Ichnanthus hoffmannseggii ( Roemer & Schultes 1817: 450 ) Döll (1877: 287)

Silva, Christian, Davidse, Gerrit, Schnadelbach, Alessandra Selbach & Oliveira, Reyjane Patrícia De, 2016, Systematics of Ichnanthus hoffmannseggii and allies (Poaceae, Paspaleae) based on molecular and morphological evidence, Phytotaxa 267 (4), pp. 263-278 : 265-273

1. Ichnanthus hoffmannseggii ( Roemer & Schultes 1817: 450) Döll (1877: 287) . ( Figs. 1 , 4A,B , 5 , and 7A− F)

Basionym: Panicum hoffmannseggii Roemer & Schultes (1817: 450) .

Lectotype:— BRAZIL. Pará, s.d., Sieber s.n. (B!), designated by Stieber (1982: 94), corrected from “ holotype ” . Isolectotypes: BM!, BR!, US-78075! (fragm. ex B).

Panicum villosum Hoffmanns. ex Roemer & Schultes (1817: 450) , nom. inval., as synonym of Panicum hoffmannseggii .

Panicum eriophorum Link ex Nees (1829: 148) , nom. inval., as synonym of Panicum hoffmannseggii .

Annual plants, herbaceous, caespitose, without rhizomes, 0.15−0.85 m tall. Culms erect or decumbent and geniculately ascending, rooting at the lower nodes; internodes sparsely to densely pilose, trichomes 0.4−2.2 mm long, slightly papillose-based or not papillose-based; nodes densely pilose, trichomes 0.4−3.6 mm long, papillose-based or not papillose-based. Leaves distributed along the culms; leaf sheaths entirely densely pilose, rarely with short trichomes on the surface and longer trichomes on the margins and towards the base, trichomes 0.3−7.5 mm long, not papillose-based to strongly papillose-based; collar pilose, trichomes 0.4−2.4 mm long, not papillose-based to strongly papillose-based; leaf blades 2.5−12.5 × 0.4−2.9 cm, ovate-lanceolate, elliptic-lanceolate to broadly lanceolate, apex acuminate, base rounded to subcordate, margins slightly to strongly sclerified and scabrous, densely pilose on both surfaces, rarely with sparse and short trichomes on both surfaces, trichomes 0.25−6.4 mm long, not papillose-based to strongly papillose-based; pseudopetioles absent; ligules membranous-ciliate, 1.7−2.9 mm long, membranous portion 0.1−0.4 mm. Inflorescence paniculate, 4.5−17 × 2−12 cm, open, terminal and axillary, partially included to entirely exserted from the leaf sheaths, 1(−2) inflorescences per sheath; peduncle 2−11.5 cm long; primary branches alternate, perpendicular to oblique to the main axis, the longest ones 2−9 cm long, the lower ones longer than the apical ones; lower levels of branching alternate, secund, the longest branches 1.5−2.8 cm long, the lower branches longer than the apical ones; peduncle, main axis, and branches densely pilose, trichomes 0.3−6 mm long, papillose-based, rarely not papillose-based; spikelets in pairs, the lower ones with pedicels 0.5−1.1 mm long, the apical ones with pedicels 1.6−6.8 mm long. Spikelets 3.5−4.5 × 0.9−1.2 mm, elliptic-lanceolate, regularly distributed along the branches, green to stramineous, densely pilose, rarely glabrescent, trichomes 0.3−3.8 mm long, strongly papillose-based, rarely not papillose-based; internodes between glumes conspicuous, ca. 0.2 mm long; lower glume 2.6−4.5 mm long, shorter, equal to or slightly longer than the upper one, 3-nerved, apex acuminate to shortly caudate; upper glume 3.1−4.4 mm long, slightly shorter to slightly longer than the anthecia, 5(−7)-nerved, apex acute to acuminate; lower anthecium staminate or sterile; lower lemma 3−4 mm long, 5-nerved, apex acute to acuminate, glabrous; palea 3/4 of the length of the lemma, hyaline; upper anthecium 2−2.5 × 0.6−1.1 mm, elliptic to widely elliptic at maturity, white to dark brown with dark spots at maturity, glabrous, smooth, with papillae at the base of the upper lemma, and at the base and apex of the upper palea, conspicuous only at maturity; upper lemma slightly curved at the apex; rachilla appendages 0.4−0.7 mm long, winged, membranous and hyaline in the young anthecium, fleshy and light brown at maturity, ca. 1/4 the length of the anthecium, apex rounded, base slightly narrower than the apex, emerging from a curved stipe (rachilla) present at the base of the upper lemma, ca. 0.1 mm long, inconspicuous; lodicules 2, ca. 0.4 mm long; stamens 3, filaments white, anthers white or yellow; stigmas 2, vinaceous. Caryopsis ca. 1.9 × 1 mm, elliptic to slightly obovate, pale yellow; hilum linear, ca. 1/2 the length of the caryopsis.

