Geraniales

Classification System: APG IV

Superregnum: Eukaryota
Regnum: Plantae
Cladus: Angiosperms
Cladus: Eudicots
Cladus: Core eudicots
Cladus: Rosids
Cladus: Eurosids II
Ordo: Geraniales
Familiae: Francoaceae – Geraniaceae – Greyiaceae – Melianthaceae – Vivianiaceae
Name

Geraniales Juss. ex Bercht. & J. Presl Prir. Rostlin, 221. (1820)

Typus: Geranium L. Sp. Pl. 2: 676. (1753)

Synonyms

Francoales
Greyiales
Ledocarpales
Melianthales

Note: See discussion page, but the 5 family circumscription is not supported by APGIV (2016).
References

Jussieu, A.L. de 1820. O Prirozenosti Rostlin 221.
Angiosperm Phylogeny Group. 2016. An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV. Botanical Journal of the Linnean Society 181(1): 1–20. DOI: 10.1111/boj.12385 Reference page. Favours two families.
Applequist, W.L. 2012. Report of the nomenclature committee for vascular plants: 64. Taxon 61(5): 1108–1117. There was no consensus for or against a proposal to superconserve Melianthaceae against Francoaceae.
Jeiter, J., Weigend, M. & Hilger, H.H. 2017. Geraniales flowers revisited: evolutionary trends in floral nectaries. Annals of Botany 119(3): 395-408. DOI: 10.1093/aob/mcw230 Open access Reference page. Favours five families.
Palazzesi, L., Gottschling, M., Barreda, V., & Weigend, M. 2012. First Miocene fossils of Vivianiaceae shed new light on phylogeny, divergence times, and historical biogeography of Geraniales. Biological Journal of the Linnean Society 107: 67–85. DOI: 10.1111/j.1095-8312.2012.01910.x ResearchGate Reference page. Favours five families.
Reveal, J.L. 2009. (1915) Proposal to conserve Melianthaceae, nom. cons., against Francoaceae (Magnoliophyta), a superconservation proposal. Taxon 58(4): 1373–1374.
Stevens, P.F. 2001 onwards. Angiosperm Phylogeny Website. Version 14, July 2017 [and more or less continuously updated since]. Online. Reference page. Favours two families.
Sytsma, K.J., Spalink, D. & Berger, B. 2014. Calibrated chronograms, fossils, outgroup relationships, and root priors: re‐examining the historical biogeography of Geraniales. Biological Journal of the Linnean Society 113(1): 29-49. DOI: 10.1111/bij.12297 PDF Reference page. Ambiguously favours two families.
Tropicos.org 2014. Geraniales. Missouri Botanical Garden. Published on the internet. Accessed: 2014 Jan. 25.

Vernacular names
العربية: غرنوقيات
azərbaycanca: Ətirşahçiçəklilər
беларуская: Геранекветныя
català: Geranial
čeština: kakostotvaré
dansk: Storkenæb-ordenen
Deutsch: Storchschnabelartige
فارسی: شمعدانی‌سانان
עברית: גרנאים
hrvatski: Iglicolike
magyar: Gólyaorrvirágúak
日本語: フウロソウ目
한국어: 쥐손이풀목
lietuvių: Snaputiečiai
македонски: Здравцовидни
norsk: Storkenebbordenen
polski: Bodziszkowce
русский: Гераниецветные
slovenčina: pakostotvaré
svenska: Näveordningen
తెలుగు: జిరానియేలిస్
українська: Геранієцвіті
Tiếng Việt: Bộ Mỏ hạc
中文: 牻牛儿苗目

Geraniales are a small order of flowering plants, included within the rosid subclade of eudicots. The largest family in the order is Geraniaceae with over 800 species. In addition, the order includes the smaller Francoaceae with about 40 species. Most Geraniales are herbaceous, but there are also shrubs and small trees.

Flower morphology of the Geraniales is rather conserved. They are usually perfectly pentamerous and pentacyclic without fused organs besides the carpels of the superior gynoecium. The androecium is obdiplostemonous. Only a few genera are tetramerous (Francoa, Tetilla, Melianthus). In some genera some stamens (Pelargonium) or a complete whorl of stamens are reduced (Erodium, Melianthus). In the genera Hypseocharis and Monsonia there are 15 instead of the usual ten stamens. Most genera bear nectariferous flowers.[2] The nectary glands are formed by the receptacle and are localised at the bases of the antesepalous stamens.[2][3]

The economic importance of Geraniales is low. Some species of the genus Pelargonium (Geraniaceae) are cultivated for their aromatic oil used in the perfume industry. Some other species, also mostly within Geraniaceae, have horticultural or medicinal uses. A Paleobotanic record is missing.

