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Key to Cyperus, Key ZD: Key to Cyperus, subgenus Cyperus -- Spikelets quadrangular to subterete; scales 1-8, persistent rachillas deciduous, articulate at the base

Cyperaceae

Cyperus

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(c) Montes de Oca, Joseph - CC-BY-NC, permission granted to NCBG
(c) Bradley, Keith
1 Culms, rays, and leaves bluish-glaucous, papillose; leaves harshly scabrid on margins and midrib; spikes with numerous well-developed branches or secondary rays; spikelets dark reddish brown at maturity
1 Culms, rays, and leaves green, not papillose; leaves glabrous to scabridulous; spikes solitary or with a few reduced basal branches; spikelets various.
(c) Fleming, Gary P.
  2 All but the uppermost spikelets strongly (-weakly in some forms of C. hystricinus) reflexed; scales (3.8-) 4-4.9 (-5.4) mm long, terminal involute and prolonged distally, midrib often bent downward; achenes linear to linear-oblong, (2.2-) 2.5-3 (-2.3) mm long; [section Umbellati, subsection Longistylis, C. retrofractus complex]
    3 Spikes mostly at least 2× as long as wide, slightly (or not) narrowed to base, spikelets reflexed to spreading; culms, leaves, and bracts glabrous except for marginal prickles on leaves and bracts; rachilla wing clasping achene for its entire length (concealing it in abaxial view); floral scales golden-brown when mature
    3 Spikes mostly about as long as wide and strongly narrowed to the base, spikelets reflexed; culms scabrous, bracts scabridulous and also often hirtellous; rachilla wing clasping achene for <3/4 its length; floral scales greenish to brown or reddish-purple-striped.
      4 Rays scabrid throughout their lengths; leaves and bracts puberulent; longest ray equal to or longer than the longest bract; culms roundly trigonous to subterete; terminal scale spinose distally; scales 1-2 (-3); spikes tight, burlike, turbinate
      4 Rays glabrous or rarely with a few hairs just below the spike; leaves and bracts with hairs confined to abaxial midrib; longest ray shorter than longest bract; culm sharply trigonous; terminal scale involute but not spinose; scales (2-)3-6; spikes often loose, sometimes more elongate
(c) Campos, Aidan
(c) Fleming, Gary P.
  2 Spikelets ascending to spreading, or reflexed in the lower half of spike; scales 1.8-5.3 mm long, terminal scale not modified as above; achenes ellipsoid to narrowly oblong, 1.2-2.4 (-2.6) mm long (linear-oblong in C. refractus).
(c) Mesaglio, Thomas - CC-BY
        5 Spikes densely cylindric, all sessile or 1-3 (-5) lateral spikes on short horizontal rays; spikelets ellipsoid (rarely oblong-ellipsoid), with 1-2 (-4) scales; [section Umbellati, subsection Brevistyli].
          6 Spikes dense, the rachis obscured when pressed; achenes lacking a stipitate base, apiculate at the apex; [native of Mexico and Central America, introduced northward along the Gulf and Atlantic coasts]
          6 Spikes relatively loose, the rachis usually visible when pressed; achenes cuneate to a stipitate base, broadly rounded to acute at the apex; [native, s. TX, uncertain reports]
(c) Fleming, Gary P.
        5 Spikes oblong to ovoid (cylindric in C. tetragonus), primarily on well-developed rays; spikelets oblong-lanceoloid to linear, with (1-) 2-8 scales.
             7 Spikelets very closely spaced, obscuring the rachis in life and when pressed; [section Umbellati, subsection Brevistylis]
               8 Spikes broadly and appearing (because of longer spikelets) relatively loosely ellipsoid to ovoid-cylindric, 2.5-7 cm long × 1.5-6 cm wide; spikelets (11-) 12-25 (-28) mm long; achenes 2.2-3 mm long (if < 2.5 mm long, noticeably falcate).
                 9 Spikes dense, ellipsoid to obovoid, convex, with 50-90 spikelets; scales 3-6 (-7), each 4-4.6 mm long, not red-purple striate; achenes 3-4× as long as wide, falcate
                 9 Spikes loose, broadly ovoid or ellipsoid to ovoid-cylindric, with 13-75 spikelets; scales 4-8(-11), each 4.5-5.3 mm long, red-purple striate; achenes 4-5× as long as wide, straight
               8 Spikes tightly globose or to oblong, not more than 2.5 cm long or 2 cm wide; spikelets 2-8 mm long; achenes < 2.3 mm long, straight.
