rosaceae

Rosaceae can be woody trees, shrubs, climbers or herbaceous plants.{{Cite book |last1=Heywood |first1=V.H. |title=Flowering Plant Families of the World |last2=Brummitt |first2=R.K. |last3=Culham |first3=A. |last4=Seberg |first4=O. |publisher=Firefly Books |year=2007 |isbn=978-1-55407-206-4 |location=Ontario, Canada |pages=280–282}} The herbs are mostly perennials, but some annuals also exist, such as Aphanes arvensis.{{cite book|last=Stace|first=C. A.|author-link = Stace, C. A.|year=2019|title=New Flora of the British Isles|edition=Fourth|publisher=C & M Floristics|location = Middlewood Green, Suffolk, U.K.| isbn=978-1-5272-2630-2}}{{rp|200}}{{cite web |title=Rosaceae Juss.: FloraBase: Flora of Western Australia |url=http://florabase.calm.wa.gov.au/browse/profile/22834 |work=calm.wa.gov.au |access-date=21 April 2010 |archive-date=15 March 2011 |archive-url=https://web.archive.org/web/20110315075449/http://florabase.calm.wa.gov.au/browse/profile/22834 }}

The leaves are generally arranged spirally, but have an opposite arrangement in some species. They can be simple or pinnately compound (either odd- or even-pinnate). Compound leaves appear in around 30 genera. The leaf margin is most often serrate. Paired stipules are generally present and are considered a primitive feature within the family, though they have been independently lost in many groups of Amygdaloideae (previously called Spiraeoideae). The stipules are sometimes adnate (attached surface to surface){{cite book |last=Beentje |first=H. |title=The Kew Plant Glossary, an Illustrated Dictionary of Plant Terms |publisher=Kew publishing |year=2010 |isbn=978-1-842-46422-9 |location=Kew, London, U.K.}} to the petiole. Glands or extrafloral nectaries may be present on leaf margins or petioles. Spines may be present on the midrib of leaflets and the rachis of compound leaves.

Flowers of plants in the rose family are generally described as "showy".{{cite book |editor-last1=Folta |editor-first1=Kevin M. |editor-last2=Gardiner|editor-first2=Susan E.|title=Genetics and Genomics of Rosaceae |publisher=Springer |year=2008 |isbn=978-0-387-77490-9 |edition=1 |location=New York |page=2 |author-link=Kevin Folta}} They are radially symmetrical, and almost always hermaphroditic. Rosaceae generally have five sepals, five petals, and many spirally arranged stamens. The bases of the sepals, petals, and stamens are fused together to form a characteristic cup-like structure called a hypanthium. They can be arranged in spikes, or heads. Solitary flowers are rare. Rosaceae have a variety of color petals, but blue is almost completely absent.

The fruits occur in many varieties and were once considered the main characters for the definition of subfamilies amongst Rosaceae, giving rise to a fundamentally artificial subdivision. They can be follicles, capsules, nuts, achenes, drupes (Prunus), and accessory fruits, like the pome of an apple, the hip of a rose, or the receptacle-derived aggregate accessory fruit of a strawberry. Many fruits of the family are edible, but their seeds often contain amygdalin, which can release cyanide during digestion if the seed is damaged.TOXNET: [http://toxnet.nlm.nih.gov/cgi-bin/sis/search/a?dbs+hsdb:@term+@DOCNO+3559 CASRN: 29883-15-6]

The family was traditionally divided into six subfamilies: Rosoideae, Spiraeoideae, Maloideae (Pomoideae), Amygdaloideae (Prunoideae), Neuradoideae, and Chrysobalanoideae, and most of these were treated as families by various authors.Caratini, Roger. La Vie de plantes. 1971. Encyclopédie Bordas.Lawrence, G.H.M. 1960. Taxonomy of Vascular Plants. Macmillan. More recently (1971), Chrysobalanoideae was placed in Malpighiales in molecular analyses and Neuradoideae has been assigned to Malvales. Schulze-Menz, in Engler's Syllabus edited by Melchior (1964) recognized Rosoideae, Dryadoideae, Lyonothamnoideae, Spireoideae, Amygdaloideae, and Maloideae.{{cite book | author = Schulze-Menz GK. | year = 1964 | chapter = Rosaceae | editor = Melchior H | title = Engler's Syllabus der Pflanzenfamilien | volume = II | edition = 12 | publisher = Gebrüder Borntraeger | location = Berlin | pages = 209–218 }} They were primarily diagnosed by the structure of the fruits. More recent work has identified that not all of these groups were monophyletic. Hutchinson (1964){{cite book |