Taxonomy
Origins

The botanical authority for Geraniales is given to Jussieu,[4] but since the original description did not fulfill all the rules for a valid publication and was subsequently validly published, attribution is given to both Jussieu and the subsequent publication, hence the designation Geraniales Juss. ex Bercht. & J.Presl Jussieu, who developed the concept of botanical families, described the Gerania, as a grouping of five genera, including Geranium. Although Jussieu used the term Ordo this did not correspond to current understandings of the term Order.[5][6] The subsequent attribution occurred in 1820, in the Czech text O Prirozenosti Rostlin, by Friedrich von Berchtold and Jan Svatopluk Presl, hence ex Bercht. & J.Presl.[7] However, Berchtold and Presl also only described a rad (ordo) of five genera, which they called Geraniae.[8] Other authorities have given the authority to Dumortier who described the family Geraniaceae, consisting of two tribes, Pelargonieae and Geranieae, each with three genera.[9]
Circumscription

Geraniales contain two families, 11 genera and about 830 species.[10] For a historical account of the circumscription of the order, see Price and Palmer (1993) Table 1.[11]

Under the Cronquist system (1988),[12] the Geraniales comprised the following five families:

family Geraniaceae
family Oxalidaceae
family Limnanthaceae
family Tropaeolaceae
family Balsaminaceae

While the Dahlgren system (1980) was much larger in circumscription with 16 families, only two of which were in Cronquist's construction, and placed the order in the superorder Rutiflorae:[13]

family Zygophyllaceae
family Nitrariaceae
family Peganaceae
family Balanitaceae
family Erythroxylaceae
family Humiriaceae
family Linaceae
family Ctenolophonaceae
family Ixonanthaceae
family Lepidobotryaceae
family Oxalidaceae (including Averrhoaceae)
family Geraniaceae
family Dirachmaceae
family Ledocarpaceae
family Vivianiaceae
family Biebersteiniaceae

Other modern systems include those of Takhtajan (1987) with nine families, and Thorne (1992). Thorne's system was the same as Cronquist's except that Biebersteiniaceae, Dirachmaceae, Ledocarpaceae, and Vivianiaceae were reduced to subfamilies of Geraniaceae.
Molecular phylogenetics: Angiosperm Phylogeny Group

The elucidation of the relationships within the order by morphological or cytological methods alone had proven difficult as demonstrated by the widely different treatment by various authorities. For instance Cronquist and Thorne immersed the families Biebersteiniaceae, Dirachmaceae, Ledocarpaceae, Rhynchothecaceae and Vivianiaceae within Geraniaceae (Geraniaceae sensu lato), whereas Dahlgren and others maintained them as separate taxa, maintaining a "core" Geraniaceae (Geraniaceae sensu stricto).[11] Price and Palmer (1993) were among the first investigators to apply molecular phylogenetics to this order, using the chloroplast gene rbcL.[11][14][a] This disassembled the traditional morphologically defined grouping of dicotyledons, replacing it with a series of nested clades. The Geraniales segregated in the eudicot clade, specifically in the rosid subclade.

The family circumscription of the Angiosperm Phylogeny Group (APG) of 1998 placed Geraniales Dumort. amongst the rosids with the following six families:[15]

Geraniales Dumort. 1829[16]

Francoaceae A.Juss., 1832
Geraniaceae Juss., 1789 [ + Hypseocharitaceae Wedd., 1861]
Greyiaceae Hutch., 1926
Ledocarpaceae Meyen, 1834
Melianthaceae Bercht. & J.Presl, 1820
Vivianiaceae Klotzsch, 1836

Hypseocharitaceae were a small family of eight species of the genus Hypseocharis found in the tropical mountainous regions of the Andes.[17] The APG provided the option of considering them as a separate family or subsumed into Geraniaceae. By 2003, when the APG was published, it was apparent that the small families Francoaceae, Greyiaceae and Melianthaceae were closely related and were collapsed into one family as Melanthiaceae with Francoaceae as an optional synonym. Thus the number of families was reduced to four.[18]

