Table 1.
Orders, families and genera investigated * | Presence of petals | Petals retarded in growth | Primary obdiplostemony | Secondary obdiplostemony | Size of antesepalous vs. alternisepalous stamens | Alternisepalous staminodes | Antesepalous staminodes | Presence of diplostemony in family | Presence of obhaplostemony in family | Presence of haplostemony in family | Carpel number and position if isomerous | Reference(s) |
---|---|---|---|---|---|---|---|---|---|---|---|---|
Oxalidales | ||||||||||||
Brunelliaceae Brunellia | – | n/a | type II? | – | > | – | – | + | – | – | (4–)5 alternisepalous | Matthews and Endress (2002) † |
Cephalotaceae Cephalotus | – | n/a | type II | – | > | – | – | – | – | – | 6 alternisepalous | Baillon (1865), Matthews and Endress (2002) †; Fig. 2D–F |
Connaraceae Connarus, Cnestis, etc. | + | Yes | – | type I | >/= | + (Connarus, etc.) | – | ? | – | + | 5 antepetalous | Breteler (1989), Matthews & Endress (2002), Fig. 2A |
Cunoniaceae Acsmithia, Davidsonia, Geissois, etc. | +/– | Yes | type II?‡ | type I | > | – | – | + | – | – | 2–3(4) antepetalous | Dickison (1975), Moody and Hufford (2000), Matthews et al. (2001), Schönenberger et al. (2001), Matthews and Endress (2002), Fig. 2C |
Oxalidaceae Oxalis, Biophytum, Averrhoea | + | yes | – | type I | > | + (Averrhoea) | – | – | – | – | 5 antepetalous | Matthews and Endress (2002), Fig. 1A–C |
Elaeocarpaceae§ Platytheca, Sericolea, Elaeocarpus | + | no | type I? | type I | > | – | – | – | – | + (pairs or triplets) | 2–3 | van Heel (1966), Ronse De Craene and Smets (1996), Matthews and Endress (2002) |
Saxifragales | ||||||||||||
Crassulaceae Monanthes, Semperivum, Kalanchoe etc. | + | yes | – | type I | > | + (Sempervivum) | – | – | – | + | 4–5 antepetalous | Eichler (1878), Gelius (1967), Ronse De Craene (2016) |
Haloragaceae Haloragis | +/– | yes | – | type I | = | – | – | + | – | + | 2–4 antepetalous | Eichler (1878), L. Ronse De Craene, unpubl. data |
Iteaceae Pterostemon | + | ? | – | type I? | > | + | – | – | – | – | 5 antepetalous? | Engler (1930) |
Aphanopetalaceae Aphanopetalum | +/– | + | type II? | type I? | > | – | – | – | – | – | 4 alternisepalous | Dickison (1975), Dickison et al. (1994) |
Penthoraceae Penthorum | – | n/a | type II? | – | < | – | – | – | – | – | 5–6 alternisepalous | Baillon (1865), L. Ronse De Craene, unpubl.data |
Tetracarpaeaceae Tetracarpaea | + | yes | – | type I | > | – | – | – | – | + | 4–5 antepetalous | Hils et al. (1988) |
Saxifragaceae Rodgersia, Chrysosplenium | +/– | yes | type II? | type I | > | – | – | – | + | + | 2 | Engler (1930), Gelius (1967), Klopfer (1968, 1970, 1973), Ronse De Craene et al. (1998c) |
Malpighiales | ||||||||||||
Euphorbiaceae Crozophora, Capuronia, Jatropha | – (+) | yes | type I? | – | >/< | – | – | + | + | + (Crozophora) | 2–3 | Eichler (1878), Michaelis (1924), Venkata Rao and Ramalakshmi (1968) |
Linaceae Durandea, Hugonia | + | yes | – | type I | >/< | + | – | + | – | + | 3–5 antepetalous | Narayana (1964), Schewe et al. (2011), Matthews and Endress (2011) |
Malpighiaceae Malpighia, Galphimia, etc. | + | ? | – | type I | > | – | – | – | – | + | 2–3–(–5) antepetalous | Eichler (1878), Ronse De Craene (2010) |
