Editing Rosaceae (section)

==Taxonomy==

=== Taxonomic history ===
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.<ref>Caratini, Roger. La Vie de plantes. 1971. Encyclopédie Bordas.</ref><ref>Lawrence, G.H.M. 1960. ''Taxonomy of Vascular Plants''. Macmillan.</ref> 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.<ref>{{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 }}</ref> 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)<ref>{{cite book |
 author = Hutchinson J. | year = 1964 | title = The Genera of Flowering Plants | volume = 1, Dicotyledons | publisher = Clarendon Press | location = Oxford | pages = 1–516}}</ref> and Kalkman (2004) <ref>{{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}}</ref> recognized only tribes (17 and 21, respectively). Takhtajan (1997) delimited 21 tribes in 10 subfamilies:<ref name="Takhtajan" /> 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, ''[[Rose|Rosa]]'' around 100 (including the taxonomically complex [[dog rose]]s), ''[[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.

=== Genera ===
{{main|List of Rosaceae genera}}

Identified clades include:
* Subfamily [[Rosoideae]]: Traditionally composed of those genera bearing [[fruit#Aggregate fruit|aggregate fruits]] that are made up of small achenes or [[drupelet]]s, and often the fleshy part of the fruit (e.g. [[strawberry]]) is the [[Receptacle (botany)|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.<ref name="Potter" /> 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 (plant)|Dryas]]'', ''[[Cercocarpus]]'', ''[[Chamaebatia]]'', ''[[Cowania (plant)|Cowania]]'', and ''[[Purshia]]''), most species of which form [[root nodule]]s which host nitrogen-fixing bacteria from the genus ''[[Frankia]]''.

=== Phylogeny ===
The [[phylogeny|phylogenetic relationships]] between the three subfamilies within Rosaceae are unresolved. There are three competing hypotheses:

{| {{Table}}
! | Amygdaloideae basal
! | Dryadoideae basal
! | Rosoideae basal
|-
| {{clade | 1={{clade | 1=Amygdaloideae | 2={{clade | 1=Rosoideae | 2=Dryadoideae}}}}}}
| {{clade | 1={{clade | 1=Dryadoideae | 2={{clade | 1=Amygdaloideae | 2=Rosoideae}}}}}}
| {{clade | 1={{clade | 1=Rosoideae | 2={{clade | 1=Dryadoideae | 2=Amygdaloideae}}}}}}
|}

==== Amygdaloideae basal ====
Amygdaloideae has been identified as the [[basal (phylogenetics)|earliest branching]] subfamily by Chin et al. (2014),<ref>{{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 = [[Molecular Phylogenetics and Evolution|Mol Phylogenet Evol]] | volume = 76 | pages = 34–48 | doi = 10.1016/j.ympev.2014.02.024 | pmid = 24631854| bibcode = 2014MolPE..76...34C }}</ref> Li et al. (2015),<ref>{{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 = [[Scientific Reports|Sci Rep]] | volume = 5 | page = 14023 | doi = 10.1038/srep14023 | pmid = 26354898 | pmc = 4650596| bibcode = 2015NatSR...514023L }}</ref> Li et al. (2016),<ref>{{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 }}</ref> and Sun et al. (2016).<ref>{{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 }}</ref> Most recently Zhang et al. (2017) recovered these relationships using whole [[plastid]] genomes:<ref name="Zhang">{{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 Phytologist|New Phytol]] | volume = 214 | issue = 3 | pages = 1355–1367 | pmid = 28186635 | doi = 10.1111/nph.14461| doi-access = free | bibcode = 2017NewPh.214.1355Z }}</ref>

{{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]] <!-- =Adenostomeae -->
               }}
            |2=[[Amygdaleae]]
             }}
          |label2=Kerriodae
          |2={{clade
            |1=[[Kerrieae]]
            |2=[[Exochordeae]] <!-- =Osmaronieae -->
             }}
           }}
        |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=[[Sanguisorbeae|Agrimonieae]]
               }}
            |2=[[Rubeae]]
             }}
          |2=[[Colurieae]] <!-- =Geeae -->
           }}
        |2=[[Ulmarieae]]
         }}
      |label2='''[[Dryadoideae]]'''
      |2=[[Dryadeae]]
       }}
     }}
  |2=[[Outgroup (cladistics)|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 (botany)|ovary]], and the fruits are usually achenes.<ref name="Zhang" />