Distribution and habitat:— Ichnanthus hoffmannseggii occurs in open to partially shaded and sandy areas ( Fig. 4A ) of the Brazilian Cerrado and Amazonia, where it is usually common. It was previously recorded for the Brazilian States of Pará, Maranhão, Tocantins, and Goiás ( Fig. 5 ) ( Silva et al. 2016). New records for the Brazilian States of Piauí and Bahia are reported in the present study ( Fig. 5 ).

Notes:— Roemer & Schultes (1817) described Panicum hoffmannseggii based on “ Panicum villosum Com. de Hoffmannsegge ”, which is the identification of the specimen collected by F.W. Sieber for J.C. von Hoffmannsegg in Pará ( Brazil) between 1801−1807 ( Urban 1906). This specimen, housed at B, was considered the holotype by Stieber (1983: 94), but because of the existence of duplicates at BR and US, and the uncertainty of which specimen(s) were actually used by Roemer and Schultes to describe their new species, Stieber’s designation of a holotype is best considered as an error to be corrected to a lectotypification (Art. 9.9, Melbourne Code). We do not consider the specimen “ J.C. von Hoffmannsegg am Link ” (LE-TRIN 748.1−2!) cited by Zuloaga et al. (2003) as a type because its origin is unknown. It is a mixed collection with one large individual of Ichnanthus mollis Ekman (1911: 20) (bottom left corner) and another small one of I. hoffmannseggii (top right corner). The locality, S. da Chapada [Mato Grosso], associated with this specimen by Zuloaga et al. (2003) is applicable only to I. mollis since it is known from this state, and I. hoffmannseggii has, so far, never been collected there. A new isolectotype was found in BM.

Representative specimens examined:— BRAZIL. Bahia: Formosa do Rio Preto, Fazenda Gentilio, 590 m, 11°11’50”S, 45°27’46”W, 22 April 1998 (fl), Azevedo et al. 1315 ( IBGE, MO! 2 sheets, SP!). Goiás: Pirenópolis, km 66 da estrada Padre Bernardo/Niquelândia, 15 March 1995 (fl), Pereira et al. 2729 ( IBGE, MO!, SP!). Maranhão: Balsas, 34.7 km along road from Balsas to São Raimundo das Mangabeiras, 250 m, 7°20’S, 46°0’W, 15 March 1962 (fl), Eiten & Eiten 3641 (F!, SP!) ; Balsas, Condomínio Kissy lote 23, 20 March 1999 (fl), Pereira-Silva 4123 ( CEN!) ; Carolina, Pedra Caída, Base of Morro da Baleia , 7°8’S, 47°25’W, 14 April 1983 (fl), Taylor et al. 1236 ( MO!, NY) ; Timon, BR-316 Timon-Caxias, km 40, 28 May 1980 (fl), Coradin et al. 2637 ( CEN!, SP!). Pará: Monte Alegre, alto da serra do Ereré , 15 May 1953 (fl), Lima 53-1602 ( RBR!) ; Santarém , March 1850 (fl), Spruce s.n. (‘ Panicum 31’) (F! 2 sheets, GH!, MO!) ; Santarém, Alter do Chão , savana amazônica, 20 June 1988 (fl), Miranda 94 ( CEN!, INPA) ;

