*Sipanea*, in having terminal multiflorous inflorescences, distylous flowers (although a few species of *Sipanea* remain to be confirmed to be distylous), corolla lobes contorted to the left, and capsular fruits. *Dendrosipanea* differs from *Sipanea* by being a shrub or treelet with septicidal capsules, while species of *Sipanea* are prostrate, decumbent or erect herbs or subshrubs (in a few species the stem may be woody at base) with loculicidal capsules. In the molecular phylogenetic analysis of Delprete and Cortés B. (2004) *Dendrosipanea* was retrieved at a basal position on the tree, suggesting that the woody habit is ancestral within the tribe, while *Sipanea* was found in a distant clade.

**Geographic distribution and ecology:** A genus of three species, distributed in Amazonian Colombia, Venezuela and Brazil, growing on white-sand savannas, sometimes seasonally inundated.

Key to *Dendrosipanea* species

1. Leaf blades mostly round or broadly obtuse (rarely acute) at apex, primary and secondary veins impressed above and prominent below; inflorescence shortly branched, 3-7 cm long, including peduncle and flowers (corollas 17-24 mm long; tube 9-14 mm long, 2-2.2 mm wide at base, 3-4 mm wide at mouth); lobes ovate, obtuse at apex ...2. D. revoluta

1. Leaf blades obtuse to acute or acuminate at apex, primary and secondary veins not impressed above and slightly prominent below; inflorescences long-branched, 6-16 cm long, including peduncle and flowers; lobes ovate to narrowly ovate or oblong to ovate-oblong, acute or obtuse to acute at apex ................................................................................2

2. Corollas 19-22.5 mm long; tubes 11-14 mm long, 1.6-1.9 mm wide at base, 2-2.8 mm wide at mouth; lobes ovate to narrowly ovate, 7.5-8.5 x 4.5-4.8 mm, acute at apex ............ 1. D. prancei

2. Corollas 27-30 mm long; tubes 16-20 mm long, 2-2.5 mm wide at base, 3.5-4.5 mm wide at mouth; lobes oblong to ovate-oblong, 12-16 x 5-7.5 mm, obtuse to acute at apex .. .................................................................................. 3. D. spigelioides

2-1. *Dendrosipanea prancei* Delprete, Phytotaxa 382(1): 127, fig. 1. 2018. (Figures 4C-D, 10-11).

Type: BRAZIL. Amazonas: Basin of Rio Negro, Rio Uneiuxi [ca. 0°32'S, 65°3'W, ca. 30 m alt.], 5 km above mouth, periodically flooded savanna on

sand, 8 Nov. 1971 (fl), G.T. Prance, P.J.M. Maas, D.B. Woolcott, O.P. Monteiro & J.F. Ramos 16199 (holotype INPA [Acc. No. 34418]; isotypes F [Acc. No. 1841650], MO [Acc. No. 2728504], NY, R [Acc. No. 180261], S [Acc. No. 10-28034], U [U0016880], US [Acc. No. 2868603]). (Figures 4C-D, 10-11).

*Shrubs or treelets*, 1-1.3 m tall; leafy branchlets terete to slightly tetragonal, glabrous, red, distal nodes slightly resinous, resin produced by the stipules. *Stipules* free or basally adnate to petioles, ovate-triangular to broadly triangular, 2-3 x 2-4.5 mm, bifid at upper third to half the length, lobes narrowly triangular to acuminate and reduced to two teeth in older stipules, glabrous outside, glabrous at distal portion and basal 2/3 covered by dense colleters intermixed with wavy sericeous hairs inside; margins glabrous or sometimes hispidulous-ciliolate. *Leaves* subsessile to short-petiolate; petioles 2-5 mm long, glabrous throughout; blades elliptic, oblong-elliptic to narrowly oblong-elliptic, 4-10 x 1.3-3 cm, acute to decurrent at base, acute at apex, subcoriaceous, dark green and shiny above, pale green below when fresh, drying brown above and pale brown below; glabrous throughout, margins scarcely or not revolute, primary and secondary veins not impressed above and slightly prominent below; secondary veins 6-11 each side of mid-rib. *Inflorescences* cymose, frondose, long-branched, 6-14 cm tall, (including peduncle and flowers), 3-8 cm broad, terminal branches scorpiod, expanding at fruit maturity; rachis glabrous; peduncle 2.5-5.5 cm long; secondary branches subtended by a pair of leaf-like bracts; these similar in shape to regular cauline leaves, (2.5-)3.5-7 x 0.9-2.2 cm; each flower with 1 bracteole inserted on the opposite side of the branch; bracteoles sessile, oblong-lanceolate, 1-1.5 x 0.5-0.8 mm, acuminate at apex, often with 1-2 tiny teeth on each side, glabrous, sparsely ciliolate. *Flowers* with pedicels 3-8 mm long, glabrous. *Hypanthium* turbinate to narrowly obovoid, 1.5-1.7 x 1.2-1.5 mm, glabrous. *Calyx* lobes subequal to unequal, oblong to narrowly oblong, 1.5-5 x 0.4-0.7 mm, round to acute at apex, glabrous or sparsely strigillose outside, sericeous at base inside, margins glabrous, with 2-4, linear, 0.3-0.4 mm long, colleters at each lobe sinus. *Corolla* 19-22.5 mm long; tube narrowly infundibuliform, 11-14 mm long, 1.6-1.9 mm wide at base, 2-2.8 mm wide at mouth, glabrous or with a few appressed-strigulose hairs

**Figure 10.** *Dendrosipanea prancei*. A. Branch with terminal, frondose inflorescence. B-C. Stipule shapes present on the same branch. D. Detail of inflorescence branch, with secondary branches subtended by two leaf-like bracts (i.e., pherophylls). E. Flower in anthesis. F. Dissected long-styled flower, with stamens inserted at basal portion of corolla tube. A-D: drawn from *Prance et al. 16199* (F, S); E-F: drawn from *Prance et al. 15472* (US). Illustration by Piero Delprete (Reproduced from fig. 1 of Delprete (2018) with permission of copyright holder Magnolia Press).
<figure><img src="image_14.png" /><figcaption>Figure 10.</figcaption></figure>
outside, villous and densely yellow-pubescent at mouth inside; lobes ovate to narrowly ovate, 7.5-8.5 x 4.5-4.8 mm, sometimes slightly asymmetrical and of slightly different width on the same corolla, acute at apex, glabrous outside, densely papillose inside, margins glabrous. *Short-styled flowers* unknown. *Long-styled flowers* (Prance et al. 15472): stamens included, inserted at 2.7 mm from the base of corolla; filaments 3.7 mm long; anthers linear-oblong, 3.5-3.7 x 0.2-0.3 mm, acute at both ends; style barely exserted (only branch tips beyond corolla mouth), 14-15 mm long, glabrous, style branches oblong, 2 mm long. *Infructescences, capsules and seeds* unknown.

**Notes:** *Dendrosipanea prancei* Delprete is similar to *D. spigelioides* in having leaf blades acute at apex (vs. round, broadly obtuse or rarely acute) at apex in *D. revoluta*), with primary and secondary veins not impressed

above and slightly prominent below (vs. impressed above and prominent below in *D. revoluta* Steyerm.); the former differs from the latter by the long-branched inflorescence, 6-16 cm long including peduncle and flowers (vs. shortly branched, 3-7 cm long including peduncle and flowers in *D. spigelioides*), corollas 19-22.5 mm long, tubes 11-14 mm long, 1.6-1.9 mm wide at base, 2-2.8 mm wide at mouth, and lobes ovate to narrowly ovate, 7.5-8.5 x 4.5-4.8 mm, acute at apex (vs. corollas 27-30 mm long, tubes 16-20 mm long, 2-2.5 mm wide at base, 3.5-4.5 mm wide at mouth, lobes oblong to ovate-oblong, 12-16 x 5-7.5 mm, obtuse to acute at apex in *D. spigelioides*).

**Distribution and ecology:** Only known from the Rio Uneiuxi Basin, affluent of the Rio Negro, state of Amazonas, Brazil, and the Department of Guainía, Amazonian Colombia (Figure 11); on white-sand savan-

<figure><img src="image_15.png" /><figcaption>**Figure 11.** Distribution of *Dendrosipanea prancei* (circles), *D. revoluta* (squares), and *D. spigelioides* (triangles).</figcaption></figure>
nas, seasonally flooded by black-water rivers, at 30–110 m altitude.

**Phenology:** Flowering specimens were collected in October and November.

**Suggested conservation status:** Vulnerable (VU). This species is quite rare and is only known from two collections made in 1971 near the Rio Uneiuxi, an affluent of the Rio Negro, state of Amazonas, Brazil, and one recent collection from the Municipality of Inirida, Department of Guainía, Amazonian Colombia. The two localities are quite remote and very distant from each other.

The labels of the two 1971 collections near the Rio Uneiuxi (ca. 0°32'S, 65°3'W, ca. 30 m alt.) do not report population sizes, nor the rarity of the species. Personal observations of the vegetation surrounding the Rio Uneiuxi were made in February 2018. The region of the Rio Uneiuxi was inhabited by sparse indigenous communities, and the vegetation was fairly well preserved. During the dry season, the white-sand savannas of this region are subjected to frequent extensive fires (pers. obs.), often started by the local communities. After extensive searches made in several local savannas in February 2018, this species was not found. It is possible that, as it flowers in October–November, the species was not evident in February.

The single collection in Colombia was made in 2005 near the Caño Pato, 3°8'14"N, 67°47'34"W, an affluent of the Río Inirida. It is an isolate locality, nevertheless, ca. 50 km from a main road. The specimen label reports that the species was growing in a seasonally flooded forest undercanopy, but did not report the size of the population. It is expected that the locality has the potential of being altered, as it is not far from a main road. Therefore, this species is Vulnerable (VU) following IUCN criteria (IUCN 2012, 2019).

**Additional specimens examined (paratypes):**

**COLOMBIA. Guainía:** Mun. Inirida, km 50 de la carretera Huesito (Río Inirida – Puerto Caribe – Río Guainia), Sector de Caño Pato, bosque abierto con sotobosque denso sobre llanura alluvial, 3°8'14"N, 67°47'34"W, 110 m, 3 Oct. 2005 (fl, imm fr), *D. Cárdenas & A.L. Bermudez* 16831 (COAH, NY).

**BRAZIL. Amazonas:** Basin of Rio Negro, Rio Uneiuxi [ca. 0°32'S, 65°3'W, ca. 30 m alt.], 100–200 km above mouth, black-water flooded savanna forest on sand, shrub 1.3 m tall, corolla tube green, lobes white with yellow hairs at center, 21 Oct. 1971 (fl), *G.T. Prance et al.* 15472 (INPA, MG, NY, U, US).

2-2. **Dendrosipanea revoluta** Steyerm., Mem. New York Bot. Gard. 10: 196, fig. 71. 1964. (Figures 11–12).

Type: VENEZUELA. Amazonas: Savanna on right bank of Río Pacimoni, 50 km above mouth, frequent, 100–140 m, 7 Feb. 1954 (fl), *B. Maguire, J.J. Wurdack & G.S. Bunting* 37592 (holotype NY [00131229]; isotypes P [P00729464], RB [Acc. No. RB 115187, barcode 00543545], US [00137732], VEN [Acc. No. 49433]).

*Shrubs* 0.5–4 m tall; leafy branchlets terete to slightly tetragonal, hirtellous, distal nodes slightly resinous. *Stipules* free at base, ovate-triangular to broadly triangular (at wider nodes), 2–4 mm × 2–5.5 mm, acuminate or bifid to half the length, irregularly strigose outside, sparsely pubescent at distal portion and basal 2/3 covered by dense colleters intermixed with straight serious hairs inside; margins hispidulous ciliate.