author = Hutchinson J. | year = 1964 | title = The Genera of Flowering Plants | volume = 1, Dicotyledons | publisher = Clarendon Press | location = Oxford | pages = 1–516}} and Kalkman (2004) {{cite book | author = Kalkman C. | year = 2004 | chapter = Rosaceae | editor = Kubitzki K | series = The Families and Genera of Vascular Plants | volume = 6 | title = Flowering plants—Dicotyledons: Celastrales, Oxalidales, Rosales, Cornales, Ericales | publisher = Springer-Verlag | location = Berlin Heidelberg | pages = 343–386 | isbn = 978-3-540-06512-8 | doi = 10.1007/978-3-662-07257-8 | s2cid = 12809916 | edition = 1}} recognized only tribes (17 and 21, respectively). Takhtajan (1997) delimited 21 tribes in 10 subfamilies: Filipenduloideae, Rosoideae, Ruboideae, Potentilloideae, Coleogynoideae, Kerroideae, Amygdaloideae (Prunoideae), Spireoideae, Maloideae (Pyroideae), Dichotomanthoideae. A more modern model comprises three subfamilies, one of which (Rosoideae) has largely remained the same.

While the boundaries of the Rosaceae are not disputed, there is no general agreement as to how many genera it contains. Areas of divergent opinion include the treatment of Potentilla s.l. and Sorbus s.l.. Compounding the problem is that apomixis is common in several genera. This results in an uncertainty in the number of species contained in each of these genera, due to the difficulty of dividing apomictic complexes into species. For example, Cotoneaster contains between 70 and 300 species, Rosa around 100 (including the taxonomically complex dog roses), Sorbus 100 to 200 species, Crataegus between 200 and 1,000, Alchemilla around 300 species, Potentilla roughly 500, and Rubus hundreds, or possibly even thousands of species.

{{main|List of Rosaceae genera}}

Identified clades include:

• Subfamily Rosoideae: Traditionally composed of those genera bearing aggregate fruits that are made up of small achenes or drupelets, and often the fleshy part of the fruit (e.g. strawberry) is the receptacle or the stalk bearing the carpels. The circumscription is now narrowed (excluding, for example, the Dryadoideae), but it still remains a diverse group containing five or six tribes and 20 or more genera, including rose, Rubus (blackberry, raspberry), Fragaria (strawberry), Potentilla, and Geum.
• Subfamily Amygdaloideae: Within this group remains an identified clade with a pome fruit, traditionally known as subfamily Maloideae (or Pyroideae) which included genera such as apple, Cotoneaster, and Crataegus (hawthorn). To separate it at the subfamily level would leave the remaining genera as a paraphyletic group, so it has been expanded to include the former Spiraeoideae and Amygdaloideae. The subfamily has sometimes been referred to by the name "Spiraeoideae", but this is not permitted by the International Code of Nomenclature for algae, fungi, and plants.
• Subfamily Dryadoideae: Fruits are achenes with hairy styles, and includes five genera (Dryas, Cercocarpus, Chamaebatia, Cowania, and Purshia), most species of which form root nodules which host nitrogen-fixing bacteria from the genus Frankia.