Santarém , área de cerrado próximo ao aeroporto, 32 m, 2°26’18”S, 54°47’37”W, 26 May 2014 (fl), Silva et al. 1163 ( CANB!, CEN!, F!, GH!, HSTM!, HUEFS!, ICN!, K!, MBM!, MG!, MO!, NY!, RB!, RBR!, SP!, US!) ; Santarém , próximo a Alter do Chão, 52 m, 2°32’9”S, 54°55’22”W, 27 May 2014 (fl), Silva et al. 1184 ( AAU!, BAA!, CANB!, CEN!, CEPEC!, F!, GH!, HSTM!, HUEFS!, IBGE!, ICN!, INPA!, K!, MBM!, MG!, MO!, NY!, P!, R!, RB!, RBR!, SP!, UB!, US!) ; Santarém , estrada para Ponte de Pedra, 56 m, 2°28’14”S, 54°54’36”W, 28 May 2014 (fl), Silva et al. 1196 ( AAU!, BAA!, CANB!, CEN!, CEPEC!, F!, GH!, HSTM!, HUEFS!, IBGE!, ICN!, INPA!, K!, MBM!, MG!, MO!, NY!, P!, R!, RB!, RBR!, SP!, UB!, US!) ; Soure , Fazenda Camburupy, 19−20 June 1934 (fl), Swallen 4945 ( GH!, RB!, SP!) ; Vigia , Campina do Palha, 10 August 1954 (fl), Black 54-16746 ( ALCB!, GH!, IAN) ; Vigia , 9 km SW of Vigia along PA-140 to Belém, 50 m, 0°55’S, 48°04’W, 31 March 1980 (fl), Davidse et al. 17694 (F!, MO!, NY, MG, US!). Piauí: Floriano, estrada para Itaueira, PI- 140, 562 m, 6°48’1”S, 43°01’42”W, 15 May 2012 (fl), Silva et al. 854 ( HUEFS!). Tocantins: Almas, RPPN Fazenda Minehaha, 11°8’48”S, 47°8’37”W, 21 April 2004 (fl), Felfili et al. 516 ( IBGE, ICN!) ; Babaçulândia , km 64 da BR-226 , ca. 7 km NE de Wanderlândia, 350 m, 6°50’S, 47°54’W, 16 March 1985 (fl), Valls et al. 8342 ( CEN!, ICN!, RB!, SP!, US!) ; Couto Magalhães , 16 km W of Piquizeiro along GO- 70, 400 m, 8°20’S, 49°10’W, 25 February 1980 (fl), Plowman et al. 9136 (F!, MG, MO!, NY, US!) ; Guaraí , rodovia Belém- Brasília, 29 March 1976 (fl), Hatschbach & Kummrow 38493 ( MBM!) ; Lagoa da Confusão , Ilha do Bananal, Parque Nacional do Araguaia, 250 m, 10°27’45”S, 50°28’32”W, 20 March 1999 (fl), Silva et al. 4026 ( IBGE, ICN!) ; Lajeado , estrada para Palmas, 9°58’58”S, 48°19’22”W, 28 March 2008 (fl), Santos & Pereira 1551 ( HUEFS!, HUTO) .

2. Ichnanthus oplismenoides Munro ex Döll (1877: 288) . Type:— BRAZIL. Tocantins: rio Tocantins, between São João and Funil, s.d. (fl), Burchell 9031 (holotype K!, isotypes BR! (2 sheets), GH!, K! (2 sheets), L!, LE, NY!, P!, S!, US-975271! (fragm. ex BR), US-975270! (fragm. ex LE), W!). ( Figs. 2 , 4C− E , 5 , and 7G− L)

Echinolaena oplismenoides (Munro ex Döll) Stieber (1987: 212) .