*Leaves* subsessile to short-petiolate; petioles 2–9 mm long, glabrous or sparsely hispidulous above, hirtellous below; blades narrowly oblong or oblanceolate-spathulate, 3–10 × 1–3.5 cm, cuneate at base, round to broadly obtuse or rarely acute at apex, subcoriaceous, dark green and shiny above, pale green below when fresh, drying pale brown above and pale olive-green below; glabrous or sparsely asperous-hirtellous above, densely pubescent below, margin minutely revolute, primary and secondary veins impressed above and prominent below, veins glabrous above and densely strigillose below; secondary veins 7–9 each side of midrib. *Inflorescences* cymose, often frondose, shortly branched, 3–7 cm long (including peduncle and flowers, 2–3 cm broad, expanding at fruit maturity; rachis sparsely to densely strigillose; peduncle 2–3 cm long; secondary branches often subtended by a pair of leaf-like bracts; these, (when present) similar in shape of regular cauline leaves, 2–3.5 × 0.8–1.5 cm; flowers subtended by 1 bracteole, these sessile, oblong-lanceolate, 2–3.5 × 0.4–0.8 mm, acute at apex, sparsely strigillose. *Flowers* with pedicels 1–5 mm long, sparsely to densely strigillose or hispidulous. *Hypanthium* turbinate, 2–3 × 1.5–2.5 mm, strigillose. *Calyx* lobes subequal to unequal, narrowly oblong-lanceolate, oblanceolate to spathulate-oblong, 2–9 × 0.7–3.5 mm, sparsely to moderately hispidulous outside, glabrous or hispidulous on distal portion and with a row of trichomes at base inside, margins hispidulous-ciliate, with 1–2, linear, 0.4–0.5 mm long, colleters at each lobe sinus. *Corolla* 17–24 mm long; tube narrowly infundibuliform, 9–14 mm long, 2–2.2 mm wide at base, 3–4 mm wide at mouth, sparsely strigulose or hirtellous outside, basal portion glabrous or papillose, the medio-distal portion villous and densely
<figure><img src="image_16.png" /><figcaption>**Figure 12.** *Dendrosipanea revoluta*. A. Branch with inflorescences. B–C. Stipule variation on the same branch. B. Entire stipule. C. Bifid stipule. D. Dissected short-styled flower. E. Dissected long-styled flower. A–C, E: drawn from Wurdack & Adderley 42972 (US); D: drawn from Maguire et al. 36672 (US). Illustration by Piero Delprete.</figcaption></figure>
yellow-pubescent at mouth inside; lobes ovate, 8–11 × 4–6 mm, obtuse at apex, sparsely strigulose or glabrate outside, glabrous inside, margins basally ciliate. *Short-styled flowers* (Maguire et al. 36672): stamens included, inserted at 5 mm below corolla mouth; filaments 0.7–0.8 mm long; anthers linear, 2.2–3.1 × 0.2–0.3 mm, acuminate at both ends; style included, 2.5 mm long, glabrous, style branches oblong, 1 mm long. *Long-styled flowers* (Wurdack & Adderley 42972): stamens included, inserted at 7 mm below corolla mouth; filaments 0.7–0.8 mm long; anthers linear, 2.5–2.6 × 0.2–0.3 mm, acuminate at both ends; style barely exerted (only branch tips beyond corolla mouth), 8.5–9 mm long, style branches oblong, 0.5–0.6 mm long. *Infructescence* a compound dichasium, 4.5–6.5 cm long, terminal branches scorpiod, 2.5–3.5 cm long. *Capsules* oblong-turbinate, (5-)7–10 × 3–5 mm, acute at base, round at apex, disk exceeding the calyx, dark brown, strigillose. *Seeds* irregularly polyhedral, 1.2–1.8 × 0.7–1 mm, yellowish-brown to pale brown.

**Distribution and ecology:** Occurring in the Venezuelan state of Amazonas, in the basins of the Río Atabapo and Río Pacimoni, and one collection in the Municipality of Barcelos, state of Amazonas, Brazil (Figure 11); growing in white-sand savannas seasonally flooded during the rainy season near black water rivers, at 100–200 m altitude. It is probable that this species might occur also in adjacent Colombia (Guainía Department).

**Phenology:** Flowering specimens were collected in February, March, June, October and November; with flowers and immature fruits in February and March; and with mature fruits in May.

**Suggested conservation status:** Near Threatened (NT). Known by numerous collections in the basins of Río Atabapo and Río Pacimoni, Venezuelan state of Amazonas, and one collection in the distant Municipality of Barcelos, in the Brazilian state of Amazonas. In Venezuela it was collected in 1953, 1957, 1959, 1979, 1980, 1981, 1989, 1995, and 1996. While the single collection from the surroundings of Barcelos, Brazil, was made in 1985. All of these localities are quite remote, and apparently without any immediate threat. However, Amazonian white-sand savannas are often subjects of frequent fires during the dry season. The population sizes are unknown. Using the coordinates of all collection localities, the EOO of this species is of ca. 40.000 km². Two localities in the Casiquiare Department, Venezuela, are now included within the Alto Orinoco-Casiquiare Biosphere Reserve. Therefore, taking into account that the localities of occurrence might be subjects of frequent

fires, and that the last collection of this species was made in 1996, it is classified as Near Threatened (NT) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA: Amazonas: shrubby white-sand savanna just W of Santa Cruz de Atabapo, 3°13'N, 67°23'W, 100 m, 30 Nov. 1995 (fl), P.E. Berry et al. 5779 (MO, VEN); lower Río Guasacavi (black-water river), 0–1 km W of mouth of Río Temi, 3°14'0"N, 67°23'38"W, 100 m, 6 Mar. 1996 (fl), P.E. Berry et al. 5996A (MO, VEN); Depto. Atabapo, SE bank of the middle part of Caño Yagua at Cucurital de Yagua, 3°36'N, 66°34'W, 120 m, 8 May 1979 (fr), G. Davidse et al. 17395 (MO, NY, US, VEN); Depto. Atabapo, pequeña sabana a ca. 5 km S del caserío Puruname, en el Río Puruname bajo, 3°22'N, 66°30'W, 100 m, 8 Mar. 1980 (imm fr), O. Huber 5075 (NY, VEN); Depto. Río Negro, extensa sabana en la margen derecha del bajo Río Pasimoni, 1°35'N, 66°33'W, 125 m, 8 Feb. 1981 (fl, imm fr), O. Huber & E. Medina 5850 (NY, VEN); Depto. Casiquiare, sabanas ubicadas al W de la Serra Cariche, ca. la altura de Cariche en la ribera izquierda del Río Orinoco, 2°57'N, 66°23'W, 120 m, 17 Feb. 1981 (imm fr), O. Huber 6098 (NY, VEN); Depto. Casiquiare, Ríos Pacimoni-Yatua, savanna on R bank of Río Pacimoni, 60 km above mouth, 29 Nov. 1953 (fl), B. Maguire et al. 36672 (IAN, NY, US, VEN, paratype); Sabana Pacimoni, R bank of Río Pacimoni, 50 km above mouth, 100 m, 2 Oct. 1957 (fl), B. Maguire et al. 41665 (NY, P, U, US, paratype); Depto. Atabapo, Laja Cúcuta, Río Atacavi, Pica 1, Frente No. 3, individuo de 80 cm, dominante en sabana amazónica arbustiva, flores blancas, 3°12'N, 67°24'W, 100 m, Nov. 1989 (fl, imm fr), J. Velazco 1056 (NY, PORT. US); Río Atabapo, Sabana Manacal, 15 km above Guarinumo, 125 m, 12 Jun. 1959 (fl), J.J. Wurdack & L.S. Adderley 42972 (IAN, NY, US, VEN, paratype).

**BRAZIL:** Amazonas: Mun. Barcelos, Rio Aracá, a 13 horas de barco de Barcelos, beira do Rio, 0°0'-20'S, 63°0'-30'W, 28 Jul. 1985 (fl), J.A. Silva et al. 386 (INPA, NY [2 sheets]).

2-3. *Dendrosipanea spigelioides* Ducke, Arch. Inst. Biol. Veg. 2(1): 70, figs. 8-9. 1935. (Figures 11, 13).

Emend. Delprete & Steyerm. in Steyermark et al., Fl. Venez. Guayana 8: 571. 2004.

**Type:** BRAZIL. Amazonas: Rio Curicuriary [now Rio Curicuriari; ca. 0°12'S, 66°47'W, ca. 60 m alt.], affluent of the Upper Rio Negro, ["abaixo das cachoeiras, margem inundavel"], 19 Oct. 1932
(fl, fr), *A. Ducke s.n.* (RB 24426) (holotype RB [mounted on 4 sheets, 00706257 (Sheet No. 1, fl), 00706256 (Sheet No. 2, fl), 00706258 (Sheet No. 3, fl), 00543546 (Sheet No. 4, fr)]; isotypes F [Acc. No. 779985], IAN [Acc. No. 50581], NY [2 sheets, 1731845 and 131230], INPA [Acc. No. 246872], P [P00729463], S [Acc. No. S05-1746], U [U0016882], US [00137733]]).

(=) *Dendrosipanea wurdackii* Steyerm., Mem. New York Bot. Gard. 10: 198, fig. 72a-g. 1964

Type: VENEZUELA. Amazonas: Río Guainía, occasional in Sabana El Venado, on left bank above of Caño Pimichín [2°74'N, 67°56'W], above Pimichín, 140 m, 14 Apr. 1953 (fl), *B. Maguire & J.J. Wurdack* 35558 (holotype NY [barcode 00131231]; isotype VEN [Acc. No. 49443]).

*Shrubs* 2-4 m tall; leafy branchlets terete to slightly tetragonal, glabrous or sparsely strigillose, distal nodes lightly resinous. *Stipules* free or basally adnate to petioles, ovate-triangular to broadly triangular, 2.5-3 x 2-4.5 mm, acuminate or bifid to half the length, glabrous or sparsely strigulose outside, sparsely pubescent at distal portion and basal 2/3 covered by dense colleters intermixed with straight sericeous hairs inside; margins hispidulous ciliate. *Leaves* subsessile to short-petiolate; petioles 3-10 mm long, glabrous above, glabrous or sparsely strigillose below; blades elliptic-lanceolate, oblong-lanceolate, lanceolate-ovate, narrowly oblong-lanceolate, 4-13 x (1-)2.5-4.5 cm, acute at base, acute or acuminate at apex, subcoriaceous, dark green and shiny above, pale green below when fresh, drying brown above and olive-green below; glabrous above, glabrous and with veins glabrous or sparsely strigulose below, margins scarcely or not revolute, primary and secondary veins not impressed above and slightly prominent below; secondary veins (6-)7-12 each side of midrib. *Inflorescence* cymose, frondose, long-branched, 6-16 cm long (including peduncle and flowers), 3-13 cm broad, terminal branches scorpiod, expanding at fruit maturity; rachis glabrous or sparsely strigillose; peduncle 2-5 cm long; basal secondary branches subtended by a pair of leaf-like bracts; these similar in shape of regular caul-

ine leaves, 3-5.5 x 1-2 cm; each flower with 1 bracteole inserted on the opposite side of the branch; bracteoles sessile, oblong-lanceolate, 1-2 x 0.3-0.7 mm, acuminate at apex, often with 1-2 tiny teeth on each side, sparsely strigillose, sparsely ciliolate. *Flowers* with pedicels 3-5 mm long, glabrous or sparsely strigillose. *Hypanthium* turbinate, 2-3 x 1.7-2 mm, glabrous or sparsely strigillose. *Calyx* lobes subequal or unequal, deltoid to narrowly triangular, 1-3.5 x 0.9-1.2 mm, glabrous or sparsely strigillose outside, short-sericeous inside, margins hispidulous-ciliate, with 2-4, linear, 0.3-0.5 mm long, colleters at each lobe sinus. *Corolla* 27-30 mm long; tube narrowly infundibuliform, 16-20 mm long, 2-2.5 mm wide at base, 3.5-4.5 mm wide at mouth, glabrous or very sparsely appressed-strigulose outside, basal portion glabrous or papillose, the medio-distal portion vigorous and densely yellow-pubescent at mouth inside; lobes oblong to ovate-oblong, 12-16 x 5-7.5 mm, often slightly asymmetrical and of different width on the same corolla, obtuse to acute at apex, glabrous outside, glabrous inside, margins glabrous. *Short-styled flowers* (*Maguire & Wurdack* 35558, type of *D. wurdackii* Steyerm., corolla tube 16-20 mm long, lobes 14-16 x 4-5 mm): Stamens included, inserted 6 mm below corolla mouth; filaments 2.5-3.5 mm long; anthers linear-oblong, 2.8-3.1 x 0.2-0.3 mm, acute at both ends; style included, 3-3.5 mm long, glabrous, style branches oblong-lanceolate, 1-1.3 mm long. *Long-styled flowers* (*Ducke* 356, corolla tube 18 mm long): Stamens included, inserted 5-6 mm from the base of corolla tube; filaments 4-4.5 mm long; anthers linear-oblong, ca. 4 x 0.2-0.3 mm, acuminate at both ends; style barely exserted (only branch tips exserted beyond corolla mouth), 16.5-20.3 mm long, glabrous, style branches oblong-elliptic, 2-3 mm long, round at apex. *Infructescences* a compound dichasium, 6-16 cm long, terminal branches scorpiod, 6-9 cm long. *Capsules* turbinate to oblong-turbinate, (6-)8-11.5 x (4-)5-5.7 mm, acute at base, round at apex, disk exceeding the calyx, brown, strigillose. *Seeds* irregularly polyhedral or irregularly-shaped, uncinated, dorso-ventrally compressed to tronco-conical, 1-2 x 0.6-1.3 mm, pale brown.

