Research Article |
Corresponding author: Isabel Larridon ( i.larridon@kew.org ) Academic editor: Elmar Robbrecht
© 2023 Fitiavana Rasaminirina, Isabel Larridon.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Rasaminirina F, Larridon I (2023) The genera of Cyperaceae of Madagascar. Plant Ecology and Evolution 156(3): 276-310. https://doi.org/10.5091/plecevo.98847
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Background and aims – The rise of DNA sequencing in systematics has brought more understanding of the Cyperaceae family worldwide. Through these studies, it has been possible to delineate major clades and classify its species into subfamilies, tribes, and genera. Today, we have a good understanding of the species diversity and geographic distribution of the genera. However, in the case of Madagascar, the only complete taxonomic treatment of Cyperaceae dates from 1937. Although recent monographs exist for some genera in Madagascar, the taxonomic treatment of the Cyperaceae of Madagascar has not been updated until now. Hence, the present work aims to produce an updated treatment at the generic level including descriptions and an identification key of all Cyperaceae genera in Madagascar.
Material and methods – Books and scientific articles containing descriptions of the genera of Cyperaceae of Madagascar, and information on their ecology and distribution were consulted, as well as herbarium specimens, collections, and data available from online herbaria and aggregator portals.
Key results – Madagascar is very rich in Cyperaceae. The species of the Cyperaceae family on the island are distributed in 24 genera, including one endemic genus, i.e. Trichoschoenus. Ten genera are monotypic on the island. Sixteen of the 29 genera described in the Flore de Madagascar remain accepted but the others are now either synonyms of Cyperus, or no longer apply because they are absent from Madagascar, such as Scirpus. Regarding their habitat and ecology, some genera are specific to a particular habitat, like Costularia, while others that are very broadly distributed, like Cyperus and Scleria.
Conclusion – Our study provides an updated overview of the genera of Cyperaceae of Madagascar, including an identification key, descriptions, and illustrations.
Cyperaceae, genera, identification, Madagascar, taxonomy
Madagascar, with its area of 592,000 km2, is home to ca 310 species of the Cyperaceae family (
Numerous scientific works documenting each genus of Cyperaceae occurring Madagascar have been studied. These include Sedges Genera of Africa and Madagascar by
To verify the described characteristics, herbarium specimens were consulted in the following herbaria: TAN, TEF, and the herbarium of the University of Antananarivo in Madagascar including recent collections collected by the first author and Cyperaceae recently collected by the
Kew Madagascar Conservation Centre (KMCC)
team in different localities of Madagascar. In addition, collections available from online herbaria (e.g. Kew Herbarium Catalogue of the Royal Botanic Gardens, Kew (K), Missouri Botanical Gardens’ Tropicos.org (MO), and the Vascular plants catalogue of the Muséum national d’Histoire naturelle (P), and aggregator portals such as Plants of the World Online and JSTOR Global Plants were also consulted. For studies of the ecology and distribution of each genus, online platforms such as
The identification key to subfamilies, tribes, subtribes, and genera follows the classification of
Twenty-four genera of Cyperaceae currently exist in Madagascar including one endemic genus, i.e. Trichoschoenus J.Raynal. Ten genera are monospecific on the island, i.e. Actinoschoenus Benth., Bolboschoenus (Asch.) Palla, Carpha Banks & Sol. ex R.Br., Cladium P.Browne, Coleochloa Gilly, Diplacrum R.Br., Ficinia Schrad., Hypolytrum Pers., Lepironia Pers., and Trichoschoenus. Table
Overview of the genera occurring in Madagascar and their subfamily and tribal placement following
Subfamily | Tribe | Subtribe | Genus |
Mapanioideae | Chrysitricheae | Lepironia Pers. | |
Hypolytreae | Hypolytrum Pers. | ||
Cyperoideae | Trilepideae | Coleochloa Gilly | |
Cladieae | Cladium P.Browne | ||
Bisboeckelereae | Diplacrum R.Br. | ||
Sclerieae | Scleria P.J.Bergius | ||
Carpheae | Carpha Banks & Sol. ex R.Br. | ||
Schoeneae | Oreobolinae | Costularia C.B.Clarke | |
Lepidospermatinae | Machaerina Vahl | ||
Rhynchosporeae | Rhynchospora Vahl | ||
Cariceae | Carex L. | ||
Eleocharideae | Eleocharis R.Br. | ||
Abildgaardieae | Bulbostylis Kunth | ||
Trichoschoenus J.Raynal | |||
Actinoschoenus Benth. | |||
Abildgaardia Vahl | |||
Fimbristylis Vahl | |||
Bolboschoeneae | Bolboschoenus (Asch.) Palla | ||
Fuireneae | Fuirena Rottb. | ||
Schoenoplecteae | Schoenoplectus (Rchb.) Palla | ||
Pseudoschoeneae | Schoenoplectiella Lye | ||
Cypereae | Ficiniinae | Isolepis R.Br. | |
Ficinia Schrad. | |||
Cyperinae | Cyperus L. |
Sixteen of the 29 genera described in the Flore de Madagascar are still accepted and found on the island, i.e. Actinoschoenus, Bulbostylis Kunth, Carex L., Carpha, Cladium, Costularia C.B.Clarke, Cyperus L., Diplacrum, Eleocharis R.Br., Ficinia, Fimbristylis Vahl, Fuirena Rottb., Hypolytrum, Lepironia, Rhynchospora Vahl, and Scleria P.J.Bergius. Others are now either synonyms of the genus Cyperus, such as Ascolepis Nees ex Steud., Lipocarpha R.Br., Mariscus Vahl, Pycreus P.Beauv., and Torulinium Desv., or no longer apply because the genus is absent from Madagascar, as in the case of Scirpus Tourn. ex L. Many species previously placed in Scirpus have since been moved to other genera such as Bolboschoenus and Schoenoplectiella Lye. Table
Comparison of the accepted genera in Madagascar according to
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Fimbristylis Vahl p.p. | Abildgaardia Vahl |
Actinoschoenus Benth. | Actinoschoenus Benth. |
Bulbostylis Kunth | Bulbostylis Kunth |
Carex L. | Carex L. |
Schoenoxiphium Nees | |
Carpha Banks & Sol. ex R.Br. | Carpha Banks & Sol. ex R.Br. |
Cladium P.Browne p.p. | Cladium P.Browne |
Eriospora Hochst. ex A.Rich. | Coleochloa Gilly |
Costularia C.B.Clarke | Costularia C.B.Clarke |
Ascolepis Nees ex Steud. | Cyperus L. |
Courtoisina Nees (as Courtoisia) | |
Cyperus L. | |
Kyllinga Rottb. (as Kyllingia) | |
Lipocarpha R.Br. | |
Mariscus Gaertn. | |
Pycreus P.Beauv. | |
Queenslandiella Domin | |
Remirea Aubl. | |
Torulinium Desv. | |
Diplacrum R.Br. | Diplacrum R.Br. |
Eleocharis R.Br. (as Heleocharis) | Eleocharis R.Br. |
Scirpus L. p.p. | |
Ficinia Schrad. | Ficinia Schrad. |
Fimbristylis Vahl p.p. | Fimbristylis Vahl |
Fuirena Rottb. | Fuirena Rottb. |
Hypolytrum Pers. | Hypolytrum Pers. |
Scirpus L. p.p. | Isolepis R.Br. |
Lepironia Pers. | Lepironia Pers. |
Cladium P.Browne p.p. | Machaerina Vahl |
Rhynchospora Vahl | Rhynchospora Vahl |
Scirpus L. p.p. | Schoenoplectiella Lye |
Scirpus L. p.p. | Schoenoplectus (Rchb.) Palla |
Taxon not included but Scirpus L. p.p. | Bolboschoenus (Asch.) Palla |
Taxon not yet described | Trichoschoenus J.Raynal |
Acriulus Ridl. | Scleria P.J.Bergius |
Scleria P.J.Bergius |
With respect to their habitat and ecology, there are those that are specific to one habitat, such as the genus Costularia, and others that are very widely distributed, such as the genera Cyperus and Scleria. Our study provides an updated overview of the genera of Cyperaceae of Madagascar, including an identification key, descriptions, and illustrations.
