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.exBercht. & 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]
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 superorderRutiflorae:[13]
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 chloroplastgenerbcL.[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 eudicotclade, specifically in the rosidsubclade.
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]
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.
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]
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. JSTOR2399846.
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.