**Taxonomic notes:** Ducke (1935), along with the description of *Dendrosipanea spigelioides* Ducke, indi-

**Figure 13.** *Dendrosipanea spigelioides*. *A*. Branch with inflorescence and flowers in anthesis. *B*. Dissected corolla of long-styled flower, with exserted style. *C*. Dissected corolla of short-styled flower, with included style. *D*. Longitudinal section of ovary, disk and calyx, showing fleshy placental extension adnate along the entire length of the septum, and peltate ovules. *E*. Septicidal capsule, half portion showing septum and long disk. *F-G*. Stipule variation on the same branch. *F*. Entire stipule. *G*. Bifid stipule. *A, D, F, G*: drawn from *Ducke 356* (F); *B*: drawn from *Ducke 356* (S); *C*: drawn from *Maguire & Wurdack 35558* (NY); *E*: drawn from *Ducke s.n* (RB 24426) (RB). Illustration by Piero Delprete.
<figure><img src="image_17.png" /><figcaption>Figure 13.</figcaption></figure>
cated his own unnumbered collection with the RB accession No. 24426, without indicating the herbarium of deposit. At the time of the publication he worked at the Botanical Garden of Rio de Janeiro (RB) and he did not use his own collection numbers, but rather the RB accession numbers, as it was commonly done in those days at that institution. This gathering was subsequently distributed, and duplicate specimens with RB accession No. 24426 are now at F, INPA, NY, P, RB, S and US. Steyermark (1964: 199) cited the type of this name as "Brazil. Amazonas: Rio Curicuriary affluentis Rio Negro superioris; ad ripas inundatas; 19 October 1932, *A. Ducke 24426* (isotype NY). - The isotype specimen at NY shows minor variations from the description of the species. According to the material examined, this species has a larger, more laxly branched inflorescence than either of the other two species of the genus." Article 9.10 of the *Code* (Turland et al. 2018) states that "The use of a term defined in the *Code* (Art. 9.1, 9.3 and 9.5-9.9) as denoting a type, in a sense other than that in which it is so defined, is treated as an error to be corrected (for example, the use of the term lectotype to denote what is in fact a neotype)." Because Ducke in the protologue of *D. spigelioides* cited "leg. A. Ducke, 19-10-1932, H.J.B.R. n° 24.426," Steyermark assumed that Ducke cited a sole specimen at RB, which, according to him, was the holotype. With this assumption, Steyermark (1964) by citing "*A. Ducke 24426* (isotype NY)" was correct in using the term isotype as he assumed that it was a duplicate of the holotype at RB, hence it cannot be corrected. At RB are present four sheets with accession No. 24426; these sheets are kept together in the same folder, and only one of them (sheet No. 1, with barcode 00706257), has the original label handwritten by Ducke (the other three sheets have no label); therefore, according to Art. 8.3 of the *Code* (Turland et al. 2018), they are a single specimen with multiple preparations. To each sheet has been assigned a different barcode. This specimen, mounted on four sheets, is the holotype of *Dendrosipanea spigelioides*.

Seyermark (1964) separated *Dendrosipanea wurdackii* from *D. spigelioides* based on the shape and size of the calyx lobes and details of the pubescence of both vegetative and reproductive structures. However, according to Delprete and Steyermark (2004a, p. 571), the variation in these characters is continuous, and these taxa are here confirmed to be synonymous.

**Distribution and ecology:** It has been collected in Amazonian Venezuela on white-sand savannas, along streams, and adjacent seasonally flooded forests, usually

in hummocks locally called "tatucos", and Brazil, near the Rio Curicuriari, Upper Rio Negro Basin (Figure 11); growing in seasonally flooded white-sand savannas, and seasonally flooded low vegetation or forests at river margins, at 90-200 m altitude.

**Phenology:** Flowering specimens were collected in February, April, June, July, October and November; with flowers and mature fruits in February.

**Suggested conservation status:** Vulnerable (VU). A rare species known by three collections in Amazonian Venezuela, and three collections in contiguous Amazonian Brazil, always on white-sand savannas. In Brazil it was collected in the state of Amazonas in 1932 (type of *D. spigelioides*) and 1936 (paratype of *D. spigelioides*) near the Rio Curicuriari, ca. 0°12'S, 66°47'W, and in 1987 near the Rio Içana, ca. 0°23'N, 67°33'W, in the Municipality of São Gabriel da Cachoeira.

In Venezuela it was collected in the state of Amazonas, in 1953 (type of *D. wurdackii*) in the savanna El Venado, basin of the Río Guainía, near Caño Pimichín, ca. 2°74'N, 67°56'W; in 1982 in a savanna at ca. 15-20 km S of Tama-Tamá, ca. 2°31'N, 66°16'W; and in 1992, in a savanna near the town of San Juan de Ucata, 4°20'25"N, 67°44'12"W.

None of the specimen label report the size of the populations of this species, and no recent observations has been made in the collection localities to assess their state of preservation. During the dry season, Amazonian white-sand savannas are subject to frequent fires, often started by the local communities. Therefore, this species is classified as Vulnerable (VU) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** **VENEZUELA:** Amazonas: Edge of lagoon below Raudal de Ucata, above town of San Juan de Ucata, 4°20'25"N, 67°44'12"W, 120-150 m, 20 Jun. 1992 (fl), *P.E. Berry et al. 5162* (MO, VEN); Dto. Río Negro, ca. 15-20 km S of Tama-Tamá, transecto desde sabana de arena blanca con drenaje laterítico hasta bosque medio denso inundado, con tatucos, 2°31'N, 66°16'W, 90 m, 5 Nov. 1982 (fl, young fr), *F.M. Guánchez 1996* (MO, TFAV).

**BRAZIL:** Amazonas: Rio Curicuriari [ca. 0°12'S, 66°47'W, ca. 60 m alt.], above waterfalls, flooded river margin, 23 Feb. 1936 (fl, fr), *A. Ducke 356* (F, MO, NY, R, US); Mun. São Gabriel da Cachoeira, Rio Içana, Jauacanã, arbusto, na mata de igapó, 0°23'N, 67°33'N, 2 Nov. 1987 (fl), *C. Farney & W. Rodrigues 1861* (MO, NY).
### 3. LIMNOSIPANEA

*Limnosipanea* Hook.f., Ic. pl. 11: 38, pl. 1050. 1868

K.Schum. in Mart. et al., Fl. Bras. 6(6): 252-256, tab. 123. 1889; Steyerm. in Mem. New York Bot. Gard. 17: 282-284. 1967; Steyerm. in Lasser and Steyermark, Fl. Venezuela 9(1): 384-390, fig. 61. 1974; Delprete and Steyerm. in Steyermark et al., Fl. Venez. Guayana 8: 638-639, figs. 500-501. 2004; Delprete in J.A. Rizzo, Fl. Goiás Tocantins 40(2): 619-629, fig. 54. 2010.

Type: lectotype (designated by Steyermark (1967: 282), see notes below): *Limnosipanea spruceana* Hook.f.

*Herbs*, terrestrial, semi-aquatic (amphibious) or aquatic (in *L. spruceana*); erect, prostrate or decumbent; ephemeral to short-lived (2-4 months life span) when terrestrial or semi-aquatic, or perennial when growing in permanently submersed areas; aerial portion (i.e., not submerged) of stems sparsely pubescent. *Stipules* reduced to a line between the petioles, sometimes obsolete, persistent, withering on the stem (no abscission layer is formed); with 1 to 3 colleters inserted just below the margin, on the internal side, with the tips exerted beyond the stipule margin, or, when stipules obsoletes, 1 or 2 colleters inserted at the sinuses between leaf bases; in submersed stems, stipules and colleters absent. *Leaves* opposite or whorled, sessile, membranaceous or chartaceous, domatia absent; homophyllous with 2-3 per node in terrestrial plants; heterophyllous in aquatic or semi-aquatic plants, submerged leaves (when present) 3-6(-8) per node, linear to acicular, aerial leaves 2-5 per node per node, linear to lanceolate. *Inflorescence* terminal, cymose, few-many-flowered, frondose or not, or uniflorous. *Flowers* homostylous, (4-5)5(-6)-merous. *Hypanthium* narrowly ellipsoid to narrowly obovoid, densely hispid. *Calyx* persistent; tube extremely reduced or absent; lobes free or nearly so, subequal, oblong, lanceolate to linear, aristate at apex (arista caducous) or not, with 1 minute colleter in each lobe sinus. *Calycophylls* absent. *Corolla* hypocrateriform, white, cream-white, pinkish-white, pink to lilac; tube externally glabrous or strigose, internally vilious, with a pubescent ring at orifice; lobes left-contorted, ovate or oblong-ovate, margin entire or undulate, obtuse or round at apex. *Stamens* exerted; filaments attached near the mouth of the corolla tube, glabrous; anthers yellow, narrowly elliptic or oblong, round at base and at apex, dorsifixed near the middle. *Pollen* 3(4)-colporate, exine shallowly foveolate or shallowly reticulate-foveolate, without perforations or finely perforate. *Ovary* with a stalked placenta; stalk inserted at the middle of the sep-

tum, terminating with a hemi-ellipsoid extension; ovules inserted on the placental extension. *Style* exerted, filiform, glabrous; branches oblong or ovate. *Fruits* capsular, dehiscing loculicidally, crustaceous. *Seeds* many per locule, horizontal, small, 3-5-angular; testa foveolate.

**Notes on designation of lectotype species:** Joseph Dalton Hooker (1868: 38. t. 1050) published *Limnosipanea* Hook.f., and described three species in this genus: *L. spruceana* Hook.f., *L. schomburgkii* Hook.f., and *L. palustris* (Seem.) Hook.f., without indicating the type species. Standley (N. American Fl. 32(2): 94. 10 May 1921) indicated *L. spruceana* as the type of the genus, without any explanation. Article 10.7 Ex. 11 of the *Code* (Turland et al. 2018) states that "In considering the typification of *Achyranthes* L. [...] in the *North American Flora*, Paul C. Standley (in J. Wash. Acad. Sci. 5: 72. 1915) selected *A. repens* L. as type stating that "there seems, moreover, no doubt as to the type of the genus *Achyranthes* under the *American Code* of nomenclature", noting that, as a result, "the name *Achyranthes* must be used in a sense other than that in which it has generally been employed in recent years". As a result of this published statement of acceptance of the *American Code*, not only is Standley's selection of *A. repens* superseded by that of *A. aspera* L. by Hitchcock (in Sprague, Nom. Prop. Brit. Bot.: 135. 1929), but types cited in Standley's other publications (e.g. in Britton, N. Amer. Fl. 21: 1-254. 1916-1918) are superseded under Art. 10.5." As Standley followed a clearly mechanical method and accepted the *American Code*, his species lectotypification of *Limnosipanea* should be superseded. Searching subsequent specialized literature, the following type species citation for *Limnosipanea* is that by Steyermark (Mem. New York Bot. Gard. 17(1): 282. 1967), who designated *L. spruceana* as the type of the genus, which is here followed.

**Distribution:** Genus ranging from Panama through Colombia, Venezuela, Guyana, Brazil to Bolivia. The three species of this genus are terrestrial or semi-aquatic (amphibious), commonly found in seasonally inundated fields, at margins of small water courses, and swamps with *Mauritia flexuosa* L.f. palms, or in permanently inundated areas (aquatic habit). Two species of this genus can assume different habit and can be heterophyllous, depending on the habitat where they grow. For example, when *L. spruceana* grows as a partially submerged plant in inundated areas, the submerged leaves are whorled, 3-6(-8) per node, linear to acicular, and the aerial leaves are opposite or whorled, 2-5 per node, lanceolate to ovate. On the other hand, when the same species grows in areas not inundated, the plants are terrestrial and the
leaves are homophyllous and similar to those of the aerial portion of the partially submerged plants.

*Limnosipanea*, and particularly *L. spruceana*, is often overlooked as a genus of Rubiaceae because the stipules are so reduced that they appear to be absent, its semi-aquatic habit and whorled leaves, which are unusual in this family.

Key to *Limnosipanea* species

1. Leaves homophyllous or heterophyllous, 2-6(-8) per node; submerged leaves (when present) 3-6(-8) per node, linear to acicular, without evident secondary veins or sometimes with 1 secondary vein on each side of midrib; aerial leaves 2-6 per node, linear to lanceolate, with 1-2(-3) secondary veins on each side of midrib.................. 3. *L. spruceana*

1. Leaves homophyllous, whorled, 3-4 per node, at basal nodes and opposite and ternate at distal nodes, or leaves opposite throughout the whole plant; leaf blades ovate, narrowly ovate, oblong-ovate, elliptic to lanceolate, without evident secondary veins (rarely with 1-2 secondary veins on each side of midrib) .................................................... 2

2. Plant prostrate or decumbent; leaves 3-4 per node at basal nodes and opposite at distal nodes, or opposite throughout the whole plant; flowers with pedicels (0.5)-1-2.5 mm long; corollas 11.5-16(-18) mm long, pink to lilac, tube 7-9 mm long, glabrous outside, lobes 4.5-8 × 2-4 mm; capsule pedicellate, sparsely hispid........ 1. *L. erythraeoides*

2. Plant erect; leaves commonly opposite throughout the whole plant, or rarely ternate; flowers commonly sessile (rarely short-pedicellate in central flowers); corollas 5.7-10 mm long, white to pinkish-white, tube 3.5-6.5 mm long, strigose outside, lobes 2.2-3.5 × 0.7-0.9 mm; capsule sessile, hispid-setose..................................................2. *L. palustris*

3-1. *Limnosipanea erythraeoides* (Cham.) K.Schum. in Martius & A.G.Eichler, Fl. Bras. 6(6): 253, tab. 123. 1889. (Figures 14-15).