(c) Ward, Scott G
(c) Fleming, Gary P.
                   10 Spikes uniformly convex throughout, globose or globose-ellipsoid, so dense as to appear nearly smooth in silhouette; scales 3.5-4.6 (-4.8) mm long, acute
(c) Bradley, Keith
                   10 Spikes cylindric, oblong, ellipsoid, or ovoid, mostly less dense; scales 1.8-3.6 mm long (or, if longer, mucronate).
                     11 Terminal scale of spikelet reduced, sterile, involute, with a minute but distinct mucro to 0.3 mm long, usually uncinate (hooked); fertile scales (if more than one) not overlapping on the same side of the rachilla, usually red-brown or orangeish laterally at maturity; leaf blades 1-3 mm wide; [Cyperus retroflexus complex]
                       12 Fertile floral scales 1.9-3.0 (-3.3) mm long; rachilla wing usually membranaceous throughout; rachilla lacking lateral nerves; longest spikelets 2.8-5.8 (-8.0) mm long; terminal sterile floral scale of spikelet often much reduced, less than 2/3 the length of fertile floral scales; longest peduncle less than 2.7 (-3.9) cm long; plants diminutive, culms 3-35 (-45) cm tall
                       12 Fertile floral scales (2.8-) 3.0-3.9 mm long; rachilla wing usually chartaceous beyond clasped achene angle, border membranaceous; rachilla with two lateral nerves, one along each side of the median; longest spikelets 4.9-9.0 (-11.3) mm long; terminal sterile floral scale usually not greatly reduced, 2/3 or more the length of fertile floral scales; longest peduncle (0.5-) 2.4-6.8 cm long; plants more robust, culms usually >25 cm tall
                     11 Terminal scale of spikelet not differentiated; scales acute, greenish-hyaline to brownish or straw-colored laterally; scales overlapping; leaf blades 1.5-5 mm wide.
                          13 Spikes relatively loosely ovoid, 8-20 mm wide, with (5-) 10-50 (-70) spikelets, lacking basal branches; spikelets (2.2-) 3.5-10 mm long, angular in ×-section, greenish-hyaline to pale yellow-green; bracts horizontal
                          13 Spikes densely oblong-ellipsoid to ovoid, 6-8 (-10) mm wide, with (13-) 30-200 (-274) spikelets, occasionally with short basal branches; spikelets 2.2-4 (-4.5) mm, subterete, straw-colored to brown at maturity; bracts horizontal or ascending to erect; [Cyperus retrorsus complex]
                            14 Achenes oblong-fusiform, stipitate, narrowed to both ends; spikes densely oblong to oblong-cylindric, often 3× or more as long as wide; scales ascending (the tips free), usually straw-colored; bracts horizontal
                            14 Achenes oblong, usually abruptly constricted at the ends; spikes ovoid to oblong, to 2× as long as wide (rarely longer in C. retrorsus); scales appressed, usually reddish-brown at maturity; bracts ascending to erect (sometimes horizontal in C. nashii).
                              15 Spikes > 1.5× as long as wide, each with (42-) 50-200 (-274) spikelets; achenes 0.4-0.5 (-0.6) mm wide; stems moderately cespitose; [widespread and weedy]
                              15 Spikes < 1.5 × as long as wide, each with (13-) 30-92 spikelets; achenes 0.5-0.7 mm wide; stems single to slightly cespitose; [Florida scrub, northward and westward in coastal dunes]
                                16 Rays 3-6 (-8); anthers 0.2-0.5 (-0.7) mm long; achenes 1.3-1.6 mm long × 0.5-0.6 mm wide
             7 Spikelet attachments spaced apart at least proximally, rachis not obscured when pressed and usually also visible in life.