==== Dryadoideae basal ====
Dryadoideae has been identified as the earliest branching subfamily by Evans et al. (2002)<ref>{{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}}</ref> and Potter (2003).<ref>{{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}}</ref> Most recently Xiang et al. (2017) recovered these relationships using [[Cell nucleus|nuclear]] [[transcriptomes]]:<ref>{{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 = [[Molecular Biology and Evolution|Mol Biol Evol]] | volume = 34 | issue = 2 | pages = 262–281 | doi = 10.1093/molbev/msw242 | pmid = 27856652 | pmc = 5400374}}</ref>

{{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]] <!-- =Osmaronieae -->
                   }}
                |2=[[Sorbarieae]] <!-- =Adenostomeae -->
                 }}
               }}
            |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=[[Sanguisorbeae|Agrimonieae]]
                |2=[[Potentilleae]]
                 }}
              |2=[[Roseae]]
               }}
            |2=[[Colurieae]] <!-- =Geeae -->
             }}
          |2=[[Rubeae]]
           }}
        |2=[[Ulmarieae]]
         }}
       }}
    |label2='''[[Dryadoideae]]'''
    |2=[[Dryadeae]]
     }}
  |2=[[Outgroup (cladistics)|outgroup]]
   }}
}}

==== Rosoideae basal ====
Rosoideae has been identified as the earliest branching subfamily by Morgan et al. (1994),<ref>{{cite journal |vauthors=Morgan DR, Soltis DE, Robertson KR | year = 1994 | title = Systematic and evolutionary implications of ''rbcL'' sequence variation in Rosaceae | journal = [[American Journal of Botany|Am J Bot]] | volume = 81 | issue = 7 | pages = 890–903 | jstor = 2445770 | doi = 10.2307/2445770 }}</ref> Evans (1999),<ref>{{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 }}</ref> Potter et al. (2002),<ref>{{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 Systematics and Evolution|Plant Syst Evol]] | volume = 231 | issue = 1–4 | pages = 77–89 | doi = 10.1007/s006060200012| bibcode = 2002PSyEv.231...77P | s2cid = 35829880 }}</ref> Potter et al. (2007),<ref name="Potter">{{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 }}</ref> Töpel et al. (2012),<ref>{{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 }}</ref> and Chen et al. (2016).<ref>{{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 }}</ref> The following is taken from Potter et al. (2007):<ref name="Potter" />

{{Clade
|1={{clade
  |label1=Rosaceae
  |1={{clade
    |label1='''[[Rosoideae]]'''
    |1={{clade
      |label1=Rosodae
      |1={{clade
        |1={{clade
          |1={{clade
            |1=[[Sanguisorbeae|Agrimonieae]]
            |2={{clade
              |1=[[Potentilleae]]
              |2=[[Roseae]]
               }}
             }}
          |2=[[Colurieae]] <!-- =Geeae -->
           }}
        |2=[[Rubeae]]
         }}
      |2=[[Ulmarieae]]
       }}
    |2={{clade
      |label1='''[[Amygdaloideae]]'''
      |1={{clade
        |1={{clade
          |1={{clade
             |1={{clade
               |1=[[Sorbarieae]] <!-- =Adenostomeae -->
               |2={{clade
                 |label1=Malodae
                 |1={{clade
                   |1=[[Maleae]]
                   |2=[[Gillenieae]]
                    }}
                 |2=[[Spiraeeae]]
                  }}
                }}
             |label2=Kerriodae
             |2={{clade
               |1=[[Kerrieae]]
               |2=[[Exochordeae]] <!-- =Osmaronieae -->
                }}
              }}
          |2={{clade
             |1=[[Amygdaleae]]
             |2=[[Neillieae]]
              }}
           }}
        |2=[[Lyonothamneae]]
         }}
      |label2='''[[Dryadoideae]]'''
      |2=[[Dryadeae]]
       }}
     }}
  |2=[[Outgroup (cladistics)|outgroup]]
   }}
}}

The sister relationship between Amygdaloideae and Dryadoideae is supported by the following shared biochemical characters not found in Rosoideae: production of [[cyanogenic glycoside]]s and production of [[sorbitol]].<ref name="Zhang" />