(=) *Sipanea erythraeoides* Cham., Linnaea 9: 242. 1834. Original type: BRAZIL: "Brasilia inter tropicos", s.d., *Sellow s.n.* (B, destroyed). **Neotype (here designated):** BRAZIL. Goiás: Near Caiapônia, "in arenosis humidiusque, Aldeia dos Cayapos" [sandy humid savanna, near the indigenous village of the Cayapos (now the town of Caiapônia, ca. 16°57'S, 51°48'W)], Aug. 1826 (fl), *L. Riedel 409* (BR [barcode 00000824532]; isoneotypes BM [barcode BM000614341], FI [barcode FI018882], P [barcode P00729267], US [Acc. No. 254488]).

*Herb*, terrestrial or semi-aquatic (amphibious), prostate or decumbent, single-stemmed or sparsely branched; main

stems 8-20 cm long; stems sparsely to densely appressed pubescent. *Stipules* reduced to a line between the petioles, sparsely appressed pubescent, with 2 or 3 colleters inserted just below the margin on the internal side, with the tips exserted beyond the margin. *Leaves* homophyllous, 3-4 per node at basal nodes and opposite at distal nodes, or opposite throughout the whole plant, sessile; blades ovate, narrowly ovate, elliptic to lanceolate, 2.5-8(-10) × 0.5-3.5 mm, acute to obtuse at base, acute to obtuse at apex, chartaceous to membranaceous, pale green, drying olive-green, glabrous above and below, only the midvein evident (rarely with 1-2 secondary veins evident on each side). *Inflorescences* 1-3-flowered, or 2-3-chotomous and pauciflorous, 3-15 mm long, frondose, with leaf-like bracts subtending secondary branches; bracteoles subtending each flower elliptic-lanceolate, narrowly lanceolate to linear, 1.5-2.5 × 0.2-0.4 mm, glabrous. *Flowers* pedicellate; pedicels (0.5-1)1-2.5 mm long. *Hypanthium* narrowly obovoid, 1-1.5 mm long, hispid, with white, erect hairs 0.3-0.7 mm long. *Calyx* lobes narrowly lanceolate, narrowly oblanceolate to linear, 2.8-4.5 × 0.3-0.5 mm, glabrous throughout, sometimes with a few hispid hairs at margins, with 1 deltoid, 0.1 mm long colleter at each lobe sinus. *Corolla* hypocrafteriform, 11.5-16(-18) mm long, pink to lilac; tube narrowly cylindrical, 7-9 mm long, glabrous outside, sparsely villose at mouth inside, glabrous below; lobes ovate to oblong-ovate, 4.5-8 × 2-4 mm, obtuse to round at apex, glabrous. *Filaments* 1.5-2 mm long, anthers oblong-elliptic, 0.2-0.3 × 0.1 mm. *Style* 10-13 mm long, glabrous; style branches ovate, 0.2-0.3 mm long. *Capsules* pedicellate, globose, 2.5-3 × 2.5-3 mm, sparsely hispid. *Seeds* 0.2-0.5 × 0.2-0.5 mm, strongly foveolate.

**Taxonomic observations:** Chamisso (1834: 242) in the protologue of *Sipanea erythraeoides* Cham. cited the gathering "Brasilia inter tropicos. Sellow." When he published this name he worked at Berlin. The original material at B was destroyed during WW2 (R. Vogt, pers. comm.). In addition, no original specimen with this name collected by Sellow is present at M (A. Fleischmann, pers. comm.). After exhaustive searches in several other herbaria, no original material of *S. erythraeoides* collected by Sellow could be found. Therefore the specimen *Riedel 409* at BR, barcode 00000824532, is here designated the neotype of this name, and the duplicates of this gathering at BM, FI, P, and US are isoneotypes.

This species is easily distinguishable from the others of the same genus, by its prostrate or decumbent habit, pedicellate flowers, and pink to lilac corollas.

**Geographic distribution and ecology:** Species restricted to Brazil. It has been collected in the states of Bahia, Goiás, Distrito Federal, Mato Grosso, Mato Gros-
<figure><img src="image_18.png" /><figcaption>**Figure 14.** *Limnosipanea erythraeoides*. A. Habit. B. Leaf. C. Node with base of two leaves, and extremely reduced stipule with three partially exserted colleters. D. Dissected flower, with a flower bud, subtended by two leaves. A–D: drawn from Macedo 5537 (US). Illustration by Piero Delprete.</figcaption></figure>
so do Sul, Minas Gerais, Rondônia and São Paulo (Figure 15). Ephemeral to short-lived (2–4 months life span), prostrate herb, terrestrial or semi-aquatic, commonly found in inundated open fields, at margins of small water courses, and swamps with *Mauritia flexuosa* L.f. palm communities.

**Phenology:** Flowering and fruiting from June to November, which in Central Brazil correspond to the dry season, with one collection in December which corresponds to the beginning of the rainy season.

**Suggested conservation status:** Least Concern (LC). This species occurs in numerous Brazilian states (see geographic distribution and specimens cited). It is a prostate herb, commonly found in seasonally inundated open fields, at margins of small water courses,

and swamps. The size of the populations is unknown. In addition, it is poorly collected due to its small size and short life span. Taking into account the numerous sites of occurrence, this species is here treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** BRAZIL: Bahia: Near Salvador, “Habitat in aquaticis uliginosis ad Soteropolin, Prov. Bah.”, Dec. [1818], *C.F.P. Martius Iter Brasil. 2884* (M M0198161); without locality, “Prov. Bahiensis”, s.d. [Oct.–Nov. 1818], *C.F.P. Martius Iter Brasil. 2885* (M M0198163); between Lages and Caetité, “Habitat in campis humidis ad Lages et Caiteté [ca. 14°4’S, 42°28’W]”, Oct.–Nov. [1818], *C.F.P. Martius Iter Brasil. 2886* (M M0198162). **Distrito Federal:** Ca. 25 km E of Brasília, low gallery forest, creek margin, wet soil, 21 Aug. 1964 (fl), *H.S. Irwin & T.R. Soderstrom 5429* (NY,

<figure><img src="image_19.png" /><figcaption>**Figure 15.** Distribution of *Limnosipanea erythraeoides*.</figcaption></figure>
UB). **Goiás**: Rd. Aruanã-Araguapaz, at 22.4 km from Aruanã [ca. 14°55'S, 51°1'W], swamp near the river, with *Bacopa myriophylloides* and *Drosera*, amphibious plant, 15 Nov. 1999 (fl), C.P. Bove et al. 594 (R); rd. Caiapônia-Iporá, 54 km from jct, at river margin, inside water, submersed leaves red, emerged leaves green, 19 Nov. 1999 (fl), C.P. Bove et al. 672 (HUNI, R); Mun. Goiânia, Dec. 1936 (fl), A.C. Brade 15461 (RB); rd. Caiapônia-Jataí, left margin, near spring with sulphur water, 14 Oct. 1968 (fl), S. Fonseca & E. Onishi 1542 (F, RB, UB); without locality, 21 Dec. 1873 (fl), A. Glaziou s.n. (P P00729458); Pouso Alto, nos pantanos, 24 Aug. 1894 (fl, fr), A. Glaziou 21340 pro parte (mixed collection; mixed with some individuals of *Acisanthera pulchella*, *Melastomataceae*) (R); "Ponte Lavrado à Cuba," 18 Aug. 1894 (fl), A. Glaziou 21692 (BR, P [2 sheets], R, S); Mun. Jataí, locality Balsamo, 19 Jul. 1951 (fl), A. Macedo 3334 (S, US); Mun. Quirinópolis, Serra da Confusão do Rio Preto, Faz. Boa Sorte [ca. 18°26'S, 50°26'W], no meio da vereda, em local alagado, 757 m, 8 Aug. 2014 (fl, fr), I.L.M. Resende 3363 (SPF); Mun. Quirinópolis, Serra da Confusão do Rio Preto, Faz. Boa Sorte, no meio da vereda, em local alagado, 757 m, 3 Oct. 2014 (fl, fr), I.L.M. Resende 4027 (SPF); Goiânia, near EFO-MARGO, base of Morro de Santo Antônio, 5 Sep. 1968 (fl), J.A. Rizzo & A.M. Barbosa 2127 (NY, UB, UFG); Mun. Morrinhos, rd. Morrinhos-Caldas Novas [ca. 17°43'S, 49°4'W], Córrego Samambaia, 29 Aug. 1970 (fl), J.A. Rizzo & A.M. Barbosa 5457-A (UFG); "Goyaz," without locality, 1816-1821 (fl, fr), A. Saint-Hilaire Catal. C1 747 (P [P00729269]). **Mato Grosso:** Poconé, Rodovia Transpantaneira, 178.4 km de Cuiabá, Pantanal, solo úmido, 116 m, 16°37'42"S, 56°46'17"W, 1 Dec. 2005 (fl), R.M. Harley et al. 55369 (HUEFS); Coxim [ca. 18°30'S, 54°45'W], May 1911 (fl), F.C. Hoene 3220 (R), 3222 (R), 3223 (R), 3225 (R); Barra do Rio Camaré, Juruena, Ilha do meio do rio, Jan. 1912 (fl), F.C. Hoene 5181 (R); Porto Velho do Rio Arinos, planta rasteira das margens do rio e de um lago, Nov. 1914 (fl), J.G. Kuhlmann 1486-K (F, R); 1º de Fevereiro, Rio Cautario, margens do rio, Jan. 1919 (fl), J.G. Kuhlmann 2362-K (B†, R; photo-B at MO, NY); Mun. Santa Cruz do Xingu, Parque Estadual do Xingu, limite NE do parque, subindo o Rio Fontourinha, 9°42'19"S, 52°21'16"W, 250 m, 1 Mar. 2011 (fl), D.C. Zappi et al. 3037 (RB). **Minas Gerais:** Uberlândia, Clube Caça e Pesca Itororó [ca. 18°55'S, 48°17'W], campo úmido, borda do brejo, 13 Aug. 1993 (fl), A.A. Arantes 44 (HUFU); Uberlândia, Clube Caça e Pesca Itororó, 29 Sep. 1994 (fl), A.A. Arantes 203 (HUFU); "Inter Tejuco et Rio Paranahyba," s.d. (fl), W.J. Burchell 5814 (BR, P); Mun. Campina Verde, Arenitos, várzea, sandy soil, 19 Jun. 1989 (fl), A. Macedo 5537 (NY, US); Mun. Campina Verde, brejo, 19°32'8"S, 49°29'11"W, 2 Jul. 2005 (fl), A. Macedo 5913 (NY, US); "Habitat in rudis campis uliginosis ad Buritizaes, crescit Chap. do Par., Provinciae MG", Sep. [1818], C.F.P. Martius Iter Brasil. 2907 (M M0198164); "Prov. Minas Geraes", without locality, s.d. [February-Sep. 1818], C.F.P. Martius Iter Brasil. 2908 (M M0198160); "olho d'agoa" [Olho d'Agua], 1816-1821 (fl, fr), A. Saint-Hilaire s.n. (P P00729270); "Minas Gerais," without locality, 1816-1821 (fl, fr), A. Saint-Hilaire Catal. C1 977bis (P [P00729272]); "Minas Gerais," without locality, 1816-1821 (fl, fr), A. Saint-Hilaire Catal. B1 1817 (P [P00729271]), 1929 (P [P00729274]); Campina Verde, Faz. São João do Lajeado, 20 Jul. 2003 (fl, fr), J.F. dos Santos 5841 (US). **Rondônia:** Rio Guaporé, Fazenda Santa Rosa, mata, 11 Jun. 1952 (fl, fr), G.A. Black & E. Cordeiro 52-14823 (IAN). **São Paulo:** Without locality, 1833 (fl), C. Gaudichaud Herbier Imperial du Brésil N. 99 (P); São Carlos, brejos dos cerrados [ca. 22°S, 47°53'W], 22 Apr. 1963 (fl), M. Kuhlmann 3077 (RB, SP); Mogy-Guaçu [Mogi Guaçu, ca. 22°22'S, 46°56'W], 27 Jul. 1889 (fl, fr), A. Lofgren 1337 (SP). **Tocantins:** Mun. Lagoa da Confusão, Ilha do Bananal, Parque Nacional do Araucoia, área ca. 3 km da sede do parque, Lago Rufino, solo escuro, floresta de galeria mesofítica, 10°26'48"S, 50°28'25"W, 210 m, 21 Mar. 1999 (fl), M. Aparecida da Silva et al. 4034 (RB). **Without state:** Without locality, s.d., A. Glaziou s.n. (P [barcode 00729458]).