Annual plants, herbaceous, caespitose, without rhizomes, 0.15−1 m tall. Culms erect or decumbent and geniculately ascending, rooting at the lower nodes; internodes densely pilose, trichomes 0.4−4.5 mm long, papillose-based or not papillose-based; nodes densely pilose, trichomes 0.4−7.2 mm long, not papillose-based to strongly papillose-based. Leaves distributed along the culms; leaf sheaths densely pilose, rarely glabrous or with short trichomes on the surface and longer trichomes on the margins and towards the base, trichomes 0.3−6 mm long, not papillose-based to strongly papillose-based; collar pilose, trichomes (0.1−) 0.4−3 mm long, not papillose-based to strongly papillose-based; leaf blades 2−15 × 0.6−5 cm, ovate-lanceolate, elliptic-lanceolate to broadly lanceolate, apex acute to acuminate, base rounded to subcordate, margins strongly sclerified and scabrous, densely pilose on both surfaces, or glabrous, with or without trichomes along the margins abaxially, trichomes 0.3−6 mm long, not papillose-based to strongly papillose-based; pseudopetioles absent; ligules membranous-ciliate, 1−3.9 mm long, membranous portion 0.1−0.4 mm. Inflorescence paniculate, pseudoracemose, 4−32 × 1−4.5 cm, terminal and axillary, partially included to entirely exserted from the leaf sheaths, 1−3 inflorescences per sheath; peduncle 2−32 cm long; primary branches alternate, oblique to the main axis, the longest ones 1.5−4.5 cm long, the lower ones longer than the apical ones; lower levels of branching alternate, secund, contracted, the longest branches 0.9−1.5 cm long, the lower branches longer than the apical ones; peduncle, main axis, and branches sparsely to densely pilose, trichomes 0.1−6.3 mm long, not papillose-based to strongly papillose-based; spikelets in pairs, the lower ones with pedicels 0.2−0.5 mm long, the apical ones with pedicels 0.6−1.3 mm long. Spikelets 5.6−15 × 1.2−2 mm, ovate-lanceolate to lanceolate, regularly distributed along the branches, green to stramineous, densely pilose, trichomes 0.3−6 mm long, not papillose-based to strongly papillose-based; internodes between glumes conspicuous, ca. 0.2 mm long; lower glume 5−15 mm long, shorter, equal to much longer than the upper one, 3-nerved, apex acuminate to aristate; upper glume 5.1−7.7 mm long, longer than the anthecia, 5-nerved, apex acute to caudate; lower anthecium staminate or sterile; lower lemma 4.2−5 mm long, 5-nerved, apex acute, ciliolate on the margins at the apex; palea 3/4−4/5 of the length of the lemma, hyaline; upper anthecium 2.8−3.2 × 1−1.3 mm, elliptic to slightly obovate at maturity, white to dark brown with dark spots at maturity, glabrous, entirely covered with papillae, more conspicuous at maturity; upper lemma slightly curved at the apex; rachilla appendages 0.9−1.2 mm long, winged, membranous and hyaline in the young anthecium, fleshy and light brown at maturity, 1/4−1/3 the length of the anthecium, apex rounded, base slightly narrower than the apex, emerging from a curved stipe (rachilla) present at the base of the upper lemma, ca. 0.3−0.4 mm long, conspicuous; lodicules 2, ca. 0.3 mm long; stamens 3, filaments white, anthers yellow; stigmas 2, vinaceous. Caryopsis ca. 2.1−2.3 × 1.1−1.2 mm, elliptic, pale yellow; hilum linear, ca. 3/5 the length of the caryopsis.