1. | Basic inflorescence unit (= spicoid) usually comprising 2, strongly keeled and opposite basal bracts (rarely 1 and unkeeled), with a further (0–)1–13(–100) scale‐like bracts, the bracts subtending 1 stamen, the whole unit with a terminal pistil | 2 (Mapanioideae) |
– | Basic inflorescence unit (= spikelet) consisting of a rachilla bearing few to many glumes that may or may not subtend a flower | 3 (Cyperoideae) |
2. | Leaf blade absent; inflorescence always a single spike; predominantly in temperate and subtemperate heathlands and swaps | Chrysitricheae (Lepironia) |
– | Leaf blade present, inflorescence variable, relatively broad with 3 prominent veins; usually forest plants | Hypolytreae (Hypolytrum) |
3. | All florets unisexual | 4 |
– | At least some florets bisexual | 7 |
4. | Female (or more rarely bisexual) spikelet enclosed in a sac-like prophyll (thus forming the utricle or perigynium), usually clustering to form a spike or part of a bisexual spike | Cariceae (Carex) |
– | Female spikelet not enclosed in a sac-like prophyll or perigynium | 5 |
5. | Inflorescence paniculate, densely spiked with many spindly spikelets and with 1 or 2 florets; nutlet fusiform with long beak, surrounded at the base by 3 small long fimbriate scales opposite the sides of the nutlet; plant of granitic rocks | Trilepideae (Coleochloa) |
– | Inflorescence variable, but not as above; nutlet not surrounded at base by 3 small long fimbriate scales opposite the sides of the nutlet | 6 |
6. | Inflorescence a group of sessile to short-stalked spikelets developing in leaf axils (except the lowest 1 or 2 leaves); all spikelets unisexual, the female spikelet 1-flowered, with no evidence of the other sex, the top of the rachilla completely reduced; male flower usually 1-stamened; contraligule usually undeveloped; nutlet surrounded by two subterminal and opposite glumes | Bisboeckelereae (Diplacrum) |
– | Inflorescence variable, often paniculate or spike-like on upper part of stem; spikelet bisexual with 1 female floret; if all spikelets unisexual, then female spikelets with reduced male flowers, or with reduced rachilla apex; male flower usually with 3 stamens; contraligule usually well developed; nutlet usually large, well distinct at maturity, smooth or ornamented, egg-shaped, pale and often white, with hypogynium (hardened gynophore) at base sometimes cupuliform (sometimes reduced) | Sclerieae (Scleria) |
7. | Robust perennial; leaf with very scabrous and sharp margins; inflorescence terminal, paniculate or briefly corymbiform | Cladieae (Cladium) |
– | Plants of variable size, if robust the leaf sometimes scabrous but not sharp; inflorescence terminal or pseudolateral, anthelate or capitate | 8 |
8. | Style base distinct, often thickened and/or distinctly fimbriate | 9 |
– | Style base neither distinct nor thickened and non-fimbriate | 11 |
9. | All leaves reduced to bladeless sheaths; inflorescence always reduced to a terminal spikelet without involucral bracts; flower without adaxial and bifid hypogynous scale | Eleocharideae (Eleocharis) |
– | At least a few leaves or involucral bracts with well-developed leaf blades; inflorescence variable, usually composed of several spikelets, if inflorescence reduced to a terminal spikelet then at least 1 involucral bract larger than the glumes or flower with 1 hypogynous scale, adaxial, flattened, more or less obovate, bifid at apex | 10 |
10. | Inflorescence paniculate or corymbiform, rarely capitate and then the spikelet with several empty basal glumes and only the upper glumes fertile; perianth present or absent, formed of 6 hairs | Rhynchosporeae (Rhynchospora) |
– | Inflorescence anthelate or capitate; spikelets usually with all glumes fertile; if some basal glumes empty, spikelets with many flowers; perianth absent or flower subtended by a bifid scale | 15 (Abildgaardieae) |
11. | Leaf ligulate; glumes distichous and glabrous | 12 (Schoeneae) |
– | Leaf eligulate or reduced to sheath; if ligulate, glumes spirally arranged and hairy | 13 |
12. | Culms terete, elliptical or partially flattened | Oreobolinae (Costularia) |
– | Culms highly variable, commonly biconvex, but ranging from flat to terete or quadrangular or occasionally biconvex | Lepidospermatinae (Machaerina) |
13. | Glumes and spikelets persistent on the plant | 14 |
– | Glumes deciduous or the spikelet deciduous as a whole or in pieces | 19 |
14. | Inflorescence a reduced panicle; glumes distichously arranged; perianth present | Carpheae (Carpha) |
– | Inflorescence anthelate, often reduced to a single spikelet; glumes spirally arranged; perianth absent | 15 (Abildgaardieae) |
15. | Leaf sheath apex with long white hairs; style base mostly enlarged and persistent; style glabrous | Bulbostylis |
– | Leaf sheath apex without long white hairs, an adaxial ligule of minute hairs present or absent; style base enlarged and mostly deciduous; style fimbriate, hispidulous, rarely glabrous | 16 |
16. | Fertile flowers per spikelet more than 2 | 17 |
– | Fertile flowers spikelet 1(–2) | 18 |
17. | Lowermost involucral bract glume-like, inflorescence of 1–3 spikelets; nutlets 2–3.4 mm long (always trigonous, stipitate) | Abildgaardia |
– | Lowermost involucral bract not glume-like, inflorescence of 1–many spikelets; nutlets 0.5–1.8 mm (rarely up to 2.2 mm) long (many species trigonous some biconvex, stipitate or not) | Fimbristylis |
18. | Style base persistent, spreading over apex of nutlet; leaves, stem and glumes with many long erect hairs | Trichoschoenus |
– | Style base deciduous with rest of style, slightly enlarged | Actinoschoenus |
