Saxifragaceae, corresponding to subfamily Saxifragoideae and its 30 genera, is now placed within the Saxifragales. There, with three other subfamilies it forms the Saxifragaceae alliance, while Penthoroideae and Tetracarpaeoideae are closely related within the core Saxifragales as shown in the cladogram. The remaining subfamilies are all transferred to more distant orders within the rosid and asterid clades.

Subdivision

Numerous attempts have been made to subdivide Saxifragoideae. These have included dividng the family by the placentation of the ovules, as either parietal or axile. None of these has been supported by molecular data.
Molecular data indicate that the family can be considered as a number of informal clades, with two main lineages, saxifragoids and heucheroids and further subdivision of heucheroids into nine subclades or groups:
;Clades
The clades and subclades are related as shown in the cladogram:

Genera

Saxifragaceae s.s. has about 33–35 genera and about 640 species. About half of the genera are monotypic, but Saxifraga has about 400 species, and has generally been divided into sections.

Evolution and biogeography

The crown group of Saxifragaceae diversified at about 38 Mya, with the two main lineages diversifying arising at about 30 Mya. The present day heucheroid diversity dates later than the Miocene. Ancestral Saxafragaceae emerged in either East Asia or Western North America, with subsequent dispersal West to Europe and south to South America. From the Eocene to the late Miocene these ancestral land masses were joined by the Bering Land Bridge facilitating plant migration.

Etymology

The family and type genus name are derived from the two Latin words saxum, and frango, but the exact origin is unknown, although surmised to refer to either growing in crevices in rocks or medicinal use for kidney stones.

Distribution and habitat

Primarily Northern hemisphere temperate and arctic regions, and also tropical montane, including the Americas, Europe, North Africa and temperate and subtropical Asia to Luzon and New Guinea. In the Americas it extends south to central Mexico and the Andes to Tierra del Fuego. Centres of diversity are western N America, East Asia and the Himalayas. The largest concentration of genera are in the Pacific North West. Some Saxifraga and Chrysosplenium are circumboreal. Others have disjunct distributions between E Asia and N America, while other taxa have separate narrow distributions in southern S America. Some species are found in wet woodlands, swamplike conditions and wet cliff edges.

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