Distribution and habitat:— Ichnanthus oplismenoides occurs in open and sandy areas ( Fig. 4C− E ) of the Brazilian Cerrado. It was previously recorded for the Brazilian States of Maranhão, Tocantins, and Goiás ( Fig. 5 ) ( Filgueiras & Oliveira 2016; as Echinolaena oplismenoides ). New records for the Brazilian States of Pará and Mato Grosso are reported in this study ( Fig. 5 ).

Notes:— As mentioned above, this species was transferred to Echinolaena by Stieber (1987) due to the presence of racemose inflorescences, lower glume equal to or exceeding the upper glume, and upper floret with winged appendages. However, it was recently reestablished in Ichnanthus in our previous phylogenetic study ( Silva et al. 2015) because it was part of a clade with other representatives of the genus, due to the presence of paniculate inflorescences (“pseudoracemose”, not racemose; see Figs. 2A,B , 4C− E ), and because winged appendages in the upper anthecium ( Figs. 2D− G , 7G− J ) are synapomorphic to Ichnanthus . Three new isotypes were found in GH, NY, and W respectively.

Representative specimens examined:— BRAZIL. Goiás: Posse, Fazenda das Araças, 765 m, 13°57’42”S, 46°22’11”W, 7 March 2001 (fl), Silva et al. 4908 ( CEN!, IBGE, MO!). Maranhão: Barra do Corda to Grajaú, 1−5 March 1934, Swallen 3663 ( GH!, RB!, SP!) ; Carolina to Balsas, 20−25 March 1934 (fl), Swallen 4063 ( SP!) ; Carolina, Serra Morro do Chapéu , 12 June 1966 (fl), Prance 2099 ( MO!, NY, US!) ; Carolina, rodovia MA-010, Serra Portal da Chapada , 296−350 m, 7°11’13”S, 47°25’22”W, 19 May 2012 (fl), Silva et al. 891 ( BAA!, CANB!, CEN!, CEPEC!, F!, GH!, HSTM!, HUEFS!, IBGE!, ICN!, INPA!, K!, MBM!, MG!, MO!, NY!, R!, RB!, SP!, US!) ; Carolina, MA-010, sentido Estreito, Balneário do Dodô , 231 m, 7°5’41.6”S, 47°26’38”W, 20 May 2012 (fl), Silva et al. 923 ( AAU!, BAA!, CANB!, CEN!, CEPEC!, F!, GH!, HSTM!, HUEFS!, IBGE!, ICN!, INPA!, K!, MBM!, MG!, MO!, NY!, P!, R!, RB!, RBR!, SP!, UB!, US!) ; Estreito, 220 m, 7°11’S, 47°25’W, 17 March 1985 (fl), Valls et al. 8366 ( CEN, ICN!, RB!). Mato Grosso: Santa Terezinha, Ilha do Bananal, Monte de Areia, 190 m, 10°27’S, 50°28’W, 21 March 1999 (fl), Silva et al. 4050 ( IBGE, MO!, US!). Pará: Alto dos Montes, região do Araguaia, 17 June 1953 (fl), Fróes 30068 ( SP!). Tocantins: Guaraí , campo cerrado, 26 March 1976 (fl), Hatschbach & Kummrow 38348 ( MBM!) ; Guaraí , 10 km N of Guaraí along BR- 153, 400 m, 8°44’S, 48°31’W, 26 February 1980 (fl), Plowman et al. 9160 ( MG, MO!, NY!, US!) ; Miracema do Tocantins , rodovia TO-010, 236 m, 9°36’8”S, 48°24’43”W, 1 February 2012 (st), Silva et al. 834 ( HUEFS!) ; Tocantinópolis, Ribeirão do Côrrego , 55 km SW of Estreito along the BR-153 highway, 480 m, 6°49’S, 47°49’W, 27 February 1980 (fl), Plowman et al. 9204 ( MO!, NY!, US!) .