19. | Perianth absent; glumes usually distichous or reduced to 1 per spikelet (and then spikelet bract larger or smaller than glume), rarely spirally arranged; ligule always absent; if glumes spirally arranged and primary involucral bract erect, then stem capillary, not compressible and less than 1 mm wide | 20 (Cypereae) |
– | Perianth present or absent, glumes always spirally arranged; ligule present or, if absent, the inflorescence always with erect primary involucral bract and cylindrical, compressible stem, more than 1 mm wide | 23 |
20. | Glumes usually spirally arranged; anatomy C3. If distichously arranged, then 2 or more parallel veins prominently visible on glume and/or nutlet bearing a distinct gynophore | 21 (Ficiniinae) |
– | Glumes usually distichously arranged; anatomy C3 or C4. If spirally arranged, then either with elongated anther filaments, usually pale or white head-like inflorescences or parallel veins not strongly visible on glume and no gynophore | Cyperinae (Cyperus) |
21. | Leaf blades and inflorescence bracts scabrid, nutlets with gynophore | Ficinia |
– | Leaf blades and inflorescence bracts smooth, nutlets without gynophore | 22 |
22. | Glumes cartilaginous | Ficinia |
– | Glumes chartaceous to hyaline | Isolepis |
23. | Culm with more than 3 nodes | 24 |
– | Culm nodeless or with less than 1–3 nodes above the base | 25 |
24. | Glumes puberulent, apex entire to emarginate or deeply 2-fid, awned or mucronate | Bolbochoeneae (Bolboschoenus) |
– | Glumes often pubescent, apex entire and mucronate to awned | Fuireneae (Fuirena) |
25. | Inflorescence bracts often large, erect, stem-like, rarely leaf-like, and patent to reflexed | Schoenoplecteae (Schoenoplectus) |
– | Inflorescence bracts culm-like, erect, or patent while fruiting, rarely short, rigid and sheathing | Pseudoschoeneae (Schoenoplectiella) |
Lepironia mucronata Rich. [= Lepironia articulata (Retz.) Domin]
Medium-sized to tall perennials; rhizomes woody, creeping with thick roots. Culms scapose, erect, terete, with transverse septa. Basal leaves, reduced to a sheath, open in front, the margins overlapping, eligulate. Involucral bracts 1, large, subulate, cylindric, erect, culm-like. Inflorescence a single spike, pseudolateral, with many spirally arranged imbricate glume-like bracts; basal glume-like bracts empty, most subtending spicoids. Spicoids with 2 outer strongly keeled glumes and many non-keeled glumes, most subtending 1 stamen and a solitary apparently terminal female flower; rachilla thick and spongy. Florets unisexual; perianth absent. Stamens 1; anthers linear, apiculate; filaments highly accrescent. Style deeply 2-fid, long, slender; base not distinct, slightly thickened, persistent. Nutlets obovoid, dorsiventrally compressed, plano-convex, ± winged along the margins, beaked.
Lepironia occurs in Madagascar, and in tropical and subtropical Asia to the Western Pacific (
Hypolytrum latifolium Rich. ex Pers. [= Hypolytrum nemorum (Vahl) Spreng.]
Perennials, moderately robust to robust; rhizomatous or stoloniferous; rhizomes often woody, roots coarse. Culms trigonous or cylindrical, either laterally scapose and bearing reduced leaves at the base (cataphylls), or central with 1-several nodes and bearing well-developed leaves. Leaves eligulate, 3-ranked, sometimes pseudopetiolate; leaf sheath of basal leaves open adaxially; leaf sheath of cauline leaves tubular; blade linear to oblong-lanceolate or reduced (often in basal leaves), with 3, well developed principal veins, margin entire or many small teeth, gradually or abruptly narrowed at apex. Involucral bracts small, scale-like, or large and leaf-like, not sheathing, patent to reflexed. Inflorescence terminal, paniculate, corymbose or capitate, the ultimate branches subtending small clusters of spikes. Spicoids composed of 2(–3) floral bracts, each subtending 1 stamen (male flowers), often connate to varying degrees, the lower two followed by a bare pistil (female flower). Florets unisexual; perianth absent. Stamens 1; anthers oblong to linear, latrorsely dehiscent, without extended connective tip; filaments filiform, exceeding spicoid bract. Style 2-fid, exserted; base distinct or not, thickened or not, more or less persistent. Nutlets obovoid to ellipsoid, dorsiventrally biconvex, smooth, costate, wrinkled, tuberculate or spongious.
Hypolytrum is widely distributed in the tropics (
Hypolytrum nudicaule. A. Habit. B. Inflorescence. C. Plant base. All photos taken on Nosy Boraha Island by Justine Faure (https://www.inaturalist.org/observations/132556611), reproduced with permission from the photographer.
Coleochloa abyssinica (Hochst. ex A.Rich.) Gilly
Tufted or cushion-forming perennials; rhizomes extensively branched, infrequently forming a small caudex. Culms scapose or nearly so, compressed below, sometimes subcylindrical; erect or slightly wanting, basal part of culms covered by remains of old leaf sheaths. Leaves basal only, distichous; leaf sheath open on ventral side; ligule a line of fine hairs; contraligule not developed; blade flat or inrolled, deciduous. Involucral bracts more or less leaf-like; sheaths partly closed. Inflorescence terminal, paniculate, with few to many spikelet-like spikes; spikes bisexual, with numerous lateral spikelets subtended by small, densely spirally arranged glume-like bracts. Spikelet unisexual (male) or bisexual; lateral spikelets with an irregularly shaped prophyll. Glumes distichous, persistent, the larger 1–2 each subtending a flower, enclosed by the wings of the next glume. Basal spikelets with 1–2 male florets, apical spikelets mostly with 1 male and 1 female floret, more rarely completely female. Perianth surrounding the style base by 3 small, long fimbriate hypogynous scales, deciduous with the fruit. Stamens 2–3. Style 3-fid; base not distinct, tapering, persistent. Nutlets fusiform, trigonous, long beaked, surface smooth.
