Amaranthaceae

Amaranthaceae (incl. Chenopodiaceae)

Recent molecular phylogenetic research strongly suggests the inclusion of Chenopodiaceae in Amaranthaceae (APG 1998-2009). This taxonomy became widely accepted in recent times (see for instance van der Meijden 2005, Stace 2010) and is adopted here as well.

Polycnemum L. is only remotely related to most members of Amaranthaceae s.l. and is possibly better segregated (see for instance Kadereit & al. 2003). Polycnemum majus A. Braun is an extinct native species (Lawalrée 1953, Lambinon & al. 2004) and P. arvense L. was last seen an alien in Belgium in 1903 (Verloove 2006). Similarly, Einadia Raf. is here tentatively accepted as a distinct genus (pending additional studies) but it might well be a member of Australian Chenopodium (Kadereit & al. 2010).

The generic limits of Chenopodium s.l. and related taxa have considerably changed as a result of recent molecular phylogenetic studies (see Fuentes-Bazan & al. 2012 for an overview). Dysphania R. Br. (for species with aromatic, glandular hairs) had already been segregated before on morphological grounds (Mosyakin & Clemants 2002) and this is confirmed by molecular data. Chenopodium, as traditionally understood, finally turns out to fall apart in seven distinct genera and the Belgian representatives (native as well as non-native) should be classified as follows: species preceded by an * are non-native in Belgium

Lipandra Moq. with a single, native species:

L. polysperma (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium polyspermum L.)

Oxybasis Kar. & Kir., with 5 species (4 native, 1 non-native, preceded by an asterisk):

O. chenopodioides (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium chenopodioides (L.) Aellen)

O. glauca (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium glaucum L.)

* O. macrosperma (Hook. f.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium macrospermum Hook. f.)

O. rubra (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium rubrum L.)

O. urbica (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium urbicum L.)

Blitum L., with a single native species and three non-native species (preceded by an asterisk):

Blitum bonus-henricus (L.) Reichenb. (syn.: Chenopodium bonus-henricus L.)

* Blitum capitatum L. (syn.: Chenopodium capitatum (L.) Ambrosi)

* Blitum nuttallianum Schult. (syn.: Monolepis nuttalliana (Schult.) E. Greene)

* Blitum virgatum L. (syn.: Chenopodium foliosum Aschers.)

Dysphania R. Brown, with nine non-native species (preceded by an asterisk):

* D. ambrosioides (L.) Mosyakin et Clemants (syn.: Chenopodium ambrosioides L.)

* D. botrys (L.) Mosyakin et Clemants (syn.: Chenopodium botrys L.)

* D. carinata (R. Brown) Mosyakin et Clemants (syn.: Chenopodium carinatum R. Brown)

* D. cristata (F. Muell.) Mosyakin et Clemants (syn.: Chenopodium cristatum (F. Muell.) F. Muell.)

* D. glomulifera (Nees) P.G. Wilson (syn.: Chenopodium myriocephalum (Benth.) Aellen)

* D. multifida (L.) Mosyakin et Clemants (syn.: Chenopodium multifidum L.)

* D. pseudomultiflora (J. Murr) Verloove et Lambinon (syn.: Chenopodium pseudomultiflorum (J. Murr) Uotila)

* D. pumilio (R. Brown) Mosyakin et Clemants (syn.: Chenopodium pumilio R. Brown)

* D. schraderiana (Schult.) Mosyakin et Clemants (syn.: Chenopodium schraderianum Schult.)

Teloxys Moquin-Tandon

* Teloxys aristata (L.) Moq. (syn.: Chenopodium aristatum L.)

Chenopodiastrum S. Fuentes, Uotila & Borsch, with two native and one non-native species (preceded by an asterisk):

C. hybridum (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium hybridum L.)

C. murale (L.) S. Fuentes, Uotila & Borsch (syn.: Chenopodium murale L.)

* C. simplex (Torrey) S. Fuentes, Uotila & Borsch (syn.: Chenopodium simplex (Torrey) Raf.)

