| # The grass genera of the world |

L. Watson, T.D. Macfarlane, and M.J. Dallwitz

### Cenchrus L.

From the Greek kegchros (millet, Panicum miliaceum).

Type species: Type: C. echinatus L.

Including Echinaria Fabric., Nastus Lunell, Raram Adans.

Excluding Odontelytrum, Pennisetum, Snowdenia

Habit, vegetative morphology. Annual, or perennial (‘sand-burrs’); rhizomatous, or stoloniferous, or caespitose, or decumbent. Culms 5–100(–150) cm high; herbaceous; branched above. Culm nodes glabrous. Culm leaf sheaths compressed, or rounded. Culm internodes solid, or hollow. Leaves not basally aggregated; non-auriculate. Leaf blades linear, or linear-lanceolate; narrow; flat, or folded; without cross venation; persistent; rolled in bud. Ligule a fringed membrane to a fringe of hairs.

Reproductive organization. Plants bisexual, all with bisexual spikelets; with hermaphrodite florets. The spikelets of sexually distinct forms on the same plant; hermaphrodite, or hermaphrodite and sterile (if the ‘bristles’ are taken to include vestigial spikelets). Apomictic, or reproducing sexually.

Inflorescence. Inflorescence a false spike, with spikelets on contracted axes (the spikelets in prickly glomerules (burrs) composed of coalescing spines representing modified branchlets). Primary inflorescence branches inserted all around the main axis. Inflorescence with axes ending in spikelets. Rachides angular or compressed. Inflorescence espatheate; not comprising ‘partial inflorescences’ and foliar organs. Spikelet-bearing axes very much reduced (to burrs); disarticulating (but the main axis persistent); falling entire (i.e., the burrs falling). Spikelets with ‘involucres’ of ‘bristles’ (the bristles coalescing, by contrast with Pennisetum). The ‘bristles’ nearly always spiny, markedly coalescent basally (not spiny, merely ciliate, in C. ciliaris); deciduous with the spikelets. Spikelets not secund.

Female-fertile spikelets. Spikelets lanceolate, or ovate; compressed dorsiventrally; falling with the glumes (i.e., in the burrs). Rachilla terminated by a female-fertile floret. Hairy callus absent.

Glumes present; two; very unequal; shorter than the adjacent lemmas, or long relative to the adjacent lemmas; awnless; very dissimilar, or similar (hyaline or membranous). Lower glume 1–5 nerved. Upper glume 1–7 nerved. Spikelets with incomplete florets. The incomplete florets proximal to the female-fertile florets. The proximal incomplete florets paleate, or epaleate (rarely). Palea of the proximal incomplete florets fully developed, or reduced (rarely). The proximal incomplete florets male, or sterile. The proximal lemmas awnless; 1–7 nerved; more or less equalling the female-fertile lemmas; less firm than the female-fertile lemmas; not becoming indurated.

Female-fertile florets 1. Lemmas similar in texture to the glumes to decidedly firmer than the glumes (firmly membranous, dull, papery or leathery); smooth; not becoming indurated; yellow in fruit; entire; awnless; hairless; non-carinate; having the margins lying flat on the palea; with a clear germination flap; 3–7 nerved. Palea present; relatively long; entire; awnless, without apical setae; textured like the lemma; not indurated; 2-nerved. Lodicules absent. Stamens 3. Anthers not penicillate. Ovary apically glabrous. Styles fused, or free to their bases. Stigmas 2.

Fruit, embryo and seedling. Fruit small; compressed dorsiventrally. Hilum short. Embryo large; waisted. Endosperm hard; without lipid; containing only simple starch grains. Embryo without an epiblast; with a scutellar tail; with an elongated mesocotyl internode. Embryonic leaf margins overlapping.

Seedling with a short mesocotyl. First seedling leaf with a well-developed lamina. The lamina broad; curved; 21–30 veined.

Abaxial leaf blade epidermis. Costal/intercostal zonation conspicuous. Papillae absent. Long-cells similar in shape costally and intercostally. Mid-intercostal long-cells rectangular; having markedly sinuous walls, or having straight or only gently undulating walls. Microhairs present; panicoid-type; (45–)54–84(–114) microns long; 6–11.5(–12.6) microns wide at the septum. Microhair total length/width at septum 6–10(–11.6). Microhair apical cells (27–)30–60(–72) microns long. Microhair apical cell/total length ratio 0.5–0.69. Stomata common; 30–45 microns long, or 36–52 microns long (C. echinatus). Subsidiaries markedly triangular. Guard-cells overlapping to flush with the interstomatals. Intercostal short-cells common, or absent or very rare. Costal short-cells conspicuously in long rows. Costal silica bodies ‘panicoid-type’; cross shaped, or butterfly shaped, or dumb-bell shaped.