Coleochloa occurs in tropical and southern Africa and Madagascar (
Coleochloa setifera. A. Habit, entire plant. B. Habit. C. Part of culm and leaf sheath. D. Leaf sheath opened to show ligule. E. Nutlet split open, seed enclosed. F. Seed. G. Nutlet, some bristles removed. H. Bristles detached from base of nutlet. I. Three fascicles of spikelets. J. Mature anther. K. Spikelet and bract, abaxial view. L. Spikelet, bract removed, adaxial view. M. Spikelet, glumes displaced, adaxial view. A–D, I–L from Browning 560; E–H from Pawek 13626a. Scale bars: A = 250 mm; B = 40 mm; C–M = 1 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Cladium jamaicense Crantz [= Cladium mariscus subsp. jamaicense Kük.]
Robust perennials, up to several meters tall; rhizomatous, sometimes with swollen stolons. Culms with few-noded, internodes hollow. Leaves basal and cauline, V-shaped to flat, midribs and margins scabrid, eligulate. Involucral bracts leaf-like, sheathing. Inflorescence terminal or some lateral, paniculate; partial inflorescences anthelate. Spikelets numerous, short stalked or sessile. Glumes few to many, spirally arranged, persistent, increasing in length, with 2–3 upper glumes fertile. Lower floret mostly functionally male, upper florets bisexual. Perianth absent. Stamens 2–3. Style 2–3-fid, with a thickened persistent base. Nutlets ovoid, with a thick corky beak, surface smooth to wrinkled. Embryo small and poorly developed, broadly obovate in outline, with a basal, poorly developed root cap and without a leaf primordium.
Cladium is a cosmopolitan genus (
Cladium mariscus subsp. jamaicense. A. Habit. B. Inflorescence. C. Plant base, longitudinal section. D. Detail of leaf. E. Nutlet and upper unisexual floret with 2 filaments and rudimentary gynoecium. F. Spikelet. All from Ward 9082. Scale bars: A = 25 cm; B, C = 5 cm; D = 1 cm; E, F = 2 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Diplacrum caricinum R.Br.
Small to medium-sized annuals or tufted, rarely stoloniferous perennials. Culms scapose or leafy, often short. Leaves eligulate; blade linear or lanceolate, alternate, simple, often reddish purple abaxially. Primary bracts leaf-like, sheathing. Inflorescence paniculate or capitate; partial inflorescences anthelate or capitately contracted. Spikelets many, lateral spikelets usually male, terminal spikelets usually female. Glumes male spikelets with few distichous, persistent glumes, each subtending a male floret; female spikelet with 2 distichous, persistent glumes, sometimes deciduous with the fruit, surrounding a pseudo-terminal female floret. Florets unisexual. Bristles absent. Stamens 1; anthers sightly oblongs. Pistil seated on a basal trilobed disc, lobes opposite the 3 main ribs; style 3-fid ; base not distinct, not thickened, deciduous. Nutlets subglobose to ovoid, inconspicuously 3-ribbed, beak short, surface smooth, ribbed, or reticulate.
Diplacrum is widely distributed in the tropics and tubtropics (
Diplacrum africanum. A, C. Habit. B Leaf. D. Inflorescence. E. Spikelets, male and female with bract and prophyll. F. Prophyll. G–H. Female spikelet, with and without nutlet. I–J. Nutlet, lateral and basal view. B–D, G–J from Robinson 2851; A, E, F from Bidgood et al. 3967. Scale bars: A, B = 25 mm; C = 10 mm; D–H = 1 mm; I, J = 0.5 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Scleria flagellum-nigrorum P.J.Bergius
Habit variable, from tiny annuals with fibrous roots to perennial climbers more than 10 meters tall; stoloniferous rhizome or tubers; aerial adventitious roots at stem nodes (adaptation to flooded habitats). Culms trigonous or triquetrous, noded bearing leaves often without ligules, sometimes with a contraligule. Leaves alternate, tristichously arranged, often persistent at the base, and finely serrate at least along the distal third of the margins, rarely smooth: sometimes abruptly narrowed down or pseudopraemorse; sheaths sometimes winged, usually topped by a contraligule, opposite to the blade. Inflorescence bracts leaf-like and sheathing, setaceous, or glume-like; spikelet bract usually setaceous, rarely glume-like. Inflorescence variable, usually paniculate, but often with contracted partial inflorescences. Spikelets bearing flowers of one or both sexes, the bisexual ones with one basal female and one to few male flowers above; female spikelet similar but upper part reduced to 1–2 empty scales or wanting; male spikelet lacking basal female flower and with more male flowers. Glumes in androgynous or bisexual spikelets the lower part is female with distichously arranged glumes (a few may be empty), upper part male with few to many spirally arranged glumes. Florets always unisexual, enclosed by at least three glumes. Bristles absent. Stamens 1–3, anthers often linear, more or less apiculate. Style 3-fid; ovary surrounded at the base by a variously shaped (sometimes reduced) lobed hypogynium, which is shed with the fruit. Nutlets globose to ovoid, variously sculptured and ornamented, usually white, sometimes beaked, subtended by a cupule, frequently surrounded by a hypogynium.
Scleria is widely distributed in the tropics and subtropics up to North America (
Scleria bulbifera. A. Habit showing rhizome. B. Junction of lamina and sheath. C. Androgynous spikelet with lower prophyll and one glume removed; remainder displaced to show lower female floret and upper male florets. D. Glomerule of three spikelets. E. Nutlet. A–D from Hilliard & Burtt 13394; E from Hilliard & Burtt 18036. Scale bars: A = 40 mm; B–E = 2 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Carpha alpina R.Br.
Perennials, small to medium-sized (rarely tall), tufted, mat-forming; rhizomatous or rarely stoloniferous. Culms erected, trigonous to more or less cylindrical, scapose or with a few nodes. Leaves basal and cauline, eligulate. Lower primary bracts more or less leaf-like, sheathing. Inflorescence open to condensed paniculate with (sub)capitate partial; inflorescences with few to many spikelets. Spikelets with 3–6 distichous, persistent glumes of increasing length. Lower glumes empty, larger 1–2(–3) glume(s) each subtending a floret, enclosed by the wings of the next glume. Floret bisexual, sometimes upper or lower floret male. Bristles 6, ciliate to partly fimbriate or plumose, shorter than to much longer than the fruit, deciduous with the fruit. Stamens 2–3, anthers conspicuously greenish yellow. Style 3-fid, base not distinct, slightly thickened, persistent, often scabrid. Nutlets narrowly (ob)ovoid to oblong, trigonous, often with a long beak, surface smooth or finely reticulate.