The remaining species belong with Chenopodium s.str. In this new, narrower circumscription it is monophyletic.

In the key beneath this new generic circumscription for Chenopodium and related genera is applied.

1. Leaves scale-like, not developed (i.e. forming succulent sheaths around stem). Stem succulent and jointed, especially when young. Plants confined to saline habitats (native) === Salicornia

1. Leaves not scale-like, well developed. Stem never jointed. Plants of various habitats === 2

2. All leaves opposite. Flowers in axillary whitish heads === Alternanthera

2. Leaves mostly alternate, rarely some or most opposite (in the latter case inflorescences never axillary and whitish) === 3

3. Leaves linear-subulate, (semi-) cylindrical, usually succulent. Plants mostly confined to saline habitats (native) === Salsola and Suaeda

3.Leaves flat, wider and not succulent === 4

4. Hairs branched to stellate === 5

4. Hairs simple or absent ==== 6

5. Hairs stellate. Perianth segments 3-5. Utricle with apical wings, slightly flattened. Monoecious. Leaves shortly petiolate === Axyris

5. Hairs branched. Perianth segments absent or 1. Utricle with winged margins, strongly flattened. Dioecious. Leaves sessile === Corispermum

6. Basal rosette always present at anthesis. Plant entirely glabrous === 7

6. Basal rosette usually absent at anthesis. Indumentum often present === 8

7. Rosette leaves usually 20-50 cm long, stem leaves not hastate at base. Flowers bisexual with 5 tepals. Fruit without teeth or spines. Seeds horizontal === Beta

7. Rosette leaves rarely exceeding 20 cm, stem leaves often hastate at base. Flowers unisexual: male in dense spikes or panicles, female axillary. Fruit often with apical teeth or spines. Seeds vertical === Spinacia

8. Female flowers without perianth but enclosed by two accrescent bracts === 9

8. Female flowers with perianth === 10

9. Fruiting bracteoles almost entirely fused, becoming very hard. Silvery or greyish-white subshrubs with leaves with entire margins (native) === Halimione

9. Fruiting bracteoles usually free to base or fused to above middle, thin and papery or hardening. Leaves green or greenish-white, reddish or rarely more or less silvery (if exceptionally fruiting bracteoles almost entirely fused and becoming very hard, than annual herb with leaf margins irregularly dentate or with basal lobes) === Atriplex

10. Perianth segments of bisexual flowers transversally winged in fruit. Leaves entire, narrowly lanceolate to almost linear, rarely wider than 5 mm === Bassia

10. Perianth segments not transversally winged (but often with a longitudinal keel!). Leaves either wider or not entire at margin === 11

11. All female flowers with 3-5 bracteoles. Perianth brownish and scarious, often regularly dehiscent at maturity === Amaranthus

11. Bracteoles usually absent or inconspicuous. Perianth usually greenish and herbaceous (sometimes succulent and red) === 12

12. Terminal branches of inflorescence without flowers, ending with acicular, sterile branches === Teloxys

12. Terminal branches of inflorescence without acicular, sterile branches === 13

13. Plant (at least in part) with glandular or glandular-vesicular hairs, aromatic === Dysphania

13. Plant glabrous or mealy, without glandular hairs, odourless or foetid, not aromatic === 14

14. Inflorescence in loose monochasial or dichasial cymes. Plant always glabrous (native) === Lipandra

14. Inflorescence in loose or dense spikes or panicles. Plant often mealy (farinose), at least when young, or glabrous === 15

15. Stem unbranched or sparingly branched. Perianth often succulent in fruit, with 1-5 segments. Stigmas 2-4. Seeds always vertical. Basal rosette sometimes present === Blitum

15. Stem usually branched. Perianth never succulent in fruit, with 3-5 segments. Stigmas 2. Seeds vertical and/or horizontal. Basal rosette always absent === 16

16. Flowers usually dimorphic, in lateral flowers with 3 perianth segments. Seeds mostly vertical, some horizontal. Stamens 1-3 === Oxybasis