The phylogenetic relationships between the three subfamilies within Rosaceae are unresolved. There are three competing hypotheses:

Amygdaloideae has been identified as the earliest branching subfamily by Chin et al. (2014),{{cite journal |vauthors=Chin SW, Shaw J, Haberle R, Wen J, Potter D | year = 2014 | title = Diversification of almonds, peaches, plums and cherries—Molecular systematics and biogeographic history of Prunus (Rosaceae) | journal = Mol Phylogenet Evol | volume = 76 | pages = 34–48 | doi = 10.1016/j.ympev.2014.02.024 | pmid = 24631854| bibcode = 2014MolPE..76...34C }} Li et al. (2015),{{cite journal |vauthors=Li HL, Wang W, Mortimer PE, Li RQ, Li DZ, Hyde KD, Xu JC, Soltis DE, Chen ZD | year = 2015 | title = Large-scale phylogenetic analyses reveal multiple gains of actinorhizal nitrogen-fixing symbioses in angiosperms associated with climate change | journal = Sci Rep | volume = 5 | page = 14023 | doi = 10.1038/srep14023 | pmid = 26354898 | pmc = 4650596| bibcode = 2015NatSR...514023L }} Li et al. (2016),{{cite journal |vauthors=Li HL, Wang W, Li RQ, Zhang JB, Sun M, Naeem R, Su JX, Xiang XG, Mortimer PE, Li DZ, Hyde KD, Xu JC, Soltis DE, Soltis PS, Li J, Zhang SZ, Wu H, Chen ZD, Lu AM | year = 2016 | title = Global versus Chinese perspectives on the phylogeny of the N-fixing clade | journal = Journal of Systematics and Evolution | volume = 54 | issue = 4 | pages = 392–399 | doi = 10.1111/jse.12201| s2cid = 88546939 | doi-access = free | bibcode = 2016JSyEv..54..392L }} and Sun et al. (2016).{{cite journal | year = 2016 | title = Phylogeny of the Rosidae: A dense taxon sampling analysis | journal = Journal of Systematics and Evolution | volume = 54 | issue = 4 | pages = 363–391 | doi = 10.1111/jse.12211 |author=Sun Miao |author2=Naeem Rehan |author3=Su Jun-Xia |author4=Cao Zhi-Yong |author5=Burleigh J. Gordon |author6=Soltis Pamela S. |author7=Soltis Douglas E. |author8=Chen Zhi-Duan | doi-access = free | bibcode = 2016JSyEv..54..363S }} Most recently Zhang et al. (2017) recovered these relationships using whole plastid genomes:{{cite journal |vauthors=Zhang SD, Jin JJ, Chen SY, Chase MW, Soltis DE, Li HT, Yang JB, Li DZ, Yi TS | year = 2017 | title = Diversification of Rosaceae since the Late Cretaceous based on plastid phylogenomics | journal = New Phytol | volume = 214 | issue = 3 | pages = 1355–1367 | pmid = 28186635 | doi = 10.1111/nph.14461| doi-access = free | bibcode = 2017NewPh.214.1355Z }}

{{Clade

|1={{clade

|label1=Rosaceae

|1={{clade

|label1=Amygdaloideae

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|label1=Malodae

|1={{clade

|1=Maleae

|2=Gillenieae

}}

|2=Spiraeeae

}}

|2=Sorbarieae

}}

|2=Amygdaleae

}}

|label2=Kerriodae

|2={{clade

|1=Kerrieae

|2=Exochordeae

}}

}}

|2=Neillieae

}}

|2=Lyonothamneae

}}

|2={{clade

|label1=Rosoideae

|1={{clade

|label1=Rosodae

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|1=Potentilleae

|2=Roseae

}}

|2=Agrimonieae

}}

|2=Rubeae

}}

|2=Colurieae

}}

|2=Ulmarieae

}}

|label2=Dryadoideae

|2=Dryadeae

}}

}}

|2=outgroup

}}

}}

The sister relationship between Dryadoideae and Rosoideae is supported by the following shared morphological characters not found in Amygdaloideae: presence of stipules, separation of the hypanthium from the ovary, and the fruits are usually achenes.