### 3-2. *Limnosipanea palustris* (Seem.) Hook.f., Ic. pl. 11: 38. 1868. (Figures 16A-E, 17).

(=) *Sipanea* [as "Sipania"] *palustris* Seem., Bot. Voy. Herald 136. 1854; non *Sipanea palustris* (A. Rich.) J.H.Kirkbr., Brittonia 49: 360. 1997, *comb. illeg. super-fl.* (= *Sipanea wilson-brownei* R.S.Cowan); non *Bertiera palustris* A.Rich. ex DC. (Sep. 1830: 392 [see *Neobertiera palustris*, below]).

Type: PANAMÁ: Near Panama City, swamps, s.d. [1846-1849], B.C. Seemann 347 (*lectotype* BM [barcode BM 000614301], **here designated**; isotype type K [without barcode]; photo-BM at BM, MO, NY, US).

(=) *Limnosipanea schomburgkii* Hook.f., Ic. pl. 11: 38. 1868.

Type: GUYANA. ["Roraima", sic!], Rovuma River, s.d. [1842-1843], R.H. Schomburgk ser. II, 464 (= M.R. Schomburgk 744) (*lectotype* BM [BM000614338], **here designated**; isotype types F [Acc. No. 766892], G [2 sheets, barcodes G00379139,
G00379140], K [without barcode], P [barcode P00729461], US [barcode 00588519]; photo-P at NY; photo-G at F [Neg. No. 25681]).

(=) *Limnosipanea schomburgkii* var. *robustior* Pilg., Bot. Jahrb. Syst. 30: 198. 1901 [title page of the volume reports “1902” but the third page of the same volume reports that “Heft 2 (pp. 129–288)” was published in “2 July 1901”].

Type: BRAZIL. Mato Grosso: Near Cuiabá “gesellig in kleinen Complexen auf trockner Wiese bei Cuyabá,” Apr. 1899 (fl), *R. Pilger 400* (holotype B [barcode B100673663]).

*Herbs*, terrestrial or semi-aquatic (amphibious), erect, 5-40 cm tall, single-stemmed or sparsely branched; stems sparsely to densely appressed pubescent, or pubescent and with two kinds of hairs, antrorsely appressed and spreading intermixed. *Stipules* reduced to a line between the petioles, sparsely appressed-pubescent, with 1 to 3 colleters inserted just below the margin on the internal side, with the tips exerted beyond the margin. *Leaves* homophyllous, opposite or ternate, or sometimes ternate at basal nodes and opposite at distal nodes of the same plant, sessile; blades narrowly ovate, oblong-ovate, elliptic to lanceolate, 5-18 × 1-8 mm, round, obtuse to subobtuse at base, acute at apex, chartaceous, pale green below, drying olive-green, glabrous or sparsely appressed pubescent above and below, midvein strigose to glabrate above and below, margins sometimes ciliate; secondary veins 2-4 each side of midrib. *Inflorescence* 0.5-12 cm long, 2-4-dichotomously branched, with one flower at the base of each bifurcation, 2-many flowered; frondose, with leaf-like bracts subtending secondary branches; bracteoles subtending flowers elliptic-lanceolate to narrowly lanceolate, 4-6.5 × 1-1.5 mm, strigose-pubescent, margins ciliate. *Flowers* sessile or short-pedicellate in central flowers. *Hypanthium* narrowly ellipsoid to narrowly obovoid, 0.7-1 mm long, densely hispid, with white, erect hairs ca. 1 mm long. *Calyx* lobes narrowly lanceolate to linear, 2.5-5 × 0.3-0.6 mm, sparsely hirsute outside, glabrous inside, with 1 broadly triangular to deltoid, 0.05-0.1 mm long colleter at each lobe sinus. *Corolla* hypocrateriform, white to cream-white, 5.7-10 mm long; tube narrowly cylindrical, 3.5-6.5 mm long, strigose outside, sparsely villose at mouth

inside, glabrous below; lobes oblong-ovate, 2.2-3.5 × 0.7-0.9 mm, obtuse to round at apex, glabrous on both sides. *Filaments* 1.5-1.7 mm long; anthers oblong, 0.7-0.8 × 2.5-3 mm. *Style* 5.5-7.5 mm long, glabrous, style branches narrowly oblong, 0.5-0.8 mm long. *Capsules* sessile, globose, 2.5-3.5 × 2-2.5 mm, hispid-setose. *Seeds* 0.2-0.4 × 0.2-0.4 mm, strongly foveolate.

**Taxonomic observations:** Seeman (1854: 136) did not indicate his collection number or the herbarium of deposit for *Sipanea* [as “Sipania”] *palustris* Seem. Delprete (2010b: 624) indicated the type of this name as “Seeman 347 (holótipo, K; isotipo K, fotos em NY).” Lorence et al. (2012: 138) indicated as type “Seeman 347 (foto MO! ex BM)”. According to the *Code*, as they were published after 2001, both Delprete’s and Lorence’s statements cannot be interpreted as inadvertent lectotypifications because they did not include “here designated” or a similar expression. Therefore, the specimen *Seeman 347* at BM, barcode BM 000614301, is here designated as the lectotype of this taxon.

Hooker (1868: 38) along with the description of *Limnosipanea schomburgkii* Hook.f. cited the gathering “Schomburgk 464 (744)” without citing the herbarium of deposit. Hooker’s citation corresponds to the gathering *R.H. Schomburgk ser. II, 464* (= *M.R. Schomburgk 744*). A specimen at BM, barcode BM000614338, has the label “464, Schomburgk, Coll. 1842-3, Roraima, Brit. Guiana” is here designated as the lectotype of this name. Duplicate specimens are at F, G, K (with the handwritten label “464 (744), Sipanea, Roraima, British Guiana, Schomburgk 1843”), P and US are isolectotypes.

Pilger (1901) described *Limnosipanea schomburgkii* var. *robustior* Pilg. and cited the collection *Pilger 400* from Mato Grosso, Brazil. As he worked at Berlin, and there is only a specimen of *Pilger 400* in that herbarium, it is the holotype specimen. This variety fits within the morphological variation of the species and no variety is recognized in *L. palustris*.

Steyermark (1967: 282) differentiated *Limnosipanea schomburgkii* by the entire calyx lobes, opposite leaves, and its occurrence in “Southern Venezuela, probably British Guiana [Guyana] and Brazil” keeping *L. palustris* separated because of the bi-aristate calyx lobes, whorled leaves (3-4 per node), and restricted to Panama. As already stated by Delprete (2010b), after a detailed analysis of specimens from the whole distribution, these charac-

**Figure 16.** A-E. *Limnosipanea palustris*. A. Habit. B. Flower in anthesis. C. Dissected corolla. D. Longitudinal section of ovary and calyx. E. Dehiscent capsule. F-L. *Limnosipanea spruceana* var. *spruceana*. F. Habit. G. Dehiscent capsule. H. Longitudinal section of ovary and calyx, with colleters at lobe sinuses. I. Seeds. J. Dissected corolla. K. Young fruit subtended by two leaf-like bracts. L. Node with whorled leaves. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 61. 1974, as *L. schomburgkii* Hook.f.).
<figure><img src="image_20.png" /><figcaption>Figure 16.</figcaption></figure>
ters were found to be variable throughout the geographic range, and the two taxa are treated as synonymous.

**Distribution and ecology:** Distributed in Panama, Colombia, Venezuela, Guyana, and Brazil, on the Guiana and Brazilian Shields (Figure 18). Short-lived semi-aquatic or terrestrial herb, sprouting just after the water of the flooded area has dissipated, in seasonally inundated areas in open savannas and along streams, at 100-1200 m altitude.

*Limnosipanea palustris* (Seem.) Hook.f. has been collected at least six times in populations intermixed with *L. spruceana* var. *spruceana*: **COLOMBIA:** *Guaviare*: Río Guaviare, San José del Guaviare [ca. 2°34′N, 72°38′W], sabana, 240 m, 13 Nov. 1939 (fl, fr), *J. Cuatrecasas* 7714 (COL, F, US, mixed collection). **Meta:**

Río Meta, Orocué, 140 m, sabana, 3 Nov. 1933 (fl, fr), *J. Cuatrecasas* 4363 (COL, F, P, US; mixed collection); **VENEZUELA:** *Bolívar*: Morichal, 2 km E of Río Ori-noco, between Rio Horeda and Cerro Gavilan (Cerro Carichana), 100 m, 17 Dec. 1955 (fl, fr), *J.J. Wurdack & J.V. Monachino* 39942 (F, MO, NY, S, US, VEN; mixed collection), 39944 (NY, VEN; mixed collection). **GUY-ANA:** Rupuruni District, Chaakoitou, 29 Oct. 1979 (fl, fr), *P.J.M. Maas & L.Y.T. Westra* 4100 (*L. palustris*; K, MO, NY, P, S, U), 4101 (*L. spruceana*; BBS, F, NY, S, U).

**Phenology:** In Guyana, flowering and fruiting during the dry season, from August to October. In Central Brazil, flowering and fruiting from the end of the rainy season to the beginning of the dry season, from April to June.

<figure><img src="image_21.png" /><figcaption>**Figure 17.** Distribution of *Limnosipanea palustris*.</figcaption></figure>
**Suggested conservation status:** Least Concern (LC). Species ranging from Panama, Colombia, Venezuela, Guyana to Brazil. In Brazil it has been collected in the states of Amazonas, Bahia, Goiás, Maranhão, Mato Grosso, Pará, Piauí, Roraima, and Tocantins. It is a short-lived, semi-aquatic or terrestrial herb, usually found in seasonally inundated areas in open savannas and along streams. The size of the populations is unknown. In addition, it is poorly collected due to its small size and its short life span. Considering the numerous sites of occurrence, without imminent threats, this species is here treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

**Specimens studied:** COLOMBIA: Arauca: Mun. Arauca, corregimiento Caracol, Vda. La Maporita, Finca La Maporita, 6,933297 N, -70,461718 W, 117 m, sabana alta abierta, 10 Dec. 2015 (fl, fr), J.C. Arias et al. 3070 (COL); Mun. Arauca, corregimiento Caracol, Vda. La Maporita, Finca La Maporita, 6,946514 N, -70,452555 W, 122 m, sabana alta abierta, 11 Dec. 2015 (fl, fr), J.C. Arias et al. 3100 (COL). **Casanare:** Río Meta, Orocué [ca. 1°48'N, 72°45'W], 140 m, sabana, 3 Nov. 1933 (fl, fr), J. Cuatrecasas 4363 pro parte (Mixed collection; COL, F [2 sheets], P, US). **Guaviare:** Río Guaviare, San José del Guaviare, sabana, 240 m, 13 Nov. 1939 (fl, fr), J. Cuatrecasas 7714 pro parte (Mixed collection; COL, F, US). **Meta:** 73 km W of Las Gaviotas, along rd to Puerto Gaitán [ca. 4°18'N, 72°4'W], low lying savanna along drainage channel now dried to mud at this time of the year, 210 m, 30 Dec. 1973 (fl, fr), G. Davidse 5386 (COL, F, NY); Boca de Monte, Llanos de San Martín [ca. 3°41'N, 73°41'W], 300 m, open savanna, 16 Aug. 1950 (fl, fr), S. Galen Smith & J.M. Idrobo 1389 (COL, F), 1398 (US); Cabuyaro, 10 Jan. 1899 (fl, fr), T.A. Sprague 57 (BM); Llano de San Martín, La Quebradita, 200 m, 16 Aug. 1950 (fl, fr), J. Triana 3248(2) (COL, F). **State Unknown:** without locality, 1844 (fl, fr), J. Goudot s.n. (P P00728115).

**VENEZUELA. Amazonas:** Depto. Aтуres, 56 km NE of Puerto Ayacucho, 170 m, 22 Nov. 1984 (fl), R. Kral et al. 71775 (NY, TFAV, US). **Apure:** "plaines marecageuses entre l'Apure et l'Arauca," 1894 (fr), F. Geay s.n. (P [P00729254]); "savane marecageuses du Guaritico," 1894 (fr), F. Geay s.n. (P [P00729253]); Depto. Paez, Módulo F., Corrales de la Unellez, Carretera a La Victoria, 28 Dec. 1985 (fl), B. Stergios & P. Stergios 8802 (PORT, US). **Bolívar:** "Orenoque" [Rio Orinoco], 1844 (fr), J. Goudot s.n. (FI-Webb); Morichal, 2 km E of Río Orinoco, between Rio Horeda and Cerro Gavilan (Cerro Carichana), 100 m, 17 Dec. 1955 (fl, fr), J.J. Wurdack & J.V. Monachino 39942 pro parte (Mixed collection;

F, NY, S, US, VEN), 39944 (NY, VEN). **Guárico:** Dto. Infante, Parque Nacional Aguaro-Guariquito, Morichal Charcote, 9°24'-32'N, 67°64'-68'W, 70 m, Dec. 1981 (fl), F. Delascio, R. Montes & G. Davidse 11121 (NY, US, VEN); Dto. Infante, Parque Nacional Aguaro-Guariquito, Saladillal Los Galapagos, 9°04'-08'N, 67°52'-56'W, 60 m, Dec. 1981 (fl, fr), F. Delascio et al. 11234 (MO, NY, VEN).