Ochnaceae Ouratea, Sauvagesia | + | no | – | type I | > | + (Sauvagesia) | – | + (Ouratea) | – | + | 3–5 antepetalous | Farrar and Ronse De Craene (2013), Fig. 1G–I |
Rhizophoraceae Anopyxis, Bruguieria, Ceriops, Crossostylis | + (–) | yes | ? | type I | >, =,< | – | – | + | – | – | 2–5(–20) ¶ antepetalous/antesepalous | Baillon (1862), Juncosa and Tomlinson (1987), Juncosa (1988), Setoguchi et al. (1996), Matthews and Endress (2011) |
Fabales | ||||||||||||
Surianiaceae Suriania | + | yes | – | type I | > | + (occ. incomplete) | – | – | – | 5 antepetalous | Tschunko and Nickerson (1976), Bello et al. (2007) | |
Ericales | ||||||||||||
Clethraceae Clethra | + | no | - | type I | > | – | – | + | – | – | 3–4 | Leins (1964), Caris (2013) |
Diapensiaceae Galax, Pyxidanthera, etc. | + | yes | - | type I | > | + | – | – | – | + (Pyxidanthera) | 3 | Caris (2013) |
Ericaceae Erica, Monotropa, etc. | + | yes/no | – | type I | = | – | – | + | – | + | 4–5 antepetalous | Payer (1857), Leins (1964), Caris (2013) |
Sapotaceae Palaquium, Sideroxylon, etc. | + | no | – | type III | < | – | + | + | + | – | 4–5–∞ antesepalous/antepetalous | Caris (2013), Kümpers et al. (2016), Fig. 3D–F |
Styracaceae Styrax, Pterostyrax | + | no | – | type I | >/= | – | – | + | – | + (Pamphilia) | 2–5 antepetalous | Dickison (1993), Caris (2013) |
Geraniales | ||||||||||||
Geraniaceae Geranium, Erodium, etc. | + | yes | – | type I | > | + (Erodium) | – | – | – | + | 5 antepetalous | Endress (2010), Ronse De Craene et al. (1993) |
Melanthiaceae Francoa, Greyia | + | yes | – | type I | > | – | – | – | – | + | 4–5 antepetalous | Ronse De Craene and Smets (1999) |
Vivianaceae Viviania, Rhynchotheca | + (–) (Rhynchotheca) | yes? | – | type I | > | – | – | – | – | + | (2–)3–5 antepetalous | Weigend (2005), L. Ronse De Craene, pers. observ. |
Zygophyllales | ||||||||||||
Zygophyllaceae Tribulus, Balanites, etc | + | yes | – | type I | > | – | – | – | – | ? | 4–5 antepetalous | Payer (1857), Ronse De Craene and Smets (1995) |
Malvales | ||||||||||||
Malvaceae -Byttnerioideae Melochia, Byttneria, Theobroma, Lasiopetalum | + | yes | type I | – | < | – | + | – | + (Hermannia, Melochia, Lasiopetalum) | – | 5 antepetalous, antesepalous (when obhaplostemony) | van Heel (1966), Payer (1857), Ronse De Craene and Smets (1995), Fig. 3A–C |
Malvaceae - Grewioideae Triumfetta annua, T. bartriania | + | yes | – | type I? | > | – | – | – | – | + | 5 antepetalous | van Heel (1966) |
Caryophyllales | ||||||||||||
Asteropeiaceae Asteropeia | + | + | – | type I? | > | – | – | – | – | – | 3 | Schatz et al. (1999), L. Ronse De Craene, unpubl. data |
Caryophyllaceae Gypsophila, Dianthus, etc. | +/– | yes | – | type I | > | + | – | + | + (Colobanthus) | + | 2–3–5 antepetalous | Eckert (1966), Rohweder (1967, 1970), Ronse De Craene et al. (1998b), Ronse De Craene (2013) |
Nepenthaceae Nepenthes | – | n/a | type II | – | > | – | – | – | + | – | 4 alternisepalous | L. Ronse De Craene, unpubl. data |
Tamaricaceae Myricaria | + | yes | – | type 1 | > | – | – | –? | – | + | 3–5 antepetalous | Payer (1857), L. Ronse De Craene, unpubl. data |
Myrtales | ||||||||||||
Combretaceae Terminalia, Thiloa, Combretum | +/– | yes | – | type II | </> | + (Thiloa) | +/– | ? | – | + | (2–3)5, antepetalous | Payer (1857), Eichler (1878), Ronse De Craene (2010), L. Ronse De Craene, unpubl. data |