3. Ichnanthus piresii Black (1950: 31) . Type:— BRAZIL. Maranhão: cerrado, a duas léguas de Carolina, encosta da Serra da Malícia, 30 July 1949 (fl), Pires & Black 1595a (holotype IAN!, isotypes RB!, US!). ( Figs. 3 , 4F− H , 5 , and 7M− R)

Annual plants, herbaceous, prostrate, without rhizomes, 0.1−0.15 m tall. Culms creeping, the flowering ones geniculately ascending, rooting at the lower nodes; internodes glabrous to pilose, trichomes ca. 0.3 mm long, slightly papillose-based; nodes densely pilose, trichomes 0.3−2.5 mm long, slightly papillose-based. Leaves distributed along the culms; leaf sheaths densely pilose, trichomes 0.1−0.2 mm long, longer on the margins and towards the apex (1−2 mm long), slightly papillose-based; collar pilose, trichomes ca. 0.3 mm long, slightly papillose-based; leaf blades 1−7 × 0.3−1.9 cm, ovate to ovate-lanceolate, apex acuminate, base rounded, margins slightly sclerified and finely scabrous, densely pilose on both surfaces, trichomes 0.2−0.4 mm long, slightly papillose-based; pseudopetioles absent; ligules membranous-ciliate, ca. 1.8 mm long, membranous portion ca. 0.1 mm. Inflorescence paniculate, pseudoracemose, 1.5−8 × 0.4−3 cm, terminal and axillary, partially included to entirely exserted from the leaf sheaths, 1−2 inflorescences per sheath; peduncle 0.5−11.8 cm long; primary branches alternate, perpendicular to oblique to the main axis, the longest ones 0.7−2.5 cm long, the lower ones longer than the apical ones; lower levels of branching alternate, secund, contracted, the longest branches ca. 1.8 mm long, inconspicuous to the naked eye, the lower branches longer than the apical ones; peduncle, main axis, and branches densely pilose, trichomes 0.3−1.7 mm long, papillose-based; spikelets in pairs, the lower ones with pedicels ca. 0.5 mm long, the apical ones with pedicels ca. 0.7 mm long. Spikelets 2.3−4.8 × 0.7−1.8 mm, ovate-lanceolate to elliptic-lanceolate, regularly distributed along the branches, green to stramineous, pilose; internodes between glumes slightly conspicuous, ca. 0.1 mm long; lower glume 2−4.8 mm long, subequal to longer than the upper one, 3-nerved, apex long acuminate to caudate, densely pilose, trichomes 0.1−1.2 mm long, papillose-based; upper glume 2.2−3 mm long, slighlty longer than the anthecia, 5-nerved, apex acuminate, densely pilose, trichomes 0.1−1.2 mm long, papillose-based; lower anthecium sterile; lemma 2−2.4 mm long, 5-nerved, apex acute to acuminate, densely pilose along the margins and at the apex, trichomes 0.1−0.8 mm long, papillose-based; palea 3/4 the length of the lemma, hyaline; upper anthecium 1.3 × 0.5−0.8 mm, elliptic to widely elliptic at maturity, white to dark brown at maturity, glabrous, smooth; lemma slightly curved at the apex; palea rounded at the base; rachilla appendages ca. 0.3 mm long, winged, membranous and hyaline in the young anthecium, fleshy and light brown at maturity, ca. 1/4 the length of the anthecium, apex rounded, base as wide as the apex throughout the entire development of the anthecium, base of the palea not covered by the wings, emerging from a very short curved stipe (rachilla), inconspicuous (<0.1 mm long); lodicules 2, ca. 0.2 mm long; stamens 3, filaments white, anthers yellow; stigmas 2, vinaceous. Caryopsis 1.2 × 0.6 mm, elliptic, whitish; hilum linear, ca. 1/2 the length of the caryopsis.