                                  17 Fertile floral scales 1-3 (-5), not overlapping on the same side of the spikelet (if more than one); terminal scale reduced, sterile and involute, mucro usually uncinate; achenes (1.7-) 1.9-2.6 mm long; [section Umbellati, Cyperus retroflexus complex]
                                    18 Longest spikelets 10-21 mm long, strongly contorted; distal scales with scabrid midrib and mucro 0.6-1.9 mm long; longest floral scale of spikelet (3.5-) 3.7-4.8 mm long; anthers 0.5-1.3 mm long; achenes > 3× as long as wide; [TX]
                                    18 Longest spikelets usually < 10 mm long, at most somewhat flexuous with curved tips; distal scales with glabrous midrib and mucro 0.1-0.3 (-0.5) mm long; longest floral scale of spikelet (2.1-) 2.5-3.5 (-4.0) mm long; anthers 0.3-0.5 (-0.6) mm long; achenes 2-3 (-3.3)× as long as wide; [more widespread]
                                  17 Fertile floral scales 3-8, reaching the next scale or overlapping on the same side of the spikelet; terminal scale not involute and sterile, mucro straight; achenes various.
                                         20 Spikelets ±terete; scales 4.5-5.3 mm long, persistent, laterally reddish brown or tinted with reddish brown if paler and stramineous; achenes usually at least half as long as floral scale; achenes linear-oblong, 2.6-3 mm long, 5-6× as long as wide, > 0.5 mm wide; anthers 1-1.5 mm long; spikes usually broadly and convexly ellipsoid or ovoid, usually unbranched (or rarely with 1-2 reduced proximal branches); [section Umbellati, subsection Brevistylis]
                                         20 Spikelets flattened; scales 3.2-4.5 (-6) mm long, laterally golden yellow to stramineous; achenes less than half as long as floral scale; achenes oblong, (1.5-) 1.8-2.4 mm long, 3-4× as long as wide, < 0.4 mm wide; anthers 0.3-0.5 mm long; spikes usually parallel-sided, ovoid-cylindric, frequently branched; [section Strigosi]
                                           21 Plants perennial, ≥ 3 dm tall; culm 1.8-3 mm in diameter; floral scales not aristate
                                       19 Scales (2-) 2.5-3.2 (-4) mm long, acute to cuspidate; spikes narrowly cylindric to ovoid, spikelets mostly 4-8mm long (if longer, scales mucronate); achenes ellipsoid, 2-3× as long as wide; culm bases not cormose.
                                               23 Spikelets oblong to linear, 4-10× as long as wide; achenes ellipsoid to narrowly oblong; spikes narrowly to broadly cylindric to oblong or ellipsoid, ≥ 12 mm wide (C. retroflexus occasionally as narrow as 8 mm wide, but not cylindric).
                                                 24 Floral scales pale olivaceous to yellowish, not mucronate (rarely with a mucro to 0.1 mm long); spikes cylindric, loose, 6-10 spikelets per 5 mm span of the rachis; culms 0.5-1 mm in diameter; leaves 0.8-2.3 (-3) mm wide; [more widespread]; [section Tetragoni]
                                                 24 Floral scales straw-colored to reddish-brown, acute or mucronate, mucro up to 0.2 mm long; spikes loosely to densely ovoid to oblong, spikelets 8-21 per 5 mm span of the rachis; culms 1-2.6 mm in diameter; leaves (1-) 3-6 (-10) mm wide; [TX].
                                                    25 Floral scales not mucronate; spikes mostly narrowly oblong (usually 2× or more as long as wide); rachis, spikelets, bracteoles, peduncles, and prophylls not reddish brown striate or maculate (at 10× magnification); [section Tetragoni]
                                                    25 Floral scales mucronate, mucro at least 0.2 mm long; spikes densely ovoid to loosely ellipsoid-oblong (to 1.5× as long as wide); rachis, spikelets, bracteoles, peduncles, and prophylls usually conspicuously reddish brown striate or maculate (at 10× magnification); [section Strigosi].