Dryadoideae has been identified as the earliest branching subfamily by Evans et al. (2002){{cite conference | title = A Rosaceae phylogeny |vauthors=Evans RC, Campbell C, Potter D, Morgan D, Eriksson T, Alice L, Oh SH, Bortiri E, Gao F, Smedmark J, Arsenault M | date = 2–7 August 2002 | publisher = Botanical Society of America, St. Louis | book-title = Abstracts | page = 108 | location = Madison, Wisconsin | conference = Botany 2002—Botany in the Curriculum: Integrating Research and Teaching}} and Potter (2003).{{cite book | author = Potter D. | year = 2003 | chapter = Molecular phylogenetic studies in Rosaceae |veditors= Sharma AK, Sharma A | title = Plant Genome: Biodiversity and Evolution | location = Enfield, NH | publisher = Scientific Publications | volume = 1, Part A: Phanerogams | pages = 319–351 | isbn = 978-1-578-08238-4}} Most recently Xiang et al. (2017) recovered these relationships using nuclear transcriptomes:{{cite journal |vauthors=Xiang Y, Huang CH, Hu Y, Wen J, Li S, Yi T, Chen H, Xiang J, Ma H | year = 2017 | title = Evolution of Rosaceae fruit types based on nuclear phylogeny in the context of geological times and genome duplication | journal = Mol Biol Evol | volume = 34 | issue = 2 | pages = 262–281 | doi = 10.1093/molbev/msw242 | pmid = 27856652 | pmc = 5400374}}

{{Clade

|1={{clade

|label1=Rosaceae

|1={{clade

|1={{clade

|label1=Amygdaloideae

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|label1=Malodae

|1={{clade

|1=Maleae

|2=Gillenieae

}}

|2={{clade

|label1=Kerriodae

|1={{clade

|1=Kerrieae

|2=Exochordeae

}}

|2=Sorbarieae

}}

}}

|2={{clade

|1=Amygdaleae

|2=Lyonothamneae

}}

}}

|2=Spiraeeae

}}

|2=Neillieae

}}

|label2=Rosoideae

|2={{clade

|label1=Rosodae

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|1=Agrimonieae

|2=Potentilleae

}}

|2=Roseae

}}

|2=Colurieae

}}

|2=Rubeae

}}

|2=Ulmarieae

}}

}}

|label2=Dryadoideae

|2=Dryadeae

}}

|2=outgroup

}}

}}

Rosoideae has been identified as the earliest branching subfamily by Morgan et al. (1994),{{cite journal |vauthors=Morgan DR, Soltis DE, Robertson KR | year = 1994 | title = Systematic and evolutionary implications of rbcL sequence variation in Rosaceae | journal = Am J Bot | volume = 81 | issue = 7 | pages = 890–903 | jstor = 2445770 | doi = 10.2307/2445770 }} Evans (1999),{{cite web | url = http://labs.eeb.utoronto.ca/dickinson/rosaceaeevolution/phylogeny.html | title = Rosaceae Phylogeny: Origin of Subfamily Maloideae | author = Evans R. | date = 1999 | website = Rosaceae Phylogeny and Evolution | publisher = Botany Department, University of Toronto | access-date = 7 July 2017 | archive-date = 8 March 2016 | archive-url = https://web.archive.org/web/20160308213223/http://labs.eeb.utoronto.ca/dickinson/rosaceaeevolution/phylogeny.html | url-status = dead }} Potter et al. (2002),{{cite journal |vauthors=Potter D, Gao F, Esteban Bortiri P, Oh SH, Baggett S | year = 2002 | title = Phylogenetic relationships in Rosaceae inferred from chloroplast matK and trnL–trnF nucleotide sequence data | journal = Plant Syst Evol | volume = 231 | issue = 1–4 | pages = 77–89 | doi = 10.1007/s006060200012| bibcode = 2002PSyEv.231...77P | s2cid = 35829880 }} Potter et al. (2007),{{cite journal |vauthors=Potter D, Eriksson T, Evans RC, Oh S, Smedmark JE, Morgan DR, Kerr M, Robertson KR, Arsenault M, Dickinson TA, Campbell CS | year = 2007 | title = Phylogeny and classification of Rosaceae | journal = Plant Systematics and Evolution | volume = 266 | issue = 1–2 | pages = 5–43 | url = http://biology.umaine.edu/Amelanchier/Rosaceae_2007.pdf | doi = 10.1007/s00606-007-0539-9 | jstor = 23655774| bibcode = 2007PSyEv.266....5P | s2cid = 16578516 }} Töpel et al. (2012),{{cite journal |vauthors=Töpel M, Antonelli A, Yesson C, Eriksen B | year = 2012 | title = Past climate change and plant evolution in Western North America: A case study in Rosaceae | journal = PLOS One| volume = 7 | issue = 12 | pages = e50358 | pmid = 23236369 | doi = 10.1371/journal.pone.0050358 | pmc=3517582| bibcode = 2012PLoSO...750358T | doi-access = free }} and Chen et al. (2016).{{cite journal |vauthors=Chen ZD, Yan T, Lin L, Lu LM, Li HL, Sun M, Liu B, Chen M, Niu YT, Ye JF, Cao ZY, Liu HM, Wang XM, Wang W, Zhang JB, Meng Z, Cao W, Li JH, Wu SD, Zhao HL, Liu ZJ, Du ZY, Wan QF, Guo J, Tan XX, Su JX, Zhang LJ, Yang LL, Liao YY, Li MH, Zhang GQ, Chung SW, Zhang J, Xiang KL, Li RQ, Soltis DE, Soltis PS, Zhou SL, Ran JH, Wang XQ, Jin XH, Chen YS, Gao TG, Li JH, Zhang SZ, Lu AM |collaboration=China Phylogeny Consortium | year = 2016 | title = Tree of life for the genera of Chinese vascular plants | journal = Journal of Systematics and Evolution | volume = 54 | issue = 4 | pages = 277–306 | doi = 10.1111/jse.12219| doi-access = free |bibcode=2016JSyEv..54..277C }} The following is taken from Potter et al. (2007):