Distribution and habitat:— Ichnanthus piresii is only known from the municipality of Carolina, Maranhão, Brazil ( Fig. 5 ). It occurs in clefts in the sandstone walls of the mountains of the Chapada das Mesas, in partially to completely shaded and sandy areas ( Fig. 4F,G ).

Notes:— This species has been considered as a synonym of Ichnanthus hoffmannseggii since the taxonomic revision of Stieber (1982), but we are reestablishing it because it differs from I. hoffmannseggii in habit, inflorescence, and size and shape of the upper anthecium and winged appendages ( Table 2). Ichnanthus piresii was referred to as I. aff. hoffmannseggii in our previous phylogenetic study ( Silva et al. 2015) because we were unaware of its identity at that time.

Representative specimens examined:— BRAZIL. Maranhão: Carolina, rodovia MA-010, Serra Portal da Chapada , 29 January 2012, Silva et al. 815 ( HUEFS!, RB!) ; ibid., 19 May 2012, Silva et al. 893 ( CEN!, F!, HUEFS!, ICN!, K!, MG!, MO!, RB!, SP!, US!).

Figures

FIGURE 1. Ichnanthus hoffmannseggii (Silva et al. 1163, by Michella D.R. Teixeira). A. Habit; B. Segment of a primary branch with lower levels of branching; C. Spikelet, lateral view; D. Young anthecium, ventral view; E. Mature anthecium, ventral view; F. Mature anthecium, lateral view; G. Mature anthecium, dorsal view.

FIGURE 2. Ichnanthus oplismenoides (Silva et al. 891, by Juliana G. Freitas). A. Habit; B. Segment of a primary branch with lower levels of branching (without spikelets); C. Spikelet, lateral view; D. Young anthecium, ventral view; E. Mature anthecium, ventral view; F. Mature anthecium, lateral view; G. Mature anthecium, dorsal view.

FIGURE 3. Ichnanthus piresii (Silva et al. 893, by Juliana G. Freitas). A. Habit; B. Segment of a primary branch with lower levels of branching (without spikelets); C. Spikelet, lateral view; D. Young anthecium, ventral view; E. Mature anthecium, ventral view; F.Mature anthecium, lateral view; G. Mature anthecium, dorsal view.

FIGURE 4. Ichnanthus hoffmannseggii (Silva et al. 1163; Santarém, Pará, Brazil). A. Habit; B. Inflorescence. Ichnanthus oplismenoides (Silva et al. 891; Carolina, Maranhão, Brazil). C,D. Habit; E. Inflorescence. Ichnanthus piresii (Silva et al. 893; Carolina, Maranhão, Brazil). F,G. Habit; H. Inflorescence. (Photos by C. Silva)

FIGURE 5. Distributions of Ichnanthus hoffmannseggii, I. oplismenoides, and I. piresii.

FIGURE 7. SEM images of the upper anthecium.A–F. Ichnanthus hoffmannseggii (Taylor et al. 1236). A.Young anthecium, ventral view; B. Mature anthecium, ventral view; C. Mature anthecium, lateral view; D. Mature anthecium, dorsal view; E. Mature anthecium, apex, ventral view; F. Mature anthecium, apex, lateral view. G–L. Ichnanthus oplismenoides (Silva et al. 891). G. Young anthecium, ventral view; H. Mature anthecium, ventral view; I. Mature anthecium, lateral view; J. Mature anthecium, dorsal view; K. Mature anthecium, apex, ventral view; L. Mature anthecium, apex, lateral view. M–R. Ichnanthus piresii (Silva et al. 893). M. Young anthecium, ventral view; N. Mature anthecium, ventral view; O. Mature anthecium, lateral view; P. Mature anthecium, dorsal view; Q. Mature anthecium, apex, ventral view; R. Mature anthecium, apex, lateral view. (Scale bars = 500 μm [A–F]; 500 μm [G–L]; 200 μm [M–R]. L = lemma; P = palea; R = rachilla; ST = stomata; W = winged appendages)