                                                      26 Spikes denser, spikelets 11-21 per 5 mm span of the rachis (at least in the upper portion), often the upper spikelets ascending; scale color bilaterally variable, reddish or reddish-brown to whitish; spikelets estipitate or stipe to 0.2 mm long; spikes ellipsoid to oblong
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Key to Cyperus, Key ZE: Key to Cyperus, subgenus Cyperus of various affinities -- Spikelets compressed; scales deciduous or persistent; rachilla persistent or deciduous after the scales

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1 Culms and leaves bluish-glaucous in life; scales overlapping, slightly spreading, glossy reddish brown at maturity; plants cespitose perennials; [FL; beaches, dunes, coastal rock]
  2 Rays absent, inflorescence a single globose spike, rachis only 1-3 mm long; scales 1.5-2.2 mm long; achenes 1-1.2 mm long, glabrous, puncticulate, or papillose; leaves and bracts 1-2.5 mm wide; culms 0.5-1 mm in diameter; [section Laxiglumi]
  2 Rays present, inflorescence of multiple well-developed spikes, though often very congested; scales 2.5-3 mm long; achenes 1.3-2 mm long, coarsely punctate; leaves and bracts (1.5-) 3-10 mm wide; culms 0.8-2.6 mm in diameter; [section Thunbergiani]
1 Culms and leaves green, not glaucous; scales divaricate to closely imbricate, mostly greenish, light brown, or straw-colored (except C. rotundus and C. setigerus); [more widespread].
    3 Rachilla wingless (except C. prolixus and C. grayi); spikelets at maturity 2-4 mm wide, scales divaricate (the spikelet appearing coarsely serrate in outline).
      4 Rachis hispidulous; spikes loosely oblong-ovoid, with spikelets spreading, the elongate, well-developed rachis clearly visible; scales 1.8-2 mm long, light brown to yellowish; achenes 1-1.2 mm long; [sect. Proceri]
      4 Rachis glabrous; spikes densely ovoid or globose, or umbellate to oblong, the rachis elongate to nearly absent; scales 1.8-3.2 mm long; achenes 1.5-2.4 mm long (1-1.2 mm long in C. cephalanthus).
        5 Rays compressed; plants robust, up to 2m tall; inflorescence diffuse, 2nd-order rays present, erect, 1-10 cm long, usually bearing well-developed 3d-order rays 0.5-5 cm long; [sect. Distantes]
        5 Rays terete to trigonous; plants mostly less robust, to 1m tall; inflorescence more compact, 2nd- and 3rd-order rays absent or poorly developed.
          6 Plants robust, culms 40-100 cm tall, sharply trigonous, scabrid to hirtellate on the angles; spikes very densely ovoid, with 25-50+ spikelets, subtended by well-developed secondary bracts and often branched at the base; achenes 1.0-1.2 mm long, ellipsoid, <1/2 as long as the scales; [coastal prairies; LA, TX]; [sect. Thunbergiani]
          6 Plants smaller, culms to 50 cm tall, terete to trigonous, not scabrid (except C. schweinitzii); spikes relatively loose, with 3-24 spikelets, unbranched and lacking secondary bracts; achenes 1.5-2.2 mm long, mostly ≥ 1/2 as long as the scales; [dry, sandy or rocky soils; collectively widespread]; [sect. Laxiglumi].
             7 Inflorescence bracts deflexed (to horizontal); spikelets in a single sessile hemispheric to globose capitate cluster, or with an additional 1-4 smaller pedunculate spikes, these densely globose with an obscure rachis < 6 mm long; [Cyperus lupulinus complex]
               8 Anthers 0.8-1 mm long; achenes 0.5-0.8 mm wide, narrowly oblong; scales usually yellowish to yellowish brown; leaves very narrow, 0.5-2 mm wide
               8 Anthers 0.3-0.6 mm long; achenes 0.8-1.2 mm wide, ±ellipsoid; scales usually off-white to pale tan to light reddish brown; leaves often broader, 1-4 mm wide.
                 9 Scales 2.5-3.8 mm long, usually fitting loosely over the mature achene, the margins spreading or loosely clasping it; spikelet with 5-22 scales
                 9 Scales 1.8-2.5 mm long, usually fitting tightly over the achene, the margins tightly clasping it; spikelets with 3-7 scales
             7 Inflorescence bracts horizontal to erect; spikes primarily pedunculate, loosely globose, densely ovoid, or more elongate.