Carpha occurs from Uganda to South Africa, Western Indian Ocean, South Japan, New Guinea to New Zealand and South America. It grows along streams, on marshy ground, and on rocks, from 1800 to 2500 m in elevation. In Madagascar, a single species of Carpha, i.e. Carpha perrieri Cherm., is known from Fianarantsoa province Matsiatra Ambony region and Toliara provinces Anosy region.
Costularia natalensis C.B.Clarke
Perennials, small to tall, tufted or more rarely shortly rhizomatous, caudex sometimes present. Culms scapose or with few nodes. Leaves usually basal and cauline; basal leaves with poorly defined sheaths; cauline leaves enveloping up to 1/2 internode length; spirodistichous, eligulate; blade sometimes deciduous, margins scabrid. Involucral bracts more or less leaf-like, sheathing. Inflorescence terminal, contracted paniculate. Spikelets few to numerous. Glumes several, distichous, deciduous, of increasing length, the upper (1–)2 glumes each subtending a floret, enclosed by the wings of the next glume. Lower floret functionally male rarely bisexual or absent, upper floret bisexual or functionally female rarely functionally male. Bristles 6, fimbriate to ciliate, mostly longer than the nutlet and deciduous with it. Stamens 3; anthers long linear, briefly apiculate or not. Style 3-fid; base often distinct (at anthesis), thickened, persistent, often scabrid. Nutlets ovoid or oblong, rounded trigonous, often 3-ribbed, more or less stipitate, beaked, surface smooth or rugulose.
Costularia occurs in southeastern Africa, Madagascar, the Mascarenes, and Seychelles. It grows on rocky ground in forest, grassland, or ericoid vegetation, sometimes along streams or in swamps, usually at higher elevations. In Madagascar, Costularia is known from north to south along the mountain ridge and high elevation areas, including the Central Highlands. The genus was recently monographed (
Costularia pantopoda var. pantopoda. A. Habit. B. Inflorescence matching the habit. C. Spikelet. D. Lowest glumes of spikelets. E. Rest of glumes lower to upper, abaxial view. F. Bisexual flower. G. Male flower. H. Nutlet. Scale bars: A–B = 3 cm, C–H = 3 mm. All from Larridon et al. 2010-0144. Drawn by Juliet Beentje, reproduced with permission from the artist, originally published in
Schoenus restioides Sw. [= Machaerina restioides Vahl]
Medium-sized to tall creeping rhizomatous or stoloniferous perennials. Culms scapose or with few nodes, flattened, angular, compressed or (sub)terete, sometimes septate. Leaves distichous, eligulate, blade ensiform to (sub)terete, sometimes septate, rarely reduced to a sheath. Primary bracts often leaf-like, sheathing. Inflorescence sometimes more or less pseudolateral, paniculate, main axis often sinuous, partial inflorescences sometimes capitately contracted. Spikelets many. Glumes 2–10 distichous, long-persistent, of increasing length, with (1–)2–4(–5) florets subtended by the larger glumes and enclosed by the wings of the next glume. Lower 1–2 floret(s) usually bisexual, upper floret(s) usually functionally male. Bristles absent or poorly developed, or up to 6, delicate, shorter than, to as long as the fruit, deciduous with the fruit. Stamens 3; filaments sometimes conspicuously elongated after anthesis. Style 3-fid, base not distinct, thickened, persistent, sometimes scabrid. Nutlets ovoid to oblong, terete to triquetrous, more or less 3-ribbed to winged, stipitate or sessile, beaked, smooth to rugose.
Machaerina occurs in Tanzania, Western Indian Ocean to Pacific, Tropical America (
Machaerina flexuosa. A. Habit (× 2/3). B. Spikes from top of inflorescence (× 3). C–D. Glume, abaxial and side view (× 10). E. Flower (× 10). F. Young flower (× 8). G. Anther (× 16). H. Nutlet (× 16). A from Andriamahay & Rakotoarison 1969; B–G from Nusbaumer et al. 1103; H from Ranirison 642. Drawn by Juliet Beentje, reproduced with permission from the artist, originally published in
Rhynchospora alba (L.) Vahl
Usually small to medium-sized perennials, rarely annuals; rhizomatous or with a poorly developed root system. Culms scapose or with 1-many nodes, rounded to trigonous. Leaves radical or radical and cauline; sheaths closed, ligule minute or absent. Involucral bracts leaf-like or reduced. Inflorescence very variable, paniculate, corymbose, anthelate or capitate, rarely pseudolateral. Glumes spirally arranged (rarely distichous), of increasing length to subequal, larger (1–)2-few glume(s) subtending a floret, enclosed by the wings of the next glume. Florets bisexual, the lowest few sterile and/or upper staminate (variable with species). Perianth bristles usually 3–6, or absent. Stamens (1–2–)3, inconspicuous. Style 2-fid, with base persistent (tubercle). Nutlets usually lenticular to globose. Embryo top-shaped in frontal view, root cap developed in a (sub)basal position, and first leaf primordium developed in a lateral position (Carex-type embryo).
Rhynchospora is a cosmopolitan genus (
Rhynchospora holoschoenoides. A–B. Habit. C. Leaf sheath. D. Leaf apex. E. Inflorescence. F. Head of spikelets. G–H. Spikelet, complete and opened to show florets. I. Glume, lateral view. J. Nutlet. A–D from Robinson 6123; E, G, I from Robinson 1048; F from Robinson 1456; J from Renvoize 5598. Scale bars: A = 250 mm; B, E = 40 mm; C, F = 5 mm; G–I = 2 mm; D, J = 1 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Schoenoxiphium
Nees (
Carex hirta L.
Generally perennials, rarely annuals, caespitose or rhizomatous, sometimes forming a compact tussock. Culms central or lateral, mostly scapose, rarely with nodes, simple, smooth or scabrid. Leaves generally present, glabrous or rarely hairy, sheath often surrounding culm, ligule or sometimes contraligule present at the junction of the sheath and the blade. Primary bracts leaf-like or not, sheathing or not. Partial inflorescences spike-like, unisexual or bisexual, with few to many spirally arranged bracts (or glumes), either subtending a female spikelet or a male floret. Inflorescence terminal, rarely pseudolateral, paniculate, often partly or completely contracted, rarely corymbose or anthelate with few to numerous spikes, less frequently reduced to a single spike, cladoprophylls sometimes swollen at the base, utriculiform and subtending a female floret; inflorescence mostly bisexual, rarely unisexual or florets dioecious. Spikes male, female, or bisexual, and then mostly male florets apically, rarely basally, or intermingled; female spikelet reduced to the rachilla and a utriculiform, flower-bearing prophyll, completely enclosing the rachilla. Glumes 0 or 1. Floret unisexual; staminate flowers without scales; pistillate flowers with 1 scale with fused margins (perigynium) enclosing flower, open only at apex. Bristles absent. Stamens 1–3, filaments distinct, anthers linear. Style 2–3-fid, exserted, base not distinct, rarely thickened, persistent. Nutlets often obovoid, trigonous, or dorsiventrally compressed, sometimes remarkably malformed.
