Hologalegina

 Yellow pea bush Sesbania cannabina, copyright Harry Rose.

Belongs within: Papilionoideae.
Contains: Fabeae, Hedysareae, Astragaleae, Lotus.

The Hologalegina are a diverse clade including the majority of temperate-climate herbaceous legumes as well as a variety of woody taxa, united by the loss of one of two copies of the 25-kb inverted repeat in the plastid genome. Within the clade, members of the tribes Loteae, Robinieae and Sesbania are united by molecular analyses as well as by the possession of a distinct determinate root nodule structure. The Robinieae bear pulvinate, paripinnate or imparipinnate leaves with stipules (if present) narrow or modified into thorns.

Milk-vetches, liquorice and locoweeds
Published 1 March 2011
Flowers of the weeping broom Carmichaelia stevensonii, from here.

Far in the distant past, I commented on a review of the New Zealand brooms of the genus Carmichaelia. The New Zealand brooms were placed by Wagstaff et al. (1999) in the tribe Galegeae, which includes more than 3000 species around the world. The vast majority of these are placed in the genus Astragalus, milk-vetches, which may itself have well over 2500 species and be the largest recognised genus of flowering plant. Other significant members of the Galegeae include the locoweeds (Oxytropis) and Glycyrrhiza, a small genus of less than twenty species that nevertheless deserves praise for being the source of liquorice*.

*I am reliably informed that there are people in this world who do not appreciate the flavour of liquorice. There is simply no accounting for taste.

Flowers of Astragalus monspessulanus, from here.

However, despite its wide recognition, phylogenetic studies of recent years have been unanimous in declaring the Galegeae polyphyletic. It is true that the vast majority of galegeans remain in a clade, with only a few relatively minor genera placed elsewhere (Wojciechowski et al. 2000). The problem is that one of those ‘minor genera’ happens to be Galega itself, the type genus and therefore sine qua non of the tribe, which is more closely related to the chickpeas of the genus Cicer. Wojciechowski et al. (2000) circumvented this issue by recognising a clade called Hologalegina, uniting the ‘Galegeae’ with other tribes such as Hedysareae, Trifolieae, Fabeae, Loteae and Robinieae. Within the Hologalegina, the taxa previously assigned to the Galegeae all belong to what is called the IRLC, the ‘Inverted Repeat-Lacking Clade’. The name of this clade refers to the loss of a copy of the 25 kb inverted repeat in the chloroplast genome that is otherwise found in almost all other land plants. Members of the IRLC are largely herbaceous, and many of the most commercially significant legumes (such as peas, beans, lentils, clover and alfalfa) belong to this clade.

Liquorice, Glycyrrhiza glabra, photographed by J. C. Schou. The flavour comes from the roots.

With the exception of the aforementioned Glycyrrhiza (which is another phylogenetically distant genus from other galegeans), most of the ‘Galegeae’ are not commercially grown. They may be commercially eradicated: species of Oxytropis get their name of ‘locoweed’ from their toxic effects on grazing livestock, and various species of Astragalus and Swainsona are also known for their toxicity. Some species are cultivated as garden plants, such as the weeping broom Carmichaelia stevensonii shown in the top photo (though many places still list it under the older name of Chordospartium stevensonii) and the kaka beak Clianthus puniceus. One particular galegean is widely regarded as the stuff that gardeners’ dreams are made of, despite being a notoriously difficult plant to grow. My partner tells me that when he lived in Kalgoorlie as a child, tourist buses would regularly appear outside his house just so that their passengers could see this plant flowering in the front yard. I speak, of course, of Swainsona formosa, Sturt’s desert pea:

Photographed by Marj Kibby.
Systematics of Hologalegina
<==Hologalegina [Galegeae]CP13
    |--+--GlycyrrhizaCP13
    |  |    |--G. acanthocarpaG04
    |  |    |--G. glabraKU02
    |  |    `--G. lepidotaCP13
    |  `--+--+--FabeaeCP13
    |     |  `--+--HedysareaeCP13
    |     |     `--AstragaleaeCP13
    |     `--+--Callerya reticulataCP13
    |        `--WisteriaCP13
    |             |--W. frutescensCP13
    |             `--W. sinensisB00
    `--+--LoteaeCP13
       |    |--+--HippocrepisCP13
       |    |  |    |--H. bicontortaD17
       |    |  |    |--H. ciliataPT98
       |    |  |    |--H. comosaH91
       |    |  |    |--H. multicaulis Batt. & Trab. 1911D17
       |    |  |    `--H. unisiliquosaCP13
       |    |  `--+--Securigera variaCP13
       |    |     `--CoronillaCP13
       |    |          |--C. coronataCP13
       |    |          |--C. emerus [incl. C. emeroides]H91
       |    |          |--C. montanaC55
       |    |          |--C. scorpioidesPT98
       |    |          `--C. variaC06
       |    `--+--+--LotusCP13
       |       |  `--AnthyllisCP13
       |       |       |--A. dilleniiB28
       |       |       |--A. hermanniae Linnaeus 1753PL04
       |       |       |--A. montanaC55
       |       |       `--A. vulnerariaCP13
       |       |            |--A. v. ssp. vulnerariaPT98
       |       |            `--A. v. ssp. rubrifloraPT98
       |       `--+--OrnithopusCP13
       |          |    |--O. compressusCP13
       |          |    |--O. perpusillusPAE98
       |          |    |--O. pinnatusH93
       |          |    `--O. sativus [incl. O. roseus]H93
       |          `--+--Hammatolobium kremerianumCP13
       |             `--DorycniumCP13
       |                  |--D. pentaphyllumCP13 [incl. D. herbaceumH91]
       |                  `--D. suffruticosumC55
       `--+--Sesbania [Sesbanieae]CP13
          |    |--S. aegyptiacaB88
          |    |--S. bispinosaPP07
          |    |--S. brachycarpaM99
          |    |--S. campylocarpaM99
          |    |--S. cannabinaB00
          |    |--S. erubescensM99
          |    |--S. exaltataH93
          |    |--S. formosaLK14
          |    |--S. javanicusB00
          |    |--S. pachycarpaCV06
          |    |    |--S. p. ssp. pachycarpaCV06
          |    |    `--S. p. ssp. dinteranaCV06
          |    |--S. puniceaH06
          |    |--S. rostratumE92
          |    |--S. simpliciusculaM99
          |    |    |--S. s. var. simipliciusculaLK14
          |    |    `--S. s. var. fitzroyensisLK14
          |    `--S. tomentosaCP13
          `--RobinieaeCP13
               |--Hebestigma cubenseCP13
               `--+--Lennea modestaCP13
                  |--+--Hybosema ehrenbergiiCP13
                  |  `--+--Poitea glyciphyllaCP13
                  |     `--GliricidiaCP13
                  |          |--G. brenningiiCP13
                  |          |--G. maculataSK02
                  |          `--G. sepiumMM96
                  `--+--RobiniaCP13
                     |    |--R. hispidaA98
                     |    |--R. neomexicanaRJ11
                     |    |--R. pseudoacaciaCP13
                     |    `--R. viscosaA98
                     `--+--Olneya tesotaCP13
                        |--+--Poissonia hypoleucaCP13
                        |  `--Sphinctospermum constrictumCP13
                        `--+--CoursetiaCP13
                           |    |--C. arboreaMM96
                           |    `--C. glandulosaCP13
                           `--+--Genistidium dumosumCP13
                              `--Peteria thompsonaeCP13

*Type species of generic name indicated

References

[A98] Anonymous. 1798. Sur le Robinia viscosa, et la substance résineuse qu’il produit. Bulletin des Sciences, par la Societé Philomathique de Paris 1 (21): 161–162.

[B28] Betrem, J. G. 1928. Monographie der Indo-Australischen Scoliiden mit zoogeographischen Betrachtungen. H. Veenman & Zonen: Wageningen.

[B88] Bouček, Z. 1988. Australasian Chalcidoidea (Hymenoptera): A biosystematic revision of genera of fourteen families, with a reclassification of species. CAB International: Wallingford (UK).

[B00] Braby, M. F. 2000. Butterflies of Australia: their identification, biology and distribution vol. 1. CSIRO Publishing: Collingwood (Victoria).

[C55] Candolle, A. de. 1855. Géographie Botanique Raisonée: Ou exposition des faits principaux et des lois concernant la distribution géographique des plantes de l’époque actuelle vol. 1. Librairie de Victor Masson: Paris.

[CP13] Cardoso, D., R. T. Pennington, L. P. de Queiroz, J. S. Boatwright, B.-E. Van Wyk, M. F. Wojciechowski & M. Lavin. 2013. Reconstructing the deep-branching relationships of the papilionoid legumes. South African Journal of Botany 89: 58–75.

[C06] Cheeseman, T. F. 1906. Manual of the New Zealand Flora. John Mackay, Government Printer: Wellington.

[CV06] Craven, P., & P. Vorster. 2006. Patterns of plant diversity and endemism in Namibia. Bothalia 36 (2): 175–189.

[D17] Diels, L. 1917. Beiträge zur Flora der Zentral-Sahara und ihrer Pflanzengeographie. Nach der Sammelausbeute des Freiherrn Hans Geyr von Schweppenburg. Botanische Jahrbücher für Systematik, Pflanzengeschichte und Pflanzengeographie 54 (Beiblatt 120): 51–155.

[E92] Eady, R. R. 1992. The dinitrogen-fixing bacteria. In: Balows, A., H. G. Trüper, M. Dworkin, W. Harder & K.-H. Schleifer (eds) The Prokaryotes: A handbook on the biology of bacteria: Ecophysiology, isolation, identification, applications 2nd ed. vol. 1 pp. 534–553. Springer-Verlag: New York.

[G04] Gibson, N. 2004. Flora and vegetation of the Eastern Goldfields Ranges: part 7. Middle and South Ironcap, Digger Rock and Hatter Hill. Journal of the Royal Society of Western Australia 87 (2): 49–62.

[H06] Henderson, L. 2006. Comparisons of invasive plants in southern Africa originating from southern temperate, northern temperate and tropical regions. Bothalia 36 (2): 201–222.

[H93] Hickman, J. C. (ed.) 1993. The Jepson Manual: Higher Plants of California. University of California Press: Berkeley (California).

[H91] Hubálek, Z. 1991. Biogeographic indication of natural foci of tick-borne infections. In: Dusbábek, F., & V. Bukva (eds) Modern Acarology: Proceedings of the VIII International Congress of Acarology, held in České Budĕjovice, Czechoslovakia, 6–11 August 1990 vol. 1 pp. 255–260. SPB Academic Publishing: The Hague.

[KU02] Khanjani, M., & E. A. Ueckermann. 2002. The stigmaeid mites of Iran (Acari: Stigmaeidae). International Journal of Acarology 28 (4): 317–339.

[LK14] Lyons, M. N., G. J. Keighery, L. A. Gibson & T. Handasyde. 2014. Flora and vegetation communities of selected islands off the Kimberley coast of Western Australia. Records of the Western Australian Museum Supplement 81: 205–244.

[M99] Matthews, M. 1999. Heliothine Moths of Australia: A guide to bollworms and related noctuid groups. CSIRO Publishing.

[MM96] Mound, L. A., & R. Marullo. 1996. The thrips of Central and South America: an introduction (Insecta: Thysanoptera). Memoirs on Entomology, International 6: 1–487.

[PP07] Pandey, R. P., & P. M. Padhye. 2007. Studies on phytodiversity of Arid Machia Safari Park-Kailana in Jodhpur (Rajasthan). Bulletin of the Botanical Survey of India 49: 15–78.

[PAE98] Pakeman, R. J., J. P. Attwood & J. Engelen. 1998. Sources of plants colonizing experimentally disturbed patches in an acidic grassland, in eastern England. Journal of Ecology 86: 1032–1041.

[PT98] Panitsa, M., & D. Tzanoudakis. 1998. Contribution to the study of the Greek flora: flora and vegetation of the E Aegean islands Agathonisi and Pharmakonisi. Willdenowia 28: 95–116.

[PL04] Pohl, G., & I. Lenski. 2004. Zur Verbreitung und Vergesellschaftung von Pennisetum orientale Rich. in Nordeuböa (Griechenland) (Poaceae, Paniceae). Senckenbergiana Biologica 83 (2): 209–223.

[RJ11] Rising, J. D., A. Jaramillo, J. L. Copete, P. G. Ryan & S. C. Madge. 2011. Family Emberizidae (buntings and New World sparrows). In: Hoyo, J. del, A. Elliott & D. A. Christie (eds) Handbook of the Birds of the World vol. 16. Tanagers to New World Blackbirds pp. 428–683. Lynx Edicions: Barcelona.

[SK02] Sumithramma, N., A. R. V. Kumar, K. Chandrashekara & D. Rajagopal. 2002. Plant selection for nesting by Oecophylla smaragdina, Hymenoptera: Formicidae: do physical features affect the choice of the plant? Journal of the Bombay Natural History Society 99 (3): 408–412.

Wagstaff, S. J., P. B. Heenan & M. J. Sanderson. 1999. Classification, origins, and patterns of diversification in New Zealand Carmichaelinae (Fabaceae). American Journal of Botany 86 (9): 1346–1356.

Wojciechowski, M. F., M. J. Sanderson, K. P. Steele & A. Liston. 2000. Molecular phylogeny of the “temperate herbaceous tribes” of papilionoid legumes: a supertree approach. In: Herendeen, P. S., & A. Bruneau (eds) Advances in Legume Systematics vol. 9 pp. 277–298. Royal Botanic Gardens, Kew.

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