| # The grass genera of the world |

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

### Axonopus P. Beauv.

From the Greek axon (axis) and pous (foot), alluding to rachides arising from a common point (digitate).

Carpet Grasses.

Type species: Type: A. compressus (Sw.) P.Beauv.

Including Anastrophus Schlecht., Cabrera Lag., Lappogopsis Steud.

Excluding Centrochloa, Ophiochloa

Habit, vegetative morphology. Annual (rarely), or perennial; stoloniferous (sometimes mat-forming), or caespitose. Culms 15–100 cm high (or more?); herbaceous; branched above, or unbranched above. Culm nodes hairy, or glabrous. Culm internodes solid, or hollow. Leaves mostly basal, or not basally aggregated; non-auriculate. Leaf blades linear-lanceolate to ovate-lanceolate; broad, or narrow; flat, or folded; not pseudopetiolate; without cross venation; persistent; rolled in bud. Ligule an unfringed membrane.

Reproductive organization. Plants bisexual, all with bisexual spikelets; with hermaphrodite florets.

Inflorescence. Inflorescence of spicate main branches; digitate (rarely), or non-digitate. Primary inflorescence branches inserted all around the main axis. Rachides hollowed. Inflorescence espatheate; not comprising ‘partial inflorescences’ and foliar organs. Spikelet-bearing axes persistent. Spikelets solitary; secund; biseriate. Pedicel apices discoid.

Female-fertile spikelets. Spikelets 1.6–5.4 mm long; oblong, or elliptic, or lanceolate, or ovate, or obovate; adaxial; compressed dorsiventrally; biconvex; falling with the glumes. Rachilla terminated by a female-fertile floret. Hairy callus absent.

Glumes one per spikelet (membranous); long relative to the adjacent lemmas; dorsiventral to the rachis (the one glume (upper) outside); awnless. Upper glume 4–5 nerved. Spikelets with incomplete florets. The incomplete florets proximal to the female-fertile florets. The proximal incomplete florets 1; epaleate; sterile. The proximal lemmas awnless; 0 nerved, or 2 nerved, or 4 nerved (the median lacking); decidedly exceeding the female-fertile lemmas; less firm than the female-fertile lemmas (membranous); not becoming indurated.

Female-fertile florets 1. Lemmas decidedly firmer than the glumes; smooth to striate; becoming indurated to not becoming indurated (papery to crustaceous); yellow in fruit, or brown in fruit; entire; blunt; awnless; hairy, or hairless (sometimes with papillae, macrohairs and microhairs); non-carinate; having the margins inrolled against the palea; with a clear germination flap; 4 nerved. Palea present; relatively long; entire; awnless, without apical setae; textured like the lemma; indurated, or not indurated. Lodicules present; 2; free; fleshy; glabrous. Stamens 3. Anthers not penicillate. Ovary apically glabrous. Styles free to their bases; free. Stigmas 2; white (e.g. A. rupestris), or red pigmented (usually?).

Fruit, embryo and seedling. Fruit small; ellipsoid; compressed dorsiventrally. Hilum short. Embryo large; waisted; without an epiblast; with a scutellar tail; with an elongated mesocotyl internode. Embryonic leaf margins overlapping.

Abaxial leaf blade epidermis. Costal/intercostal zonation conspicuous. Papillae absent. Long-cells similar in shape costally and intercostally, or markedly different in shape costally and intercostally (the costals much narrower); of similar wall thickness costally and intercostally. Mid-intercostal long-cells rectangular, or fusiform (slightly); having markedly sinuous walls. Microhairs present; panicoid-type; 39–45 microns long; 7.5–9 microns wide at the septum. Microhair total length/width at septum 4.3–5.2. Microhair apical cells 21–30 microns long. Microhair apical cell/total length ratio 0.54–0.67. Stomata common; 36–45 microns long. Subsidiaries triangular. Guard-cells overlapping to flush with the interstomatals. Intercostal short-cells common; in cork/silica-cell pairs; silicified. Costal short-cells conspicuously in long rows. Costal silica bodies ‘panicoid-type’; dumb-bell shaped, or butterfly shaped and dumb-bell shaped.

Transverse section of leaf blade, physiology. C4; biochemical type NADP–ME (1 species); XyMS–. PCR sheath outlines uneven. PCR sheath extensions present, or absent. Maximum number of extension cells 2. PCR cell chloroplasts with reduced grana; centrifugal/peripheral. Mesophyll with non-radiate chlorenchyma. Leaf blade ‘nodular’ in section; with the ribs more or less constant in size. Midrib conspicuous; having a conventional arc of bundles; with colourless mesophyll adaxially. Bulliforms present in discrete, regular adaxial groups; in simple fans. All the vascular bundles accompanied by sclerenchyma. Combined sclerenchyma girders present; nowhere forming ‘figures’. Sclerenchyma all associated with vascular bundles.

Phytochemistry. Leaves without flavonoid sulphates (1 species).

Cytology. Chromosome base number, x = 10. 2n = 20, 40, 60, and 80. 2, 4, 6, and 8 ploid.

Classification. Watson & Dallwitz (1994): Panicoideae; Panicodae; Paniceae. Soreng et al. (2015): Panicoideae; Panicodae; Paspaleae; Paspalinae. 114 species.

Distribution, phytogeography, ecology. Tropical South America.

Commonly adventive. Helophytic to mesophytic; species of open habitats; glycophytic. Savanna, forest clearings, moist and weedy places.

Economic aspects. Significant weed species: A. affinis, A. compressus. Cultivated fodder: A. affinis, A. compressus. Important native pasture species: A. affinis, A. flexuosus. Lawns and/or playing fields: A. compressus.

Rusts and smuts. Rusts — Physopella and Puccinia. Taxonomically wide-ranging species: Puccinia levis. Smuts from Ustilaginaceae. Ustilaginaceae — Sorosporium and Sphacelotheca.

References, etc. Leaf anatomical: studied by us - Axonopus affinis Chase.

Illustrations. • Axonopus affinis: Gibbs Russell et al., 1990. • Female-fertile lemma of A. affinis. • Axonopus compressus: Gardner, 1952. • Inflorescence detail (A. compressus). • Inflorescence detail (A. compressus). • Inflorescence detail (A. compressus). • Spikelet in situ (A. compressus). • Opened spikelet of A. compressus. • Caryopsis detail, embryo (Axonopus compressus). Axonopus compressus. ‘Waisted’ embryo. • Axonopus fissifolius: Sanchez-Ken, Flora del Valle Tehuacán-Cuicatlan (2011). • Axonopus flexuosus: Clayton, © Flora Zambesiaca 10 (1989). • Axonopus affinis, abaxial epidermis of leaf blade: this project. • Axonopus affinis, 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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