**GUYANA:** Rupuruni Distr., Kuyuwini Landing, Kuyuwini River, savanna, 1 km S of river, 2°05'N, 59°15'W, 150-250 m, 9 Oct. 1992 (fl, fr), M.J. Jansen-Jacobs et al. 2796 (CAY, F, MO, NY, P, U, US); Rupuruni Distr., Dadanawa, savanna, near pond, 2°49'N, 59°31'W, 120 m, 19 Aug. 1995 (fl), M.J. Jansen-Jacobs et al. 4781 (P, U, US); Rupuruni Distr., Dadanawa, savanna, near pond, 2°49'N, 59°32'W, 120 m, 19 Aug. 1995 (fl), M.J. Jansen-Jacobs et al. 5376 (BBS, MO, NY, U, US); Rupuruni Distr., Chaakoitou, near Mountain Point, just S of Kanuku Mts., 2°56'N, 59°40'W, 29 Oct. 1979 (fl, fr), P.J.M. Maas & L.Y.T. Westra 4100 (F, K, MO, NY, P, S, U).

**BRAZIL: Amazonas:** Frechal, Maruay, entre pedras graniticas, Sep. 1927 (fl, fr), P. Luetzelburg 20448 (R); Rio Piriá, Sep. 1959 (fl, fr), N.T. Silva 555 (IAN). **Bahia:** Near Rio Piau, ca. 150 km SW of Barreiras, 14 Apr. 1966 (fl, fr), H.S. Irwin et al. 14789A (UB). **Goiás:** Mun. Cavalcante, RPPN Serra do Tombador, trilha do Borá, 13°42'2"S, 47°46'32"W, 900 m, 17 Apr. 2013 (fl, fr), E. Barboza et al. 3694 (MBM); Mun. Campos Belos, estrada para Pouso Alto, ca. 18 km a direita da Usina Mosquito, 12°55'29"S, 46°25'6"W, 680 m, 26 Apr. 2001 (fl), M.L. Fonseca et al. 2612 (RB, US); Serra do Morcego [ca. 15°S, 46°W], near Córrego Estrema, ca. 38 km NE of Formosa, 800 m, 21 Apr. 1966 (fl), H.S. Irwin et al. 15198 (NY); "Bresil Central, Salinas", May-Jun. 1844 (fl, fr), M.A. Weddell 2026 (P), 2054 (P), s.n. (P 00729260). **Maranhão:** Perizes, campo, 6 Jul. 1954 (fl, fr), G.A. Black et al. 54-16543 (IAN); Itapicurú, campo alto, 8 Jul. 1954 (fl, fr), G.A. Black et al. 54-16658 (IAN, RB); Mun. Carolina, margem esquerda do Rio Farinha, próximo a ponte na BR-010, 6°51'49"S, 47°28'6'W, 150 m, 25 Apr. 2008 (fl, fr), G.P. Silva et al. 13338 (CEN); Mun. Carolina, estrada Carolina-Babaçulandia, km 8.2, margem direita do Rio Tocantins, kms marcados da Igreja São Francisco, Bairro Brejinho, 7°18'25"S, 47°30'18"W, 170 m, 22 May 2010 (fl, fr), G.P. Silva et al. 15288 (CEN); perto de Carolina [ca. 7°19'S, 47°28'W], campo cerrado, 26 May 1950 (fl), J.M. Pires & G.A. Black 2164 (IAN), 2280 (IAN); Mun. Santa Elena [ca. 2°13'S, 45°18'W], near locality Queimados, Campos da Chapada, terra firme, 9 Jul. 1978 (fl, fr), N.A. Rosa & O.C. Nascimento 2564 (IAN, MG, NY, RB, UEC, UFG). **Mato Grosso:** Chapada dos Guimaraes, Alto do Paredão, acima do Assentado, 10
rarely glabrous; perennial plants growing in permanently flooded areas (i.e., with natural springs) with the submerged stem woody at base. *Stipules* reduced to a line between the petioles or obsolete, glabrous outside, without colleters or with 1 or 2 colleters inserted just below the margin on the internal side, included or with the tips slightly exserted beyond the margin, or, when stipules obsoletes, 1 or 2 colleters inserted at leaf sinuses; in submerged stems, stipules and colleters absent. *Leaves* homophyllous or heterophyllous, dimorphic when basal portion of plant is submerged; on submerged plants, the *submerged leaves* 3–6(-8) per node, with blades linear to acicular, 5–20 × 0.4–0.7 mm, glabrous, without secondary veins (rarely with 1 secondary vein each side); the *aerial leaves* 2–5 per node, with blades linear to lanceolate, 3.5–12 × 0.3–5 mm, glabrous, with 1–2(–3) secondary veins on each side of midrib; in plants entirely terrestrial (i.e., never partially submerged), the leaves are homophyllous and similar to those of the aerial portion of the partially submerged plants. *Inflorescences* simple, unbranched, with 1–5 flowers, or branched, 2–3-chotomously branched, with one flower at the base of each bifurcation, 2–18.5 cm long, 2–multiflorous; bracteoles subtending flowers narrowly lanceolate to linear, 2–4.5 × 0.4–1.5 mm, glabrous or sparsely pubescent. *Flowers* commonly sessile, the central ones (at the base of a bifurcation) subpedicellate. *Hypanthium* narrowly ellipsoid to narrowly obovoid, 0.5–1 × 1–1.3 mm, glabrous or sparsely to densely hispid, with white hairs 0.5–0.7 mm long. *Calyx* lobes lanceolate to linear, 1.5–5.5 × 0.3–0.6 mm, aristate at apex (arista caducous) or not, glabrous throughout, sometimes with a few hispid hairs at margins, with 1 narrowly triangular to linear, 0.1 mm long colleter at each lobe sinus. *Corolla* hypocrateriform, 3–11.2 mm long, white to cream-white, rarely pale violet; tube narrowly cylindrical or narrowly imbutiform, 2–4.7 mm long, glabrous outside, with a ring of yellow hairs at mouth, glabrous below, inside; lobes ovate to oblongovate, 1–6.5 × 0.5–2.2 mm, obtuse to round at apex, glabrous on both sides. *Filaments* 0.4–0.8 mm long; anthers oblong, 0.5–0.7 × 0.2–0.4 mm. *Style* 2.5–5 mm long, glabrous; style branches oblong, 0.3–0.5 mm long. *Capsules* ovoid to subglobose, 2–4 × 1.5–2.5 mm, hispid-setose, glabrate, or glabrous. Seeds 0.2–0.3 × 0.2–0.3 mm, strongly foveolate.

**Phenology:** This species has a very short life span, and it is common to find flowers and fruits on the same individual. Its phenology varies according to the geographic positions of the populations, and corresponds to the local seasons. In the region north of the Equator (Colombia, Venezuela, Guyana, and Brazil: Roraima),

it is flowering and fruiting from September through December. In the region south of the Equator (Brazil: Amazonas, Goiás, Mato Grosso, Minas Gerais, Pará, Rondônia, and Tocantins; Bolivia: Santa Cruz) it is flowering and fruiting from May through July (August).

**Distribution and ecology:** Distributed on the Guiana and Brazilian Shields, in Colombia, Venezuela, Guyana, Brazil, and Bolivia; short-lived, semi-aquatic or terrestrial herb, coming up just after the water of the flooded area has dissipated, in seasonally inundated areas in open savannas, along streams, and *Mauritia* palm swamp communities (in Venezuela called "morichales" or "canguchales", and in Brazil called "buritizais"), at 50–900 m altitude.

**Suggested conservation status:** See under varieties.

**Habit variation:** *Limnosipanea spruceana* is a polymorphic herb (Figure 18), single-stemmed or sparsely branched, that could be terrestrial and 7–25 cm tall (Figure 18F), or semi-aquatic (amphibious) or aquatic, with the submerged portion to 40 cm long and aerial portion to 25 cm long (including inflorescence) (Figure 18A-E). The leaves can also be dimorphic (Figure 18C-D), if some of them are submerged or not. In plants partially submerged, the leaves below water (Figure 18C) (when present) are 3–6(-8) per node, linear to acicular, 5–20 × 0.4–0.7 mm, without secondary veins (rarely with 1 secondary vein on each side of the midrib); while the aerial leaves (Figure 18D) are 2–5 per node, linear to lanceolate, 3.5–12 × 0.3–5 mm, with 1–2(–3) secondary veins each side of the midrib. In terrestrial plants (i.e., not partially submerged), the leaves are homophyllous and similar to those of the aerial portion of the partially submerged plants.

The variation in life span, habit and leaf shape is correlated with the water level present during a given season. In localities seasonally inundated, during the rainy season the water level varies from 0.3 to 2 m in depth, and might reach up to 4–5 m in depth in Amazonian savannas (Figure 19A-B). As the precipitations decrease, the water level gradually drops until the soil is still moist but no longer flooded; at this precise moment the terrestrial form of this species appears (Figure 18F), sometimes forming large stands, visible from a distance due to their characteristic red stem. In this form, the leaves are opposite or whorled, 3–5 per node, and linear to lanceolate. This form is particularly short-lived (Figure 18E-F), and can grow, disperse seeds and die within 3–4 months (pers. obs.; P.G. Delprete et al. 7433-7437).

In permanently flooded areas, as in localities with natural springs, the submerged portion of the stem could
be up to 40 cm long, not branched, green, with the basal portion leafless, with adventitious roots at each node, and the distal submerged portion with whorled leaves, 3-6(-8) per node, linear to acicular (Figure 18C), and without adventitious roots (Figure 18A-D). The aerial portion of these plants is 10-25 cm long (Figure 18A), including inflorescences, branched, with stems and rachis red-vinaceous, and leaves opposite or whorled, 3-5 per node, linear to lanceolate (Figure 18D). This form is perennial (pers. obs.; *B.S. Marimon* & *P.G. Delprete BS-975*).

In other cases, in seasonally flooded areas, with water 20-30 cm deep during the rainy season, plants of *Limnosipanea spruceana* are seasonally semi-aquatic (amphibious), with dimorphic leaves (Figure 18E); the submerged portion of the plants has simple or sparsely branched stems, with 3-6(-8) leaves per node, linear to acicular; while the aerial portion is 10-20 cm long (including inflorescences), with 2-5 leaves per node, linear to lanceolate. At the end of the rainy season, when the water is completely gone, the portion of stem that was submerged is laying on the ground, decumbent, with the distal portion erect (Figure 18E); at this stage, the whole stem and the rachis are red, due to sun exposure. This form is particularly short-lived, and can grow, disperse seeds and die within 3-4 months (pers. obs.; e.g., *B.S. Marimon* & *P.G. Delprete BS-976*).

Key to the varieties of *Limnosipanea spruceana*

1. Corollas 3-6(-7) mm long, with tubes 2-3(-3.5) mm long, lobes ovate to oblong-ovate, 1-3(-3.5) × 0.5-0.7 mm; filaments 0.4-0.8 mm long; anthers 0.5-0.7 mm long; style 2.5-5 mm long....................3a. *L. spruceana* var. *spruceana*

1. Corollas 8.5-11.2 mm long, with tubes 3.5-4.7 mm long, lobes oblong to oblong-ovate, 4.5-6.5 × 1.5-2.2 mm; filaments 1.3-1.8 mm long; anthers 1 mm long; style 6.5-9 mm long .......................... 3b. *L. spruceana* var. *macrantha*

3-3a. *Limnosipanea spruceana* Hook.f. var. *spruceana*, Icon. Pl. 11: 38, pl. 1050. 1868. (Figures 4E-F, 16F-L, 18-21).

Type: BRAZIL. Pará: "Amazon R., marshy and sandy places near Pará" [Santarem, at the mouth of the Tapajós River], Aug. 1850 (fl), *R. Spruce 1027* (lectotype K [without barcode], designated by Delprete (2010b: 627); isolectotypes P [barcode 00729466], RB [ex P, barcode 00543629]; possible isolectotypes (without collector number) BM [barcode BM000614344], E [2 sheets, barcodes E00499997, E00499998], F [2 sheets, Acc. Nos. 768300 and 971288], FI-Webb [barcode FI004806], MPU [barcode MPU 021333], P [barcode 00729465]; S [Acc. No. S05-474]; photo-B and photo-K at NY).