The APG III classification (2009) was typical of newer arrangements. In this definition, Hypseocharitaceae was included within Geraniaceae, Francoaceae and Greyiaceae were included within Melianthaceae, and Ledocarpaceae was included within the Vivianiaceae.[1]

However, Considerable rearrangements took place in the 2016 APG IV system. Francoaceae was substituted for Melianthaceae, due to nomenclatural priority. The latter subsumed Vivianiaceae based on the work of Sytsma, Spalink & Berger (2014). However, there remains substantial uncertainty regarding the relationships within Francoaceae sensu stricto (s.s.), Melianthaceae (Bersama Fresen. and Melianthus L.) and Ledocarpaceae. Here, Vivianiaceae is used as a later synonym for Ledocarpaceae. This due to conflicting evidence (see Palazzesi et al., 2012). The APG chose to follow the broader circumscription for the time being till these differences are resolved.

This leaves the order Geraniales with only two families: Geraniaceae and Francoaceae (including Bersamaceae, Greyiaceae, Ledocarpaceae, Melianthaceae, Rhynchothecaceae and Vivianiaceae).[19]

The Vivianiaceae and Ledocarpaceae were included within the Geraniaceae, and the Hypseocharitaceae within the Oxalidaceae, which are now treated in the order Oxalidales. The Melianthaceae were placed within the Sapindales, the Greyiaceae and Francoaceae within the Rosales, the latter subsumed within the Saxifragaceae.

Recent comparison of DNA-fragments from species within the order resulted in the following phylogenetic tree.[20]

Geraniales	Hypseocharitaceae     Geraniaceae         Francoaceae in sensu lato:   Francoaceae (core)     Viviania and Balbisia formerly Vivianiaceae       Melianthus, Greyia and Bersama formerly Melianthaceae

 
 
 
Notes

Part of a much larger multi-institutional study of the phylogeny of seed plants (Ann. Missouri Bot. Gard. 1993)

References

APG III 2009.
Jeiter, Julius; Weigend, Maximilian; Hilger, Hartmut H. (2017-02-01). "Geraniales flowers revisited: evolutionary trends in floral nectaries". Annals of Botany. 119 (3): 395–408. doi:10.1093/aob/mcw230. ISSN 0305-7364. PMC 5314648. PMID 28025289.
Jeiter, Julius; Hilger, Hartmut H; Smets, Erik F; Weigend, Maximilian (2017-11-10). "The relationship between nectaries and floral architecture: a case study in Geraniaceae and Hypseocharitaceae". Annals of Botany. 120 (5): 791–803. doi:10.1093/aob/mcx101. ISSN 0305-7364. PMC 5691401. PMID 28961907.
Jussieu 1789, Ordo XIII Gerainia, Les Geraines p. 268
ICN 2012, 18.2 Names of families and subfamilies, tribes and subtribes.
Candolle 1813, Des familles et des tribus pp. 192–195
Tropicos 2015, Geraniales Juss. ex Bercht. & J. Presl
Berchtold & Presl 1820, Geraniae p. 221
Dumortier 1829, Geraniaceae Juss. p. 46
Christenhusz & Byng 2016.
Price & Palmer 1993.
Cronquist 1988.
Dahlgren 1980.
Chase et al 1993.
APG I 1998.
Dumortier 1829.
Watson & Dallwitz 2016, Hypseocharitaceae Weddell
APG II 2003.
APG IV 2016.

Jeiter, Julius; Cole, Theodor C.H.; Hilger, Hartmut H. "Geraniales Phylogeny Poster (GPP) - 2017". ResearchGate. doi:10.7287/peerj.preprints.3127v1. Retrieved 2017-09-27.

