‘Cones’ of Casuarina equisetifolia, photographed by Atamari.

Belongs within: Faganae.

The Casuarinaceae, sheoaks, are a family of just under 100 species of small trees that bear a superficial resemblance to conifers, with scale-like leaves and cone-like inflorescences. Species of Casuarinaceae are found in south-east Asia and Australasia; the genus Allocasuarina is restricted to Australia but Casuarina is more widespread. Species of Allocasuarina bear ‘cones’ with thinly woody, prominent bracteoles extending well beyond the body of the cone whereas cones of Casuarina species have thickly woody, convex bracteoles mostly extending only slightly beyond the cone body (Harden 1990).

The fossil record of Casuarinaceae extends at least to the early Palaeocene, with more tentative records going back as far as the Upper Triassic (Collinson et al. 1993).

Why is an oak like a cassowary?
Published 20 January 2016
Beach casuarina Casuarina equisetifolia bearing flowers and cones, copyright Atamari.

When one thinks of the Australian vegetation, one might think of towering eucalypts or hardy acacias. One might contemplate unwelcoming spinifex or vibrant grevilleas. But perhaps few groups of plants are so distinctively Australian as the Casuarinaceae, the casuarinas or she-oaks. Members of this family are also found in south-east Asia and the Pacific Islands, but it is in Australia that they reach their highest diversity.

Casuarinas are also unmistakable. They are flowering plants, but they are wind-pollinated and the flowers are highly reduced, being borne in small clusters or spikes. The clusters of fruits, when mature, look more like a miniature pine cone than anything else. The trees that bear these cones also look a bit like pines themselves, with their narrow photosynthetic branches (cladodes) bearing a superficial resemblance to pine needles. The leaves proper are reduced to tiny teeth arranged around nodes or joints on the branches. The outer layer of the cladodes is composed of a thick cortex which together with the needle-like morphology helps resist desiccation. The name of the family refers to the resemblance of their branches to the hair-like feathers of a cassowary Casuarius. Casuarinas are so distinct from other flowering plants that their affinities were long uncertain, though more recent studies have suggested a relationship to other wind-pollinated trees in families such as the Betulaceae (Steane et al. 2003).

In line with their drought-resistant mien, casuarinas are most often found growing in arid and/or coastal regions. The most widespread species, the beach she-oak Casuarina equisetifolia, is found along coastlines from the Bay of Bengal to Polynesia. Their persistance in harsh conditions is also assisted by the presence of nodules on their roots containing bacteria of the genus Frankia, that function like the Rhizobium in root nodules on legumes to fix nitrogen from the atmosphere. Casuarinas also resemble pine trees in forming a mat around their base of fallen cladodes that restricts the growth of competing vegetation.

Desert oaks Allocasuarina decaisneana, copyright Cgoodwin.

Until relatively recently, casuarinas were all classed in a single genus but most authors now recognise four genera in the family. The most distinctive, whose position as sister to the remaining genera is confirmed by molecular analyses (Steane et al. 2003), is Gymnostoma, which contains eighteen species found from south-east Asia to Queensland and Fiji. Whereas other genera of Casuarinaceae have the stomata on the cladodes hidden within deep longitudinal grooves, Gymnostoma has much shallower grooves on the cladodes and the stomata more or less exposed. As such, it is less resistant to desiccation than the other genera. Gymnostoma has four of these grooves on each cladode, corresponding to four leaf-teeth around each node, so the cladodes also tend to have a squarish cross-section.

The second-most divergent genus, Ceuthostoma, contains just two species found from Palawan and Borneo to New Guinea. Ceuthostoma resembles Gymnostoma in having four teeth around each node, but resembles the remaining two genera, Casuarina and Allocasuarina, in having the stomata hidden within deep grooves. In Casuarina and Allocasuarina, the number of teeth around each node is generally increased (up to twenty in Casuarina), meaning that the cladodes are more rounded than square. As noted by Steane et al. (2003), rounder cladodes with more grooves mean that the opening of each groove is narrower, further improving desiccation resistance. Casuarina and Allocasuarina are most readily distinguished by the appearance of their seeds, which are paler and dull in Casuarina but dark brown or black and shiny in Allocasuarina. Allocasuarina is the most diverse genus of the family, with over fifty species endemic to Australia. Casuarina contains fewer species but is more widespread. Steane et al.‘s molecular analysis suggested a division within Casuarina between two main clades, one of which was restricted to Australia while the other was primarily composed of Indomalesian species (as well as C. equisetifolia which, as noted above, is found damn near everywhere).