{{Clade

|1={{clade

|label1=Rosaceae

|1={{clade

|label1=Rosoideae

|1={{clade

|label1=Rosodae

|1={{clade

|1={{clade

|1={{clade

|1=Agrimonieae

|2={{clade

|1=Potentilleae

|2=Roseae

}}

}}

|2=Colurieae

}}

|2=Rubeae

}}

|2=Ulmarieae

}}

|2={{clade

|label1=Amygdaloideae

|1={{clade

|1={{clade

|1={{clade

|1={{clade

|1=Sorbarieae

|2={{clade

|label1=Malodae

|1={{clade

|1=Maleae

|2=Gillenieae

}}

|2=Spiraeeae

}}

}}

|label2=Kerriodae

|2={{clade

|1=Kerrieae

|2=Exochordeae

}}

}}

|2={{clade

|1=Amygdaleae

|2=Neillieae

}}

}}

|2=Lyonothamneae

}}

|label2=Dryadoideae

|2=Dryadeae

}}

}}

|2=outgroup

}}

}}

The sister relationship between Amygdaloideae and Dryadoideae is supported by the following shared biochemical characters not found in Rosoideae: production of cyanogenic glycosides and production of sorbitol.

The Rosaceae have a cosmopolitan distribution, being found nearly everywhere except for Antarctica. They are primarily concentrated in the Northern Hemisphere in regions that are not desert or tropical rainforest.

The rose family is considered one of the six most economically important crop plant families,B.C. Bennett (undated). Economic Botany: Twenty-Five Economically Important Plant Families. [http://www.eolss.net/Sample-Chapters/C09/E6-118-03.pdf Encyclopedia of Life Support Systems (EOLSS) e-book] and includes apples, pears, quinces, medlars, loquats, almonds, peaches, apricots, plums, cherries, strawberries, blackberries, raspberries, sloes, and roses.

Many genera are also highly valued ornamental plants. These include trees and shrubs (Cotoneaster, Chaenomeles, Crataegus, Dasiphora, Exochorda, Kerria, Photinia, Physocarpus, Prunus, Pyracantha, Rhodotypos, Rosa, Sorbus, Spiraea), herbaceous perennials (Alchemilla, Aruncus, Filipendula, Geum, Potentilla, Sanguisorba), alpine plants (Dryas, Geum, Potentilla) and climbers (Rosa).

However, several genera are also introduced noxious weeds in some parts of the world, costing money to be controlled. These invasive plants can have negative impacts on the diversity of local ecosystems once established. Such naturalised pests include Acaena, Cotoneaster, Crataegus, and Pyracantha.