                   10 Scales 2.2-3.2 mm long with a conspicuous cusp > 0.3 mm long (at least on the distal scales); body of scale 2.2-3.2 mm long; spikes umbellate to oblong, spikelets all erect-ascending; achenes 2-2.4 mm long, ellipsoid; plants rhizomatous, culms sharply trigonous
                   10 Scales 1.3-2.5 mm long, acute or minutely mucronate (mucro ≤ 0.2mm long); spikes mostly globose or ovoid, spikelets usually spreading or radiating in all directions (erect-ascending in C. houghtonii); achenes 1.5-2.4 mm long; plants various.
                     11 Spikelets mostly erect-ascending, the spikes umbellate to broadly ovoid; scales with distinct mucro 0.1-0.2 mm long; achenes 1-1.5 mm wide, broadly ellipsoid; anthers 0.5-0.7 mm long
                     11 Spikelets radiating in all directions, the spikes globose or subglobose; scales acute; achenes 0.7-1.2 mm wide, narrowly ellipsoid or oblong-ellipsoid; anthers 0.7-1.2 mm long.
                       12 Rachilla winged, wings 0.6 mm wide; achenes 1.5-2 mm long × 0.7-1 mm wide; scales 2-2.8 mm long; anthers 0.7-1 mm long; [Atlantic coast, south to AL]
                       12 Rachilla wingless; achenes 2-2.4 mm long × 0.9-1.2 mm wide; scales 2.5-3 mm long; anthers 1-1.2 mm long; [Midwest south to LA]
    3 Rachilla winged, wings at least 0.4 mm wide; spikelets at most 2.2 mm wide, scales overlapping, tips closely imbricate, the margin appearing smooth or slightly serrate (some forms of C. strigosus have looser scales).
                          13 Scales 1-1.5 mm long, deciduous; spikes cylindric, with (20-) 40-80+ spikelets, often with secondary rays; fibrous-rooted annuals, but often robust, roots reddish; [sect. Fastigiati]
                            14 Plants with cormose culm bases; scales 3.2-4.5 (-6) mm, 3-11 per spikelet, pale greenish-brown to straw-colored, dull; spikelets strongly compressed (at maturity), the scales loosely overlapping, tips spreading; achenes narrowly oblong, (1.5-) 1.8-2.4 mm long; [section Strigosi]
                            14 Plants lacking cormose culm bases; scales 1.8-4 (-4.4) mm long, 6-36 (-42) per spikelet, various; spikelets often only slightly compressed, scales closely imbricate, tips appressed; achenes ellipsoid to oblong, 1.1-2 mm long.
                              15 Scales pale greenish-white, deciduous, usually with dark-speckled margins visible as a prominent dark line on each side of the spikelet; plants fibrous-rooted; anthers 0.6mm long, styles 0.6-1.2 mm long, stigmas 1-1.4 (-1.7) mm long; [rare introduction, FL]; [section Subimbricati]
                              15 Scales pale yellowish to brown or deep purple, persistent or deciduous; plants with well-developed rhizomes or stolons; anthers 1-2.5 mm long, styles (0.7-) 1-3.6 mm long, stigmas (1.2-) 1.8-6 mm long; [collectively widespread]; [section Cyperus].
                                16 Culms terete at least towards the base, nodose-septate, with internal partitions every 5-50 mm, more frequent upwards; leaves usually all reduced to bladeless sheaths; bracts 2 (-4), erect, reduced, mostly < 2 cm long
                                  17 Scales deciduous, (2.2-) 2.7-4.5 (-6) mm, mostly reddish-brown; robust perennials 5-10dm tall from thick, indurate rhizomes, lacking tubers; achenes maturing, oblong
                                  17 Scales persistent, 1.8-3.4 mm long, mostly yellowish or dark glossy purplish; plants mostly 1-6 dm tall (rarely taller in C. esculentus), with slender stolons 1-2 mm in diameter, these producing terminal tubers; achenes ellipsoid or not maturing.
                                    18 Spikelets (2-) 3-14 per spike, bracts often not exceeding the inflorescence; scales glossy dark purplish, with green keels and hyaline border; culm bases indurate, stolons wiry, horizontal, 1-2 mm in diameter, producing irregular, often elongate tubers
                                    18 Spikelets (3-) 10-28 per spike, bracts exceeding the inflorescence; scales yellowish to dull reddish-brown; culm bases not indurate, stolons spongy, weak, 1 mm in diameter, producing globose tubers; [Cyperus esculentus complex].