Carex is a cosmopolitan genus (
Websteria
S.H.Wright (
Scirpus palustris L. [= Eleocharis palustris (L.) Roem. & Schult.]
Very small to medium-sized tufted annuals or large rhizomatous, stoloniferous perennials; rhizome often strong, horizontal, often producing stolons. Culms scapose, 3–4 angular, ridged or terete, occasionally septate, ancipitous or bulbously thickened at base. Leaves reduced to inconspicuous basal sheaths, rarely with a short blade, eligulate. Primary bracts absent, rarely proximal scale of spikelet resembling short bract. Inflorescence solitary terminal spikelet, usually quite short and ebracteate. Spikelets with 2–many spirally arranged, rarely subdistichous, deciduous imbricate glumes, each subtending a floret. Lowermost glume often empty or with a vegetative bud, rarely flower-bearing. Floret bisexual. Bristles barbellate, (0–)3–6(–10), or reduced to a narrow rim underlying nutlet, shorter or longer than nutlet and shed with it. Stamens 1–3; anthers linear. Style 2–3-fid, upper portion deciduous, base enlarged persistent as a conical or flattened appendage on nutlet. Nutlets obovoid, lenticular or trigonous, beaked smooth or variously ornamented with pits in longitudinal rows.
Eleocharis is a cosmopolitan genus (
Eleocharis dulcis. A. Habit (× 1/3). B. Inflorescence (× 1.5). C. Floret (× 10). D. Nutlet (× 12). A from Kirika et al. NMK 778; B from Vesey-Filtzgerald 401; C from Milne-Redhead & Taylor 9164; D from Faden et al. 96/468. Drawn by Juliet Beentje, reproduced with permission from the artist, originally published in
Scirpus capillaris L. [= Bulbostylis capillaris (L.) Kunth ex C.B.Clarke]
Small to medium-sized annuals or tufted perennials, rarely with an elongated rhizome, rarely forming a caudex; rhizome woody, variable, usually compact with swollen confluent shoot bases, occasionally elongate in uniseriate or multiseriate rows, less often of uniform thickness throughout. Culms scapose, terete, ridged and furrowed, glabrous to pilose or densely velutinous apically. Leaves eligulate (rarely ligulate), but with two lateral tufts of long white hairs at the sheath mouth, rarely reduced to a sheath. Primary bracts short, not sheathing, rarely the lowermost bract leaf-like and erect. Inflorescence terminal, rarely pseudolateral, anthelate or capitate. Spikelets few to many, or reduced to a single spikelet, often with many densely spirally arranged (rarely distichous), deciduous glumes, each subtending a floret. Florets bisexual. Perianth bristles absent. Stamens 1–3, anthers generally oblongs or linear, often acute, rarely setiferous. Style (2–)3-fid, base distinct, thickened, persistent, rarely only slightly thickened or deciduous. Nutlets obovoid to obpyriform, rounded trigonous, rarely dorsiventrally lenticular, surface with various ornamentations, rarely smooth.
Bulbostylis is widely distributed in the tropics and subtropics to Central Asia (
Bulbostylis hispidula. A. Habit. B. Glume. C. Nutlet. D. Spikelet. E. Mouth of leaf sheath. F. Young nutlet with style. All from Ward 610. Scale bars: A = 40 mm; B, C = 5 mm; D–F = 0.5 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Trichoschoenus bosseri J.Raynal
Small to medium-sized tufted perennials; thick roots. Culms scapose, densely hairy, flattened, tufted. Leaves reduced to a sheath, leaf sheath of increasing length, blade sometime present as a short mucro, eligulate. Primary bracts mostly as long as or shorter than spikelet, not sheathing. Inflorescence capitate. Spikelets many. Glumes 3, distichous, deciduous, of increasing length, the second subtending a floret, enclosed by the wings of the next. Florets bisexual. Perianth bristles absent. Stamens 3. Style deeply 3-fid, base more or less distinct, thickened, persistent. Nutlets obovoid, trigonous, with a thick short beak, surface minutely reticulate.
Trichoschoenus is the only endemic genus of Cyperaceae in Madagascar. Its only species Trichoschoenus bosseri (see
Actinoschoenus filiformis (Thwaites) Benth. [= Actinoschoenus aphyllus (Vahl) Larridon]
Small to medium-sized, tufted, shortly rhizomatous or stoloniferous perennials. Culms scapose, smooth, tufted, more or less 4-angular, bases covered by closed leaf sheaths. Leaves ligulate if present, blade very short, almost reduced to a sheath. Primary bracts small, not very conspicuous not sheathing. Inflorescence terminal, capitate. Spikelets 2 or many. Glumes 4–7 distichous, deciduous glumes, of increasing length; rachilla internodes short, somewhat elongated between the florets. Floret(s) 1(–2), subtended by the penultimate larger glume, enclosed by the wings of the next glume, bisexual. Perianth bristles absent. Stamens 3. Style deeply 3-fid, style base distinct, thickened, deciduous. Nutlets obovoid, trigonous, more or less 3-ribbed, surface smooth to slightly tuberculate.
Actinoschoenus occurs from West-Central Tropical Africa to Zambia, Western Indian Ocean, Sri Lanka to southern China and Australia (
Actinoschoenus aphyllus. All photos taken in Andrainjato, Taolagnaro District, Anosy Region by Andriambolantsoa Rasolohery (https://www.inaturalist.org/observations/145409565), reproduced with permission from the photographer.
Cyperus monostachyos L. [= Abildgaardia ovata (Burm.f.) Kral]
Small to medium-sized annuals or tufted perennials; short woody rhizome. Culms scapose, rounded, generally glabrescent, rarely scabrid. Leaves eligulate, sometimes reduced to a sheath, basal polystichous, sheaths distally open, loose, ribbed; blades mostly filiform, compressed or lunate to semicircular in cross section, margins strongly involute. Primary bracts short, not sheathing, inconspicuous. Inflorescence terminal, depauperate anthelate or capitate. Spikelets few or reduced to one single spikelet. Glumes many densely (spiro) distichous, deciduous, each subtending a floret. Floret bisexual, protandrous. Perianth bristles absent. Stamens 2–3. Style deeply 3-fid, base distinct, thickened, deciduous. Nutlets mostly obovoid, stipitate, rounded trigonous, rarely winged, surface often tuberculate.