Transverse section of leaf blade, physiology. C4; biochemical type NADP–ME (C. pauciflorus, C. incertus); XyMS–. PCR sheath outlines uneven. PCR sheath extensions absent. PCR cells with a suberised lamella. PCR cell chloroplasts with reduced grana; centrifugal/peripheral. Mesophyll with radiate chlorenchyma. Midrib conspicuous; having a conventional arc of bundles; with colourless mesophyll adaxially, or without colourless mesophyll adaxially (rarely). Bulliforms present in discrete, regular adaxial groups; in simple fans, or in simple fans and associated with colourless mesophyll cells to form deeply-penetrating fans. Many of the smallest vascular bundles unaccompanied by sclerenchyma. Combined sclerenchyma girders present, or absent; nowhere forming ‘figures’. Sclerenchyma all associated with vascular bundles.

Culm anatomy. Culm internode bundles scattered.

Phytochemistry. Leaves without flavonoid sulphates (1 species).

Cytology. Chromosome base number, x = 9 and 12. 2n = 34, 35, 36, 40, 44, 45, and 68. Chromosomes ‘small’. Mean diploid 2c DNA value 2.6 pg (?-ploidy of the one species studied unknown).

Classification. Watson & Dallwitz (1994): Panicoideae; Panicodae; Paniceae. Soreng et al. (2015): Panicoideae; Panicodae; Paniceae; Cenchrinae. 22 species.

Distribution, phytogeography, ecology. Tropical and warm temperate.

Commonly adventive. Mesophytic to xerophytic; shade species, or species of open habitats; halophytic, or glycophytic. Grassland, bush, sandy and weedy places.

Economic aspects. Significant weed species: C. biflorus, C. brownii, C. ciliaris, C. echinatus, C. incertus, C. longispinus, C. myosuroides, C. pauciflorus, C. tribuloides. Cultivated fodder: C. ciliaris (drought resistant, tolerant of heavy grazing), C. setiger. Important native pasture species: C. biflorus, C. ciliaris, C. pennisetiformis, C. setigerus.

Rusts and smuts. Rusts — Puccinia. Smuts from Ustilaginaceae. Ustilaginaceae — Sorosporium, Sphacelotheca, Tolyposporium, and Ustilago.

References, etc. Leaf anatomical: Metcalfe 1960; studied by us - C. australis R. Br., C. biflorus Roxb., C. brownii Roem. & Schultes, C. ciliaris L., C. echinatus L., C. elymoides F. Muell., C. incertus M.A. Curtis, C. longispinus (Hack.) Fern., C. setiger Vahl.

Special comments. Some species overlapping with Pennisetum.

Illustrations. • Cenchrus prieurii, as Pennisetum: Kunth (1835). • Cenchrus elymoides: Gardner, 1952. • C. elymoides: Gibbs Russell et al., 1990. • Cenchrus ciliaris: Bor, Fl. of Iraq (1968). • Ligule region of Cenchrus ciliaris. • Cenchrus ciliaris, inflorescence detail, spikelet clusters. • A spikelet cluster with bristles (C. ciliaris). Cenchrus ciliaris, a spikelet cluster subtended by ‘bristles’. • Spikelet clusters of Cenchrus setiger. • Rachis and abscising spikelet clusters of Cenchrus setiger. • Old spikelet cluster of Cenchrus setiger. • Spikelet cluster of Cenchrus longispinus. • Cenchrus myosuroides: Sanchez-Ken, Flora del Valle Tehuacán-Cuicatlan (2011). • Germination flap of C. echinatus. • Cenchrus longispinus, abaxial epidermis of leaf blade: this project.

We advise against extracting comparative information from the descriptions. This is much more easily achieved using the DELTA data files or the interactive key, which allows access to the character list, illustrations, full and partial descriptions, diagnostic descriptions, differences and similarities between taxa, lists of taxa exhibiting or lacking specified attributes, distributions of character states within any set of taxa, geographical distribution, and classifications. See also Guidelines for using data taken from Web publications.

Cite this publication as: ‘Watson, L., Macfarlane, T.D., and Dallwitz, M.J. 1992 onwards. The grass genera of the world: descriptions, illustrations, identification, and information retrieval; including synonyms, morphology, anatomy, physiology, phytochemistry, cytology, classification, pathogens, world and local distribution, and references. Version: 15th June 2025. delta-intkey.com’.

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