Bibliography

"rbcL Sequence Data and Phylogenetic Reconstruction in Seed Plants". Annals of the Missouri Botanical Garden. 80 (3): 523–786. Summer 1993. JSTOR i317486. (also at Biodiversity Heritage Library: here)
Chase, Mark W.; Soltis, Douglas E.; Olmstead, Richard G.; Morgan, David; Les, Donald H.; Mishler, Brent D.; Duvall, Melvin R.; Price, Robert A.; Hills, Harold G.; Qiu, Yin-Long; Kron, Kathleen A.; Rettig, Jeffrey H.; Conti, Elena; Palmer, Jeffrey D.; Manhart, James R.; Sytsma, Kenneth J.; Michaels, Helen J.; Kress, W. John; Karol, Kenneth G.; Clark, W. Dennis; Hedren, Mikael; Gaut, Brandon S.; Jansen, Robert K.; Kim, Ki-Joong; Wimpee, Charles F.; Smith, James F.; Furnier, Glenn R.; Strauss, Steven H.; Xiang, Qui-Yun; Plunkett, Gregory M.; Soltis, Pamela S.; Swensen, Susan M.; Williams, Stephen E.; Gadek, Paul A.; Quinn, Christopher J.; Eguiarte, Luis E.; Golenberg, Edward; Learn, Gerald H.; Graham, Sean W.; Barrett, Spencer C. H.; Dayanandan, Selvadurai; Albert, Victor A. (1993). "Phylogenetics of Seed Plants: An Analysis of Nucleotide Sequences from the Plastid Gene rbcL" (PDF). Annals of the Missouri Botanical Garden. 80 (3): 528. doi:10.2307/2399846. hdl:1969.1/179875. JSTOR 2399846.
Price, Robert A.; Palmer, Jeffrey D. (1993). "Phylogenetic Relationships of the Geraniaceae and Geraniales from rbcL Sequence Comparisons". Annals of the Missouri Botanical Garden. 80 (3): 661–671. doi:10.2307/2399852. JSTOR 2399852.
Bakker, Freek T (26 March 2003). "Geraniales (Geranium)". Encyclopedia of Life Sciences. Wiley. doi:10.1038/npg.els.0003716. ISBN 978-0470016176. Missing or empty |title= (help)
Berchtold, Friedrich von; Presl, Jan Svatopluk (1820). O Prirozenosti Rostlin. Prague: Krala Wiljma Endersa.
Candolle, A. P. de (1813). Théorie élémentaire de la botanique, ou exposition des principes de la classification naturelle et de l'art de décrire et d'etudier les végétaux (in French). Retrieved 5 February 2014.
Christenhusz, Maarten JM & Byng, J. W. (2016). "The number of known plants species in the world and its annual increase". Phytotaxa. 261 (3): 201–217. doi:10.11646/phytotaxa.261.3.1.
Cronquist, A (1988) [1968]. The evolution and classification of flowering plants (2nd ed.). Bronx, NY: New York Botanical Garden. ISBN 9780893273323.
Dahlgren, R. M. T. (February 1980). "A revised system of classification of the angiosperms". Botanical Journal of the Linnean Society. 80 (2): 91–124. doi:10.1111/j.1095-8339.1980.tb01661.x.
Dumortier, Barthélemy-Charles (1829). Analyse des familles des plantes: avec l'indication des principaux genres qui s'y rattachent (in French). Tournay: J Casterman. Retrieved 16 January 2016.
Jussieu, Antoine Laurent de (1789). Genera Plantarum, secundum ordines naturales disposita juxta methodum in Horto Regio Parisiensi exaratam (in Latin). Paris: apud viduam Herissant et Theophilum Barrois. OCLC 5161409. Retrieved 9 January 2014.
Pelletier, Bernard (2016) [2012]. Empire Biota: Taxonomy and Evolution (2nd ed.). Lulu. ISBN 978-1-329-87400-8.
Watson, L.; Dallwitz, M.J. (2016). "The families of flowering plants: descriptions, illustrations, identification, and information retrieval". DELTA – DEscription Language for TAxonomy. Archived from the original on 15 May 2016. Retrieved 1 July 2016.
ICN (2012). "International Code of Nomenclature for algae, fungi, and plants". Bratislava: International Association for Plant Taxonomy. Retrieved 2 February 2014.
"Tropicos". Missouri Botanical Garden. 2015. Retrieved 30 December 2015.
IPNI (2015). "The International Plant Names Index".

APG
APG I (1998). "An ordinal classification for the families of flowering plants". Annals of the Missouri Botanical Garden. 85 (4): 531–553. doi:10.2307/2992015. JSTOR 2992015.
APG II (2003). "An Update of the Angiosperm Phylogeny Group Classification for the orders and families of flowering plants: APG II". Botanical Journal of the Linnean Society. 141 (4): 399–436. doi:10.1046/j.1095-8339.2003.t01-1-00158.x.
APG III (2009). "An Update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG III". Botanical Journal of the Linnean Society. 161 (2): 105–121. doi:10.1111/j.1095-8339.2009.00996.x.
APG IV (2016). "An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG IV". Botanical Journal of the Linnean Society. 181 (1): 1–20. doi:10.1111/boj.12385.