In Bulgaria and parts of western Asia, the production of rose oil from fresh flowers such as Rosa damascena, Rosa gallica, and other species is an important economic industry.

The family Rosaceae covers a wide range of trees, bushes and plants.

File:Acaena magellanica magellanica 1.jpg | Buzzy burr (Acaena magellanica)

File:Alchemilla vulgaris.jpg | Common lady's mantle (Alchemilla vulgaris)

File:Aruncus dioicus 15105.JPG | Goat's beard (Aruncus dioicus)

File:Chaenomeles japonica a1.jpg | Maule's quince (Chaenomeles japonica)

File:Cercocarpus betuloides blancheae.JPG | Mountain mahogany (Cercocarpus betuloides)

File:Crataegus Submollis Flowers and Thorns.jpg| Northern downy hawthorn (Crataegus submollis)

File:Cotoneaster adpressus GotBot 2015 001.jpg | Creeping cotoneaster (Cotoneaster adpressus)

File:Dasiphora fruticosa 5698.jpg | Shrubby cinquefoil (Dasiphora fruticosa)

File:Mountainavens2.jpg | Mountain avens (Dryas octopetala)

File:Eriobotrya japonica B.jpg | Loquat (Eriobotrya japonica), a fruit tree typical by flowering in autumn

File:Exochorda racemosa1.jpg | Pearlbush (Exochorda racemosa)

(MHNT) Filipendula vulgaris - Inflorescence.jpg| Dropwort (Filipendula vulgaris)

File:FragariaMoschata.JPG | Musk strawberry (Fragaria moschata) valued for its intense aroma

File:Geum triflorum 4881.JPG | Old man's whiskers (Geum triflorum)

File:Kerria japonica 2.JPG | Kerria japonica

File:Apple blossoms.jpg | Apple tree blossoms (Malus pumila)

File:Mispel-Crataegus-germanica-001.jpg | Common medlar (Mespilus germanica)

File:Photinia fraseri B.JPG | Red Tip Photinia (Photinia x fraseri) popular for red color of its new growths

File:Physocarpus opulifolius USFWS.jpg | Common ninebark (Physocarpus opulifolius)

File:Potentilla reptans sl8.jpg | Creeping cinquefoil (Potentilla reptans)

File:PikiWiki Israel 7025 Amond blossom.jpg | Mature fruit of an almond tree (Prunus dulcis)

File:PRUNUS SPINOSA - SANT JUST - IB-481 (Aranyoner).JPG | Blackthorn (Prunus spinosa)

File:Purshia stansburiana 2.jpg | Stansbury's cliffrose (Purshia stansburyana)

File:Brosen pyracantha coccinea1.jpg | Scarlet firethorn (Pyracantha coccinea)

File:Pear-tree,katori-city,japan.JPG | Nashi pear (Pyrus pyrifolia) typical for Asian countries

File:Rhodotypos-scandens-fruit.JPG | Rhodotypos scandens, a Japanese shrub with fruits high in toxic amygdalin

File:Rosa sericea jd plt 1.jpg | The silky rose (Rosa sericea) known for its ornamental prickles

File:Rubus spectabilis 39139.JPG | Salmonberry (Rubus spectabilis)

File:Toten 2.jpg | Great burnet (Sanguisorba officinalis)

File:Sorbus alnifolia 'Submollis' JPG1La.jpg | Autumn foliage of the Korean mountain ash (Sorbus alnifolia)

File:Spiraea splendens 21648.JPG | Rose meadowsweet (Spiraea splendens)

File:Flowers of Rosa chinensis.jpg| Rosa chinensis (Rosa chinensis)

{{Reflist}}

{{Commons category|Rosaceae}}

• [http://delta-intkey.com/angio/www/rosaceae.htm Rosaceae at the DELTA Online Families of Flowering Plants] {{Webarchive|url=https://web.archive.org/web/20110514061816/http://delta-intkey.com/angio/www/rosaceae.htm |date=14 May 2011 }}

{{Angiosperm families}}

{{Taxonbar|from=Q46299}}

{{Authority control}}

Category:Rosales

Category:Rosid families

Category:Extant Cretaceous first appearances