Abildgaardia is widely distributed in the tropics and subtropics (
Abildgaardia triflora. A. Habit. B. Two leaf bases. C. Leaf apex. D. Spikelet. E. Rachilla. F, G. Glume, complete and upper abscised part. H. Young floret. I. Style and branches. J. Nutlet with filaments. All from Robinson 2038. Scale bars: A = 40 mm; B, C, H–J = 2 mm; D–G = 5 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Scirpus dichotomus L. [= Fimbristylis dichotoma (L.) Vahl]
Annuals tufted or perennials with rhizomes. Culms scapose or subscapose, terete or 3–4–5-angular or ancipitous, variously pubescent to glabrous. Leaves spirally or distichously arranged, or reduced to a sheath, ligulate or eligulate; blade linear, filiform, or rarely ensiform, usually dorsiventrally compressed and canaliculate, often adaxially cellular-reticulate. Primary bracts short, not sheathing. Inflorescence variable usually capitate or anthelate. Spikelets few to numerous, or reduced to a single spikelet, mostly cylindric. Glumes usually spirally arranged, rarely distichous, deciduous, each subtending a bisexual floret. Perianth bristles absent. Stamens 1–3. Style 2–3-fid, often flattened with fimbriate margins when 2-fid; base distinct, thickened, deciduous. Nutlets obovoid, trigonous or biconvex and often variously ornamented, trigonous when style 3-fid, lenticular when style 2-fid, smooth, tuberculate or longitudinally ribbed, not transversely wrinkled.
Fimbristylis is a cosmopolitan genus (
Fimbristylis dichotoma. A, B. Habit. C. Leaf sheath apex. D. Leaf apex. E. Spikelet. F, G. Glume, abaxial and lateral view. H. Floret. I. Nutlet. All from Brummitt 9546. Scale bars: A = 250 mm; B = 40 mm; C, E = 5 mm; D = 2 mm; F–I = 1 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Bolboschoenus maritimus (L.) Palla
Perennials with long rhizomes often forming hard ovoid tubers at tips. Culms many-noded, sharply trigonous, thickened at base. Leaves basal and cauline, eligulate; blade often reduced in lower leaves. Involucral bracts leaf-like, patent, lowermost often suberect. Inflorescence terminal (in reduced inflorescences, bract may be erect, but clearly leaf-like), a (compound) corymb-like anthela or capitate with 1-many spikelets. Spikelets with many spirally arranged, deciduous glumes, each subtending a flower. Glumes puberulent, the apex entire to emarginate or deeply 2-fid, awned or mucronate. Floret bisexual. Perianth present, formed by 3–6 parts, shorter to longer than the nutlet, bristle-like, deciduous with fruit. Stamens 3. Style 2–3-fid; base persistent, barely thickened. Nutlets obovate, dorsiventrally lenticular or trigonous. Pericarp with the three highly differentiated layers, exocarp cells often enlarged and hollow; surface smooth, epidermal cells roughly isodiametric. Embryo fungiform with three primordial leaves.
Bolboschoenus is a cosmopolitan genus (
Bolboschoenus glaucus. A. Habit. B. Inflorescence and leaves. C. Leaf sheath apex. D. Spikelet. E. Anther. F. Ovary, style and branches. G, H. Glume, adaxial and abaxial surface. I. Glume apex. J. Nutlet, abaxial view. K, L. nutlet sections. All from van Rensburg 2404. Scale bars: A = 250 mm; B = 10 mm; C = 40 mm; D–K = 2 mm. Abbreviations: ex = exocarp, m = mesocarp, en = endocarp. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Fuirena umbellata Rottb.
Annuals or rhizomatous perennials. Culms many-noded, rarely scapose, 3–5-sided, sometimes thickened at base. Leaves usually well developed, basal and cauline, ligule tubular, membranous, with blade often reduced in lower leaves (rarely all leaf blades reduced). Involucral bracts leaf-like, usually sheathing, lowermost bract sometimes erect. Inflorescence terminal (in reduced inflorescences, bract may be erect, but clearly leaf-like), paniculate to capitate with few to many spikelets. Glumes many, spirally arranged or rarely pentastichously arranged, deciduous, each subtending a flower, often pubescent, the apex entire and mucronate to awned. Floret bisexual. Perianth present, as long or shorter than nutlet, formed by 3 parts, or when 6 in 2 whorls, the inner parts scale-like, the outer parts bristle-like, rarely all parts reduced or absent or only 1 scale developed, deciduous with the fruit. Stamens 1–3. Style 3-fid, base persistent, barely thickened, if at all. Nutlets obovate, triquetrous to trigonous, frequently stipitate, smooth or variously ornamented. Embryo turbinate to weakly fungiform with a horizontally broadened scutellum, first leaf primordium not strongly outgrown, the second leaf primordium either absent or poorly developed.
Fuirena is a cosmopolitan genus (
Fuirena pubescens. A. Nutlet. B. Habit. C. Leaf sheath apex. D. Inflorescence. E. Glume lateral view. F. Floret. A–C from Taylor 36; D–F from Browning 165. Scale bars: A = 40 mm; B, C = 5 mm; D–F = 1 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Schoenoplectus lacustris (L.) Palla
Perennials with long rhizomes sometimes ending in tubers at tips. Culms scapose, trigonous to terete, thickened at base. Leaves usually reduced to a sheath, sometimes developing a ligulate blade, but rarely well developed. Involucral bracts often large, erect, stem-like, rarely leaf-like, and patent to reflexed. Inflorescence pseudolateral, rarely clearly terminal, corymb-like anthela or capitate. Spikelet 1 or few. Glumes many, spirally arranged, deciduous, each subtending a flower; puberulent to glabrous, the margins often ciliate or laciniate distally, apex entire to emarginate or deeply 2-fid, awned or mucronate. Floret bisexual. Perianth present, formed by (5–)6 parts, smooth to retorsely scabrid, bristle-like or sometimes plumose, longer or shorter than nutlet, deciduous with fruit. Stamens 2–3. Style 2–3-fid, base not thickened, persistent. Nutlets smooth, obovate, trigonous, or dorsiventrally lenticular, yellow to dark brown when mature; fruit epidermal cells isodiametric to narrowly oblong. Embryo fungiform, scutellum turbinate to rhomboid in shape, root cap lateral, first well developed and second embryonic leaves basal.