(=) "*Sipanea* [as "*Sipania*"] *limnophila* Spruce" cited in synonymy by J.D. Hooker (1868: 38, tab. 1050), not valid, *pro syn.*

Type: BRAZIL. Pará: [Santarem, at the mouth of the Tapajós River], marshy and sandy places "near Para", Aug. 1850 (fl), *R. Spruce 1027* (**lectotype** K [without barcode], **here designated**; isolectotypes P [barcode 00729466], RB [ex P, barcode 00543629]; probable isolectotypes (without collector number) BM [barcode BM000614344], P [barcode 00729465]; possible isolectotypes (without collection number) E [2 sheets, barcodes E00499997, E00499998], F [2 sheets, Acc. Nos. 768300 and 971288], FI-Webb [barcode FI004806], MPU [barcode MPU 021333], S [Acc. No. S05-474; photo-B and photo-K at NY).

(=) *Limnosipanea kuntzei* Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 280. 1931.

Type: BOLIVIA. Velasco, 200 m, Jul. 1892 (fr), O. *Kuntze s.n.* (holotype US [Acc. No. 701742, barcode 00137736]; isotypes NY [2 sheets, barcodes 000132101 and 000132102).

(=) *Limnosipanea ternifolia* Pittier, Bol. Soc. Venez. Ci. Nat. 8: 144. 1943.

Type: VENEZUELA. Guárico: Llanos de Calabozo, Laguna de Palo Seco, entre la Encrucijada y la Mis-

**Figure 18.** *Limnosipanea spruceana* var. *spruceana*. **A-D.** Perennial form growing in permanently flooded areas, with heteromorphic leaves. **A.** Habit, with basal submersed portion with adventitious roots, medial submersed portion leafless, distal submerged portion with acicular leaves, and aerial portion with whorled leaves and terminal inflorescence. **B.** Basal portion of the submerged portion of the stem with adventitious roots at each node. **C.** Distal submerged portion of the stem with whorled, acicular leaves. **D.** Aerial portion of the stem with whorled, lanceolate leaves. **E.** Form growing in seasonally flooded area, at the end of the rainy season (water completely dissipated), with decumbent stem; basal portion of stem, originally submerged, leaning on the ground, and distal portion of the stem erect, with opposite or whorled leaves. **F.** Terrestrial, short-lived form, found in seasonally inundated localities, with homomorphic leaves basally opposite and distally ternate. **A-D:** drawn from *Marimon & Delprete BS-975* (CAY); **E:** drawn from *Marimon & Delprete BS-976* (CAY); **F:** drawn from *Cuatrecasas 7714* (F). Illustration by Piero Delprete.
<figure><img src="image_22.png" /><figcaption>Figure 18.</figcaption></figure>
<figure><img src="image_23.png" /><figcaption>**Figure 19.** *Limnosipanea spruceana* var. *spruceana*. **A.** General environment. During the rainy season (July–November) the water level reach 2–3 m in depth above the ground; the plants start growing in November–December, when the water above ground has almost completely dissipated, and the seedlings are fully or partially submersed. At the moment that these photos where taken, the soil was moist but without standing water. The red patches are populations of *L. palustris*. **B.** Detail of populations of *L. palustris* (red patches). **C–D.** Habit. **E.** Detail of branches with flower buds and corollas about to open. **F.** Detail of a branch with a flower in anthesis. Photos taken by Piero Delprete on 3 January 2001, in Mun. Puerto Ayacucho, Amazonas state, Venezuela, where Delprete et al. 7433–7437 were collected.</figcaption></figure>
ión de Arriba, en terrenos periodicamente anegados, "10 Nov. 1941" [19 Feb. 1942] (fl, fr), H. Pittier 14952 [same as T. Lasser 130 (US, VEN, paratypes)] (holotype VEN [Acc. No. 15510]; isotypes F [Acc. No. 1268454], US [barcode 00137737]). While Pittier (1843: 144-145) in the publication reported the collection date "10 November 1941", the specimen label at VEN reports the collection date "19 February 1942".

(=) *Limnosipanea guaricensis* Pittier, Bol. Soc. Venez. Ci. Nat. 8: 145. 1943.

Type: VENEZUELA. Guárico: Llanos de Calabozo, en cienaga, Laguna de Palo Seco, cerca de la Misión de Arriba, en terrenos temporalmente anegados, "10 November 1941" [21 Feb. 1942] (fl, fr), T. Lasser 145 (holotype VEN [Acc. No. 15508]). While Pittier's (1943: 145) publication reports the collection date "10 November 1941" the specimen label at VEN reports the collection date "21 February 1942".

*Herbs*, terrestrial, semi-aquatic (amphibious) or aquatic, polymorphic, commonly ephemeral, or perennial when growing in permanently submerged areas; erect or decumbent, single-stemmed or sparsely branched; terrestrial plants 10–30 cm tall; semi-aquatic or aquatic plants with stems 7–65 cm long, submerged portion up to 50 cm long, aerial portion 5–15 cm long, appressed-strigose to appressed-pubescent or rarely glabrous; perennial plants growing in permanently flooded areas (i.e., with natural springs) with the submerged stem woody at base. *Leaves* homophyllous or heterophyllous, dimorphic when basal portion of plant submerged; on submerged plants, the *submerged leaves* 3–6(–8) per node, with blades linear to acicular, 5–20 × 0.4–0.7 mm, glabrous, without secondary veins (rarely with 1 secondary vein on each side of midrib); the *aerial leaves* 2–5 per node, with blades linear to lanceolate, 3.5–12 × 0.3–5 mm, glabrous, with 1–2(–3) secondary veins each side; in plants entirely terrestrial, (i.e., never partially submerged), the leaves are homophyllous and similar to those of the aerial portion of the partially submerged plants. *Inflorescences* simple, not branched, with 1–5 flowers, or branched, 2–3-chotomously branched, with one flower at the base of each bifurcation, 2–18.5 cm long, 2–multiflorous; bracteoles subtending flowers narrowly lanceolate to linear, 2–4.5 × 0.4–1.5 mm, glabrous or sparsely pubescent. *Hypanthium* narrowly ellipsoid to narrowly obovoid, 0.5–1 × 1–1.3 mm, sparsely to densely hispid, with white, erect hairs 0.5–0.7 mm long, or glabrous. *Calyx* lobes lanceolate to linear, 1.5–5.5 × 0.3–0.6 mm, aristate at apex (arista

caducous) or not, glabrous throughout, sometimes with a few hispid hairs at margins, with 1, narrowly triangular to linear, 0.1 mm long, colleter at each lobe sinus. *Corolla* hypocrateriform, 3–6(–7) mm long, white, cream-white to pink, rarely pale violet; tube narrowly cylindrical or narrowly imbutiform, 2–3(–3.5) mm long; lobes ovate to oblong-ovate, 1–3(–3.5) × 0.5–0.7 mm, obtuse to round at apex, glabrous on both sides. *Filaments* 0.4–0.8 mm long; anthers oblong, 0.5–0.7 × 0.2–0.4 mm. *Style* exserted 0.7–1.5 mm beyond corolla mouth, 2.5–5 mm long, glabrous; style branches oblong, 0.3–0.5 mm long. *Capsules* ovoid to subglobose, 2–4 × 1.5–2.5 mm, hispid-setose, glabrate, or glabrous. *Seeds* 0.2–0.3 × 0.2–0.3 mm, strongly foveolate.

**Notes:** Hooker (1868: 38), along with the original description of *Limnosipanea spruceana* cited two collections made by Spruce in the Brazilian Amazon: 1) "*Sipania* [sic!] *limnophila*, *Spruce 1027*", and 2) "Amazon River, marshy and sandy places near Pará [now Belém], *Spruce (1851)*."

Steyermark (1967) erroneously cited the type of *Limnosipanea spruceana* as "Vicinibus Santarem, Prov. Pará, Brazil, August 1850, *Spruce 677*" without making any reference to the gathering cited by Hooker in the original publication, and without citing the herbarium of deposit. Because of these errors, Steyermark's typification is invalid. Therefore, a new lectotypification was necessary, selecting a specimen out the gatherings cited by Hooker. Delprete (2010b: 627) designated the specimen *Spruce 1027* preserved at K, where Hooker worked, as the lectotype for this taxon. The lectotype specimen bears the label "1027 Sipania limnophila sp. n., Santarem marshy and sandy valley inundated by the Tapajos [River], [...]", *Spruce 1851*." The information "Sipania limnophila sp. n. [nom. illeg., pro syn.]" and "*Spruce 1851*" are in darker ink, and were probably written by an unknown author, and "1851" refers to the collection date. A duplicate specimen of *Spruce 1027* at RB is an isotype. The specimen has a label with the heading "Herb. Mus. Paris," the stamp "Ex Herbario Musei Parisiensis" and the handwriting (author unknown) "*Spruce 1027*."

Hooker (1868: 38) cited "*Sipanea* [as "*Sipania*"] *limnophila* Spruce" in synonymy under *Limnosipanea spruceana* and cited the gathering *Spruce 1027* for both names. Therefore, *L. limnophila* is not a valid name.

For additional information about the synonymy of *Limnosipanea spruceana* and *L. ternifolia* Pittier, see Delprete & Steyermark (2004b).

**Distribution and ecology:** Distributed on the Guiana and Brazilian Shields, in Colombia, Venezuela
la, Guyana, Brazil, and Bolivia (Figure 20). Short-lived semi-aquatic or terrestrial herb, coming up just after the water of the flooded area has dropped, in seasonally inundated areas in open savannas, along streams, and *Mauritia* palm swamps, or a perennial herb when growing in permanently submerged areas; growing at 50–900 m altitude.

**Phenology:** See phenology of the species (above).

**Suggested conservation status:** Least Concern (LC). This variety has the same geographic distribution as the species, and ranges from Colombia, Venezuela, and Guyana, to Bolivia and Brazil. In Brazil it is known by numerous collections from the states of Amazonas, Goiás, Mato Grosso, Minas Gerais, Pará, Rondônia, Roraima and Tocantins. It is a terrestrial, semi-aquatic (amphibious) or aquatic, polymorphic herb, commonly short-lived, or perennial when growing in permanently inundated areas.

Personal observations of a large population were made in January 2001 (*Delprete et al. 7433–7437*) at about 45 km north of the city of Puerto Ayacucho, state of Amazonas, Venezuela. That area was seasonally inundated, with water levels reaching 2–3 m above the ground during the rainy season, which in that region spans from July to October. At the time the locality was visited, at the beginning of the dry season, the water had receded, the soil was moist, without standing water, and it was dominated by herbs 30–70 cm tall. According to the information supplied by the locals, the area was subjected to sporadic cow grazing during the dry season; however, the cows were not present at that moment. The population of this species covered an area of several hectares and was estimated to be of several thousands individuals. Apparently the population was not impacted by the sporadic cow grazing, because, due to the ephemeral life span, the plants succeeded in germinating, growing and dispersing seeds before the cows were allowed in the area.

Two populations were personally observed in June 2007 in the state of Mato Grosso, Brazil. One population, near the town of Nova Xavantina (*Marimon & Delprete BS-975*), was growing in a permanent lagoon, with crystal-clear water originating from natural springs. The water depth ranged from 15 to 70 cm deep. The population was composed of perennial, amphibious individuals, with the submerged portion of the stem 10–50 long, and the aerial portion with flowers in anthesis. The population was quite healthy, esteemed to be a few hundred individuals, although their exact number was not counted.

The other population observed in the state of Mato Grosso was located in the municipality of Barra do Gar-

ça, along the road BR-158 (*Marimon & Delprete BS-976*). That area was seasonally inundated, with water levels reaching 20–30 cm above the ground during the rainy season, observed by watermarks on local vegetation, which in that region spans from the end of September to May. At the time it was visited, 10 June 2007, the water had receded, the soil was moist, without standing water, and was dominated by small herbs. The population covered an area of numerous hectares and was esteemed to be of several thousand individuals. In the area, numerous cow footprints were observed, although no cow was seen at that moment. As observed in the seasonally inundated area in Venezuela, apparently the population was not impacted by the presence of cows.

Taking into account the vast extent of occurrence (EOO > 20,000 km²), numerous sites of occurrence, and the above observations, this species is here treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** COLOMBIA: **Casanare:** Río Meta, Orocué, 140 m, sabana, 3 Nov. 1933 (fl, fr), *J. Cuatrecasas 4363 pro parte* (Mixed collection; F [2 sheets], P, US); Mun. Paz de Ariporo, Corr. La Hermosa, Finca Nicaragua, Congrial, llanura aluvial de desborde, 5°36'35"N, 70°21'48"W, 115 m, 8 Nov. 2004 (fl, fr), *J.G. Ramírez Aranço et al. 9259* (COL). **Guaviare:** Río Guaviare, San José del Guaviare [ca. 2°34'N, 72°38'W], sabana, 240 m, 13 Nov. 1939 (fl, fr), *J. Cuatrecasas 7714 pro parte* (Mixed collection; COL, F, US). **Meta:** Serrania entre los Ríos Ariari y Meta, Llano Grande, matas de monte, 320 m, 26 Nov. 1939 (fl), *J. Cuatrecasas 7885* (COL, F, US).