Lythraceae Decodon, etc. | +/– | yes | – | type I | > | – | – | + | + | + | 2–6 antepetalous/antesepalous (Lagerstroemia) | Mayr (1969), Tobe et al. (1998) |
Onagraceae Ludwigia, Epilobium | + (–) | yes | – | type I | > | – | + (Clarkia) | + | – | + (Ludwigia) | 2–4(5) antepetalous | Eichler (1878), Mayr (1969) |
Myrtaceae Heteropyxis | + | no | – | type I? | ? | – | – | + (Psiloxylon) | + | + | 2–3 (–5) antepetalous | Eichler (1878), Schmid (1980) |
Sapindales | ||||||||||||
Meliaceae Cedrella, Melia | + | no | – | type I | > | – | – | + | – | ? | 5 antepetalous | Payer (1857), Eichler (1878) |
Rutaceae Ruta, Dictamnus, etc. | + | no | – | type I | > | +(Agathosma, Flindersia) | – | + | – | + | 2–3(5) antepetalous | Payer (1857), Eckert (1966), Lal and Narayana (1994), Wei et al. (2011) |
Simaroubaceae Ailanthus, Quassia, etc. | + | no | – | type I | > | + | – | + | – | + | 2–3(5) antepetalous | Payer (1857), Eichler (1878), Eckert (1966) |
Cornales | ||||||||||||
Loasaceae Schismocarpus | + | no | – | type I | > | + | – | – | – | + | 2–3–5 antepetalous | Hufford (1989), Moody and Hufford (2000), Ronse De Craene (2010) |
Crossosomatales | ||||||||||||
Geissolomataceae Geissoloma** | – | n/a | type II? | – | > | – | – | – | – | – | 4 alternisepalous | Matthews & Endress (2005a) |
Strasburgeriaceae (Strasburgeria) | + | no | – | type I | > | – | – | – | – | – | 5 antepetalous | Matthews & Endress (2005a) |
Cucurbitales | ||||||||||||
Anisophylleaceae | + | ? | – | + | > | – | – | ? | – | – | 3–4 antepetalous | Matthews et al. (2001), Matthews and Endress (2004) |
*The following families were excluded although they were reported to be obdiplostemonous [see also Ronse De Craene and Smets (1995): Anacardiaceae, Eichler (1878), Bachelier and Endress (2009), Burseraceae, Bachelier and Endress (2009)], Humiriaceae [with increased stamen number: Narayana and Rao (1977b) – although the condition is marginal with either antesepalous or antepetalous carpels: Kubitzki (2014), Celastraceae [with antepetalous staminodes in Parnassia and possibly Brexia (Eichler, 1878; Matthews and Endress, 2005b)].
†Described as diplostemony by Matthews and Endress (2002).
‡No observations of early stamen initiation are young enough to give undisputed evidence of early obdiplostemony type II [e.g. Moody and Hufford (2000), contrary to Cephalotus (Fig. 2D–F).
§van Heel (1966) describes a condition resembling obdiplostemony in Sericolea. However, the number of antepetalous stamens is variable, with stamens occurring singly or in pairs. Other Elaeocarpaceae have antesepalous triplets or a multistaminate androecium with stamen groups in antepetalous sectors. Our unpublished data on Elaeocarpus cyaneus show a simultaneous initiation of the antepetalous stamen fascicles with the antesepalous stamens. This is correlated with a star-shaped floral apex (cf. Fig. 3A–C).
¶Antepetalous in Macarisia according to Baillon (1862), but antesepalous in Crossostylis according to Setoguchi et al. (1996).
**Matthews and Endress (2005a) described the flower as diplostemonous but the alternisepalous stamens are in outer position similar to other obdiplostemonous apetalous flowers.