Schoenoplectus is a cosmopolitan genus (
Scirpus articulatus L. [= Schoenoplectiella articulata (L.) Lye]
Annuals or perennials, tufted or with firm, short to creeping rhizomes. Culms nodeless and scapose or 1(–3)-noded above the base, trigonous, terete or rarely 7-sided. Leaves reduced to a mucronate sheath, rarely with well-developed blades, ligulate or eligulate. Involucral bracts culm-like, erect, or patent while fruiting, rarely short, rigid, and sheathing, but then appearing as a continuation of the stem. Inflorescence pseudolateral, rarely appearing terminal, a corymb-like anthela or capitate with 1-many spikelets, rarely compound-paniculate with 1-many spikelets, with a conspicuously sinuous main axis. Glumes many, spirally arranged, deciduous or persistent, each subtending a flower; scale apex entire to apiculate. Floret bisexual, rarely polygamodoecious. Perianth bristles present or absent, formed by 0–10 parts, smooth or retrorsely scabrid, bristle-like, as long as or longer than the nutlet, deciduous with the fruit. Stamens 2–3, rarely vestigial in female flowers. Style 2–3-fid, base undifferentiated, rarely distinct and somewhat thickened, persistent. Nutlets smooth or transversely rugose to distinctly ridged, obovate, trigonous to planoconvex or biconvex, dark nearing black when mature, sometimes brown.
Schoenoplectiella is a cosmopolitan genus (
Schoenoplectiella corymbosa. A. Partial habit (× 2/3). B. Inflorescence (× 2/3). C. Spikelet (× 6). D. Glume (× 10). E. Young floret (× 10). F. Floret (× 10). G. Nutlet (× 20). A from Richards 6616; B, G from Greenway & Kanuri 12546; C–F from Grimshaw 93. Drawn by Juliet Beentje, reproduced with permission from the artist, originally published in
Scirpus setaceus L. [= Isolepis setacea (L.) R.Br.]
Small to more rarely medium-sized tufted annuals or mat-forming perennials; rhizome more rarely creeping, rhizomatous or stoloniferous. Culms scapose or with few to many nodes. Leaves eligulate, of a minute lobe or elongated to form a linear blade often much reduced. Primary bracts leaf-like or short, not sheathing, lowermost bract often erect. Inflorescence often pseudolateral, capitate, rarely anthelate. Spikelets few to many or reduced to a single spikelet. Glumes with few to many usually spirally arranged (rarely distichous), mostly deciduous glumes, each subtending a floret. Floret bisexual. Perianth bristles absent. Stamens 1–3; filament ribbon-like, anther crested with minute spikes. Style deeply 2–3-fid; base not distinct, not or slightly thickened, persistent. Nutlets mostly obovoid, thick lenticular to (rounded) trigonous, often 3-ribbed, beaked, surface with various ornamentations.
Isolepis is a cosmopolitan genus (
Isolepis fluitans. A. Habit. B. Leaf sheath. C. Spikelet. D. Rachilla, apical glume attached. E. Glume, lateral view. F. Floret. G, H. Nutlet, abaxial and apical view. All from Iversen & Martinsson 89195. Scale bars: A = 40 mm; B–H = 1 mm. Drawn by Jane Browning, reproduced with permission from the artist, originally published in
Schoenus filiformis Lam. [= Ficinia filiformis (Lam.) Schrad.]
Small to medium-sized tufted perennials; frequently with a short or elongated creeping rhizome, stoloniferous, or decumbent. Culms scapose, more rarely with few to many nodes or branched, rarely thickened at the base, leafy only at the base or in all their length. Leaves often conspicuously ligulate (rarely eligulate), blade sometimes with scarious margins, or leaf reduced to a sheath. Primary bracts leaf-like (rarely bright yellow) or short, not sheathing, lowermost bract sometimes erect. Inflorescence rarely pseudolateral, mostly capitate, more rarely compacted, paniculate, spicate, or spikelets scattered along a profusely branched stem. Spikelets 1–many. Glumes with few to many spirally arranged or more rarely distichous, usually long-persistent glumes, each with a floret, or a few lower glumes empty. Floret bisexual. Perianth bristles absent. Stamens 3; anthers linear, often apiculate or setiferous. Style deeply 3-fid, sometimes 2-fid, rarely almost undivided; base not distinct, not thickened, deciduous. Nutlets mostly obovoid, rounded trigonous, rarely biconvex, base mostly surrounded by a tightly enveloping cupular to 3-lobed disc (gynophore), surface usually smooth.
Ficinia is a cosmopolitan genus (
Kyllinga
Rottb. (
Remirea
Aubl. (
Mariscus
Vahl (
Pycreus
P.Beauv. (
Lipocarpha
R.Br. (
Torulinium
Desv. ex Ham. (
Oxycaryum
Nees (
Ascolepis Nees (Nees von Esenbeck 1855: 105)
Queenslandiella
Domin (
Courtoisina
Soják (
Cyperus esculentus L.
Small to large annuals, or tufted, rhizomatous, stoloniferous or bulbiferous perennials. Culms mostly scapose, triangular to subterete, rarely winged, compressed or septate. Leaves eligulate (more rarely ligulate), sometimes reduced to a sheath; sheaths sometimes semisucculent; blade linear or rarely oblong or pseudopetiolate. Primary bracts often large and leaf-like, not sheathing, lowermost bract rarely erect. Inflorescence of few to many spikelets, in clusters called spikes, terminal, rarely pseudolateral, anthelate to capitate, rarely spicate or reduced to a single spikelet; partial inflorescences spike-like with spicately or digitately-arranged spikelets, or densely capitate. Spikelets with few to many, 2-ranked (very rarely spirally arranged) glumes, spikelet bract and prophyll more or less glume-like, each glume subtending a floret. Glumes deciduous or persistent; rachilla internodes elongated. Floret bisexual, very rarely unisexual or even dioecious. Bristles absent, lacking a gynophore. Stamens 1–3. Style (2–)3-fid, more rarely (sub)entire; base mostly not distinct, not thickened, persistent or deciduous. Nutlets most often 3-sided, with the flat side pressed against the spikelet rachilla, often obovoid or ellipsoid, rarely dorsiventrally compressed, rarely stipitate, beaked or not, surface smooth or with various ornamentations, rarely thickened, corky.
Cyperus is a cosmopolitan genus who can be found everywhere. Cyperus occurs throughout Madagascar; ca 145 species are known including e.g. Cyperus pectinatus Vahl (Figs
Cyperus pectinatus. A. Habit (× 2/3). B. Inflorescence (× 1.5). C. Spikelet (× 3). D. Glume (× 10). E. Flower (× 10). F. Nutlet (×10). A from Peter 8794; B–D, F from Greenway & Kanuri 12298; E from Richards 24601. Drawn by Juliet Beentje, reproduced with permission from the artist, originally published in
We are grateful to an anonymous member of the International Sedge Society for sponsoring a 4-month research placement for the first author in preparation of starting a PhD at University of Antananarivo in collaboration with Royal Botanic Gardens, Kew. This study was supported by the Today’s Flora for Tomorrow project funded by a generous donor through Kew Foundation. We would like to thank Benoît Loeuille and Jérémie Morel for improving the language in the French version of this paper.