**VENEZUELA:** **Amazonas:** Depto. Aтуres, Mun. Puerto Ayacucho, ca. 45 km N of Puerto Ayacucho, on rd towards Parhueña, area seasonally inundated, field dominated by grasses and small herbs 30–70 cm tall, 5°45'N, 67°30'W, 150 m, 3 Jan. 2001 (fl, imm fr), P.G. *Delprete et al. 7433* (CAY, GB, L, MO, NY, P, ufg, VEN), *7434* (B, BR, K, US, VEN), *7435* (CAY, G, GB, MO, P, US, VEN), *7436* (F, K, MG, NY, UFG, VEN), *7437* (L, MO, VEN). **Bolívar:** Dtto. Cedeño, km 12 de la carretera Caicara del Orinoco-Puerto Ayacucho, S of Caicara del Orinoco, 6°50'N, 66°30'W, 100 m, 18 Nov. 1984 (fl), G. Aymard & B. Stergios 3313 (NY, PORT); Dtto. Roscio, sabanas húmedas, morichales y bosques de Galeria en el Hato La Divina Pastora, ca. 8 km N de Santa Elena de Uairén, 4°42'N, 61°2'W, 840 m, 6 Oct. 1982 (fl, fr), O. Huber & C. Alarcon 6701 (NY, US, VEN); Morichal, 2 km E of Río Orinoco, between Rio Horeda and Cerro Gavilan (Cerro Carichana), 100 m, 17 Dec. 1955 (fl, fr), J.J. Wurdack & J.V. Monachino 39942 pro parte (Mixed
collection; F, NY, S, US, VEN), 39944 (NY, VEN). **Guárico:** Ca. 32 km SSE of Calabozo, on Finca Becerra, in and around morichal with standing water, morichal dominated by *Mauritia flexuosa*, 65 m, 6 Nov. 1973 (fl, fr), *G. Davidse* 3791 (MO, NY, VEN); Dto. Infante, Parque Nacional Aguaro-Guariquito, Congriales de La Gorra, entre La Esperanca y Mesa de Cambao, 9°12'-16'N, 67°48'-60'W, 60 m, Dec. 1981 (fl, fr), *F. Delascio* et al. 11204 (MO, NY, VEN); Llanos de Calabozo, near Mision de Arriba, sandy soil, 10 Nov. 1941 (fl, fr), *T. Lasser* 130 (US, VEN, paratypes); Morichal La Becerra, al S de la Estación Biológica, vía paso del Caballo, 5 Nov. 1988 (fl), *P. Suárez* S. 42 (NY, VEN).

**GUYANA:** Rupuruni Distr., Dadanawa, Rupununi River, around pond, 2°50'N, 59°30'W, 100-150 m, 23 Sep. 1992 (fl, fr), *M.J. Jansen-Jacobs* et al. 2615 (BBS, CAY, K, NY, P, U, US); Rupuruni Distr., Chaakoitou, near Mountain Point, just S of Kanuku Mts., 2°56'N, 59°40'W, 29 Oct. 1979 (fl), *P.J.M. Maas* & *L.Y.T. Westra* 4101 (BBS, F, NY, S, U).

**BRAZIL:** **Amazonas:** Mun. Humaitá, lagoa temporária entre igarapés Retiro e Bom Futuro, N da estrada para Manaus, a 15 km da cidade, vegetação flutuante aparecendo 20 cm acima da água, 7°31'S, 63°10'W, 11 May 1980 (fl), *J.D. Gemtchujnicov* & *A. Janssen* 366 (US); Mun. Humaitá, lagoa temporária entre igarapés Retiro e Bom Futuro, campo N da estrada 319, 657 km, vegetação flutuante aparecendo 20 cm acima da água, 7°31'S, 63°10'W, 11 May 1980 (fl), *A. Janssen* & *J.D. Gemtchujnicov* 368 (INPA, MG, RB, SP). **Goiás:** Mun. Uruaçu, 13 km NE of village of Água Branca, 27 km from BR-153 (Belém-Brasília), region of Fazenda Amarra Cachorro, campo limpo de murundum, trans. para o cerrado em toda direção, solo areno-argiloso encharcado no local, camada de turfa, area inundated by the dam of Serra da Mesa, 14°22'S, 49°1'W, 470 m, 6 Aug. 1992 (fl), *B.M.T. Walter* et al. 1956 (CEN). **Mato Grosso:** Vila Bela da Santíssima Trindade, Rod. MT-246, Pontes e Lacerda-Vila Bela da Santíssima Trindade, 35 km após ponte sobre rio na saída de Pontes e Lacerda, margem esquerda, estrada de accesso a propriedade particular, 1 km rumo ao sul, 15°8'6"S, 59°37'44"W, 239 m, 18 Jun. 2012 (fl, fr), *L.M. Borges* et al. 766 (RB, SPF); Road Barra do Garças-Xavantina, 25 km from Xavantina, standing water and margin of ponds, white fl., herb, rooted under water, 9 Jun. 1966 (fl), *D.R. Hunt* & *J.F. Ramos* 5902 (NY, P, R, SP, UB), 5904 (NY, SP, UB); near Cuiabá, Aricá, 5 Jun. 1894 (fl, fr), *G.O.A. Malme* 1662 (F, R, S[2 sheets], US), 8 May 1903 (fl, fr), 2260 (F, S), 8 May 1903 (fl), 3260 (F, R); Mun. Nova Xavantina, town of Nova Xavantina, Rua da Palha, Lagoa do Saibro, lagoa natural com água cristalina, com nascentes nas margens, água de 15-70

cm de profundidade, erva anfibia, monocaule, porção submersa de 10-50 cm de compr., caule submerso verde, com raizes adventicias brancas, caule emerso e raquis vermelho-vinoso, corolas brancas, 14°40'34"S, 52°20'2"W, 280 m, 8 Jun. 2007 (fl), *B.S. Marimon* & *P.G. Delprete* BS-975 (B, BR, C, CAY, F, G, GB, K, L, MG, MO, NY, NX, P, RB, S, UFMT, US); Mun. Barra do Garças, estrada Barra do Garças-Nova Xavantina (BR-158), km 747, campo sazonalmente inundado, neste momento sem água em cima do solo, pela vegetação é possivel deduzir que a água chegou a 20-30 cm de profundidade, erva anfibia, monocaule, porção que era submersa (agora deitada no chão) de 10-20 cm, porção emersa de 10-20 cm, caule e raquis completamente vermelhos porque expostos ao sol, corolas brancas, 15°27'17"S, 52°12'6"W, 300 m, 10 Jun. 2007 (fl), *B.S. Marimon* & *P.G. Delprete* BS-976 (B, BR, C, CAY, F, G, GB, K, L, MG, MO, NY, NX, P, RB, S, UFMT, US); Mun. Poconé, estrada Poconé-Porto Cercado, ca. 4 km após o Rio Bento Gomes, herbácea, flores alvas, frutos jovens avermelhados, frequente em pequenas ondulações do terreno com acumulo de materia organica, 27 May 1988 (fl, fr), *Pantanal do Poconé* 25 (UFMT); Poconé, Estrada para Porto Cercado, Faz. Santa Carmen, km 18, campo de murundum com mata, sazonalmente alagável, seco [no momento da coleta], 22 Apr. 1993 (fl, fr), *A.L. Prado* 2022 (UEC); Poconé, Estrada Poconé-Porto Cercado, Faz. Santa Carmen, km 18, campo sazonalmente alagável, nivel de água 5 cm, 13 May 1992 (fl, fr), *A.L. Prado* et al. 2783 (UEC). **Mato Grosso do Sul:** Coxim [ca. 18°30'S, 54°45'W], campos humidos, May 1911 (fl, fr), *F.C. Hoene* 3233 (R), 3234 (R). **Minas Gerais:** “ante Pirapora” [near town of Pirapora, ca. 17°20'S, 44°56'W], s.d. [1820], *J.E. Pohl* 3326 (F [Acc. No. 869868], W [3 sheets; Acc. Nos. W-0072337, W-0072339, W-0072340] (see below for discussion). **Pará:** Óbidos, campos ao NE do Lago Mariapixy, 19 Jul. 1912 (fl), *A. Ducke s.n.* (MG 12014) (MG); [Ilha do Marajó] Curralinho [ca. 1°48'S, 49°47'W], campo natural, 22 Aug. 1948 (fl, fr), *J.M. Pires* 1263 (IAN); Canaã dos Carajás, Serra da Bocaina - Serra dos Carajás, Lagoa do Buritiranal, margem da lagoa de canga, secando, 6°18'43"S, 49°52'21"W, 692 m, 23 Jun. 2015 (fl), *N.F.O. Mota* et al. 3406 (BHCB); Santarem, Aug. 1850 (fl, fr), *R. Spruce* 677 (B†, M [barcode M-0198159], NY [barcode 00132103], RB [without collection number, RB 17398]; photo-B at F; erroneously cited by Steyermark (1967) as the type of *L. spruceana*). **Rondônia:** [“Guaporé”], Rio Guaporé, Fazenda Santa Rosa, planta aquatica, 11 Jun. 1952 (st), *G.A. Black* & *Cordeiro, E.* 52-14889 (IAN). **Roraima:** Rio Cantá, campina, 8 Oct. 1951 (fl, fr), *G.A. Black* 51-13865 (IAN); Boa Vista (“Rio Branco”), Oct. 1913 (fl, fr), *J.G. Kuhlmann* 851 (RB [2 sheets]); Manaus-Vene-
zuela road (BR-174), estrada Manaus-Boa Vista, between Mucajaí and Boa Vista, W side of Rio Branco opposite Serra Grande, damp savanna on sand, 21 Nov. 1977 (fl), W.C. Steward et al. 142 (NY); Manaus-Venezuela road (BR-174), Malacacheta, E of Boa Vista, wet savanna on sand, 4 Dec. 1977 (fl, fr), W.C. Steward, I. Araujo, W.R. Buck, J.F. Ramos & J. Ribamar 251 (INPA, NY, P). **Tocantins:** Mun. Santa Isabel, Ilha do Bananal, Parque Nacional do Araguaia, caminho para Riozinho, campo de murundum, solo periodicamente inundável, 20 Jun. 1979 (fl, fr), F. Cardoso da Silva et al. 242 (UB).

**BOLIVIA: Santa Cruz:** Prov. Velasco, Parque Nacional Noel Kempff Mercado, pampa húmeda SW del campamento, 13°33'3"S, 61°3'31"W, 200 m, 25 May 1994 (fl, fr), E. Gutiérrez et al. 1132 (MO, NY); Prov. Velasco, Estancia Flor de Oro, W side of the Rio Guaporé (Rio Iténez), flat, grazed grass savanna near airstrip, with regularly spaced brush-covered and/termite mounds with *Cybistax antisyphilitica*, *Brosimum gaudichadii*, *Curatella americana*, *Simarouba*, on fine recent alluvium, 13°33'S, 61°1'W, 190 m, 21 Jun. 1991 (fl, fr), M. Nee 41158 (MO, NY); Velasco, Parque Nacional Noel Kempff Mercado, Pampa Toledo, 5-10 km NE del Campamento Los Fierros, sobre el camino a Tarbo, pampa húmeda, con islas arbóreas termiteras, 200 m, tallos rojizos, flor rosada, fruto pubescente y rojizo, 1 Jul. 1993 (fl, fr), M. Saldias et al. 2897 (US).

**Specimens with erroneous locality: BRAZIL:** "Rio de Janeiro: São João da Barra, dans les bourbiers [in swamps]", 6 Feb. 1876 (fl, fr), A. Glaziou 9887a (P [P00729291], R [Acc. No. 10468, barcode R010049086]).

The specimen labels of *Glaziou 9887* and *Glaziou 9887a* report a dubious collection locality, with different collection dates and different ecologies. The labels of both gatherings indicate that they were collected in São João da Barra [ca. 21°38'24'S, 41°3'3"W], which is a coastal town at the northern portion of the State of Rio de Janeiro, mostly "floresta de tabuleiro" and "restinga" vegetation. The gathering *Glaziou 9887* is *Sipanea pratensis* var. *dichotoma*; which is a variety that does not occur in the state of Rio de Janeiro (see discussion under this variety). The label of the specimen of *Glaziou 9887a* at R (Acc. No. 10468, barcode R010049086) has the heading "PLANTAS DO BRASIL CENTRAL, GLAZIOU N°" and the handwritten text "9987a. Limnosipanea Spruceana Hook., Flora Bras. VI, VI, p. 253. São João da Barra (Rio de Jan. [Rio de Janeiro]), 6 de Fevereiro de 1876. Herbacea flores rosas."

The specimen of *Glaziou 9887a* at P, barcode P00729291, has two labels. At the lower left corner is a label handwritten by Glaziou in pencil reporting "N.

9887a. São João da Barra, dans les bourbiers (Rio_J), 6 fevrier 1876. Fl. roses." The other label, at the lower right corner, has the printed heading "HERB. MUS. PARIS" and the text, handwritten in black ink by Glaziou "