Borneo ru Gymnostoma nobile, from

Fossils of Casuarinaceae date back to the Palaeocene epoch, and indicate that the family was more widespread in the past with species known from the Eocene of South America and the Miocene of New Zealand. Casuarinaceae-like pollen is also known from the Palaeogene of southern Africa and Antarctica. The South American species have been assigned to the living genus Gymnostoma; the New Zealand species, though originally assigned to Casuarina, is probably also more closely related to Gymnostoma (Zamaloa et al. 2006). Though dominant in the modern flora, the drought-resistant clade of the other three genera is probably of more recent origin, and has probably only ever been unique to the Australasian region.

And I’ve just realised that I haven’t answered the question in the title to this post. As I noted above, an alternate vernacular name for these trees to ‘casuarina’ is ‘she-oak’. I used to wonder why this should be, seeing as casuarinas look about as unlike oaks as you might care to imagine. A good summary of the solution can be found in this newspaper column from the Western Mail of 1914. Though some have suggested that ‘she-oak’ may be a corruption of an Aboriginal word (despite no such word having been put on record), the more simple explanation is that even if the tree itself doesn’t look like an oak, the wood that comes out of it does.

Systematics of Casuarinaceae

Characters (from the Angiosperm Phylogeny Website): Roots with N-fixing Frankia, rootlets clustered, of limited growth ; flavonols and biflavonoids present, flavones and myricetin absent; nodes 1:1; stomata usually tetracytic, transversely oriented; leaves 4-16-whorled, scale-like, margins entire, stipules absent; plant monoecious or dioecious, inflorescence capitate-spicate, one flower/bract, bracts and bracteoles more or less well-developed; staminate flowers with perianth [“inner bracteoles”] 2, median, androecium 1, filaments incurved in bud, anthers more or less longer than connective; pollen granular layer absent; pistillode absent; carpellate flowers with bracteoles large; perianth absent; gynoecium naked, only abaxial fertile; ovules bitegmic, outer integument 3-4 cells across, inner integument 2-3 cells across, micropyle endostomal, nucellar tracheids present, vascular bundle branched in chalaza; archesporium multicellular, embryo sac with micropylar and chalazal caecum/haustoria; fruit a samara, freed as the much accrescent bracteoles separate; seed coat adnate to pericarp; endosperm anbsent; n = 8-14.

Casuarinaceae [Casuarinales, Casuarinanae, Casuarineae]
|--Casuaroxylon anglica Goeppert & Stache 1855CBH93
|--Haloragacidites harrisii Mildenhall 1980 [=Triorites harrisii]CBH93
| |--C. bicuspidataC46
| |--C. cristata [incl. C. cambagei, C. lepidophloia]H90
| |--C. cunninghamianaB00
| |--C. decaisneanaB05
| |--C. equisetifoliaB88
| | |--C. e. ssp. equisetifoliaH90
| | `--C. e. ssp. incanaH90
| |--C. fraseranaW95
| |--C. glaucaC74
| |--C. huegelianaMH87
| |--C. humilisSM06
| |--C. junghuhnianaKHM91
| |--C. litoreaA80
| |--C. littoralisS00 [=Allocasuarina littoralisH90; incl. C. suberosaH90]
| |--C. muellerianaG76
| |--C. nobilisP88
| |--C. obesaOS04
| |--C. papuanaC78
| |--C. pauperM04
| |--C. strictaC46
| `--C. sumatranaK03
|--A. acutivalvisG04b
|--A. brachystachyaH90
|--A. campestrisB00
|--A. corniculataG04a
|--A. decaisneanaSM96
|--A. decussataM06
|--A. defungensH90
|--A. diminutaH90
| |--A. d. ssp. diminutaH90
| |--A. d. ssp. annectensH90
| `--A. d. ssp. mimicaH90
|--A. distyla [=Casuarina distyla]H90
|--A. eriochlamysG04a
|--A. fraserianaSM06
|--A. glareicolaH90
|--A. gymnantheraH90
|--A. helmsiiG04a
|--A. huegelianaSB04
|--A. humilisOS04
|--A. inophloia [=Casuarina inophloia]H90
|--A. luehmannii [=Casuarina luehmannii]H90
|--A. microstachyaOS04
|--A. nana [=Casuarina nana]H90
|--A. ophioliticaH90
|--A. paludosa [=Casuarina paludosa; incl. C. distyla var. prostrata]H90
|--A. portuensisH90
|--A. rigida [=Casuarina rigida]H90
|--A. rupicolaH90
|--A. simulansH90
|--A. strictaJ91 [=Casuarina strictaH90]
|--A. thuyoidesG04b
|--A. torulosa [=Casuarina torulosa]H90
`--A. verticillata [incl. Casuarina quadrivalvis]H90

*Type species of generic name indicated


[A80] Aoki, J. 1980. A revision of the oribatid mites of Japan. I. The families Phthiracaridae and Oribotritiidae. Bulletin of the Institute of Environmental Science and Technology, Yokohama National University 6 (2): 1–89.