16. Flowers not dimorphic, all with 5 perianth segments. Seeds all horizontal. Stamens always 5 === 17

17. Foliage and stems glabrous or very sparsely mealy (farinose) when young only. Perianth segments with distinct midvein on inner side === Chenopodiastrum

17. Foliage and stems always distinctly mealy (farinose), mostly persistent at maturity. Perianth segments without distinct midvein on inner side === Chenopodium

Additional aliens: Einadia polygonoides (J. Murr) Wilson (syn.: Chenopodium polygonoides (J. Murr) Aellen) (Aus., wool alien), Polycnemum arvense L. (Euras., wool alien), Ptilotus spec. (Aus., wool alien), Sclerolaena muricata (Moq.) Domin (syn.: Anisacantha muricata Moq., Bassia quinquecuspis (F. Muell.) F. Muell., Chenolea quinquecuspis F. Muell.) (Aus., wool alien) and Spirobassia hirsuta (L.) Freitag et Kadereit (syn.: Bassia hirsuta (L.) Aschers., Kochia hirsuta (L.) Nolte) (Euras., vector unknown).

References

A.P.G. (1998) An ordinal classification for the families of flowering plants. Ann. Missouri Bot. Gard. 85: 531-553.

A.P.G. II (2003) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants: APG II. Bot. J. Linn. Soc. 141: 399-436.

APG III (2009) An update of the Angiosperm Phylogeny Group classification for the orders and families of flowering plants. APG III. Bot. J. Linn. Soc. 161(2): 105-121.

Fuentes-Bazan S., Uotila P. & Borsch T. (2012) A novel phylogeny-based generic classification for Chenopodium sensu lato, and a tribal rearrangement of Chenopodioideae (Chenopodiaceae). Willdenowia 42(1): 5-24. [available online at: http://www.bioone.org/doi/pdf/10.3372/wi.42.42101]

Kadereit G., Borsch T., Weising K. & Freitag H. (2003) Phylogeny of Amaranthaceae and Chenopodiaceae and the evolution of C4 photosynthesis. Int. J. Plant Sci. 164(6): 959-986.

Kadereit G., Mavrodiev E.V., Zacharias E.H. & Sukhorukov A. (2010) Molecular phylogeny of Atripliceae (Chenopodioideae, Chenopodiaceae): implications for systematics, biogeography, flower and fruit evolution, and the origin of C4 photosynthesis. Am. J. Bot. 97(10): 1664-1687.

Lambinon J., Delvosalle L., Duvigneaud J. (avec coll. Geerinck D., Lebeau J., Schumacker R. & Vannerom H. (2004) Nouvelle Flore de la Belgique, du Grand-Duché de Luxembourg, du Nord de la France et des Régions voisines (Ptéridophytes et Spermatophytes). Cinquième édition. Jardin botanique national de Belgique, Meise: CXXX + 1167 p.

Lawalrée A (1953) Chenopodiaceae. In: Robyns W. (ed.), Flore Générale de Belgique, vol. 1, fasc. 2. Jardin Botanique de l’Etat, Bruxelles: 171-236.

Mosyakin S.L. & Clemants S.E. (2002) New nomenclatural combinations in Dysphania R.Br. (Chenopodiaceae): taxa occurring in North America. Ukr. Botan. Journ. 59: 380-385.

Stace C. (2010) New flora of the British Isles, 3th ed.: XXXII + 1232 p. Cambridge University Press.

Van der Meijden R. (2005) Heukels’ Flora van Nederland (23e druk). Wolters-Noordhoff, Groningen: 685 p.

Verloove F. (2006) Catalogue of neophytes in Belgium (1800-2005). Scripta Botanica Belgica 39: 89 p.

Scratchpads developed and conceived by (alphabetical): Ed Baker, Katherine Bouton Alice Heaton Dimitris Koureas, Laurence Livermore, Dave Roberts, Simon Rycroft, Ben Scott, Vince Smith