[B05] Beard, J. S. 2005. Drainage evolution in the Lake Disappointment Catchment, Western Australia—a discussion. Journal of the Royal Society of Western Australia 88 (2): 57–64.

[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).

[C46] Cleland, J. B. 1946. Some items of botanical interest in the early history of South Australia. South Australian Naturalist 23 (4): 9–11.

[C78] Clunie, N. M. U. 1978. The vegetation. In: Womersley, J. S. (ed.) Handbooks of the Flora of Papua New Guinea vol. 1 pp. 1–11. Melbourne University Press: Carlton South (Australia).

[CBH93] Collinson, M. E., M. C. Boulter & P. L. Holmes. 1993. Magnoliophyta (‘Angiospermae’). In: Benton, M. J. (ed.) The Fossil Record 2 pp. 809–841. Chapman & Hall: London.

[C74] Crowder, J. P. 1974. Exotic Plant Pests of South Florida. Bureau of Sport Fisheries and Wildlife (USA).

[G04a] Gibson, N. 2004a. Flora and vegetation of the Eastern Goldfields Ranges: part 6. Mt Manning Range. Journal of the Royal Society of Western Australia 87 (2): 35–47.

[G04b] Gibson, N. 2004b. 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.

[G76] Gross, G. F. 1976. Plant-feeding and Other Bugs (Hemiptera) of South Australia. Heteroptera—Part II. Handbook of the Flora and Fauna of South Australia.

[H90] Harden, G. J. (ed.) 1990. Flora of New South Wales vol. 1. New South Wales University Press.

[J91] Jones, D. L. 1991. New taxa of Australian Orchidaceae. Australian Orchid Research 2: 1–208.

[KHM91] Kitchener, D. J., R. A. How & Maharadatunkamsi. 1991. Paulamys sp. cf. P. naso (Musser, 1981) (Rodentia: Muridae) from Flores Island, Nusa Tenggara, Indonesia—description from a modern specimen and a consideration of its phylogenetic affinities. Records of the Western Australian Museum 15 (1): 171–189.

[K03] Kulip, J. 2003. An ethnobotanical survey of medicinal and other useful plants of Muruts in Sabah, Malaysia. Telopea 10 (1): 81–98.

[MH87] Macfarlane, T. D., S. D. Hopper, R. W. Purdie, A. S. George & S. J. Patrick. 1987. Haemodoraceae. Flora of Australia 45: 55–148.

[M06] McCaw, W. L. 2006. Asplenium aethiopicum recolonises karri forest following timber harvesting and burning. Journal of the Royal Society of Western Australia 89 (3): 119–122.

[M04] Mound, L. A. 2004. Australian Thysanoptera—biological diversity and a diversity of studies. Australian Journal of Entomology 43 (3): 248–257.

[OS04] Obbens, F. J., & L. W. Sage. 2004. Vegetation and flora of a diverse upland remnant of the Western Australian wheatbelt (Nature Reserve A21064). Journal of the Royal Society of Western Australia 87 (1): 19–28.

[P88] Polunin, I. 1988. Plants and Flowers of Malaysia. Times Editions: Singapore.

[SB04] Sage, L. W., P. A. Blankendaal, A. Moylett & K. Agar. 2004. The occurrence and impact of Phytophthora cinnamomi in the central-western Avon Wheatbelt bioregion of Western Australia. Journal of the Royal Society of Western Australia 87 (1): 15–18.

[SM06] Semeniuk, C. A., L. A. Milne, P. Ladd & V. Semeniuk. 2006. Pollen in the surface sediments of wetlands in the Becher Point area, southwestern Australia: a baseline for use in interpreting Holocene sequences. Journal of the Royal Society of Western Australia 89 (1): 27–43.

[S00] Siddiqi, M. R. 2000. Tylenchida: Parasites of plants and insects 2nd ed. CABI Publishing: Wallingford (UK).

[SM96] Southgate, R., & P. Masters. 1996. Fluctuations of rodent populations in response to rainfall and fire in a central Australian hummock grassland dominated by Plectrachne schinzii. Wildlife Research 23: 289–303.

Steane, D. A., K. L. Wilson & R. S. Hill. 2003. Using matK sequence data to unravel the phylogeny of Casuarinaceae. Molecular Phylogenetics and Evolution 28: 47–59.

[W95] Wang, Q. 1995. A taxonomic revision of the Australian genus Phoracantha Newman (Coleoptera: Cerambycidae). Invertebrate Taxonomy 9: 865–958.

Zamaloa, M. del C., M. A. Gandolfo, C. C. González, E. J. Romero, N. R. Cúneo & Peter Wilf. 2006. Casuarinaceae from the Eocene of Patagonia, Argentina. International Journal of Plant Sciences 167 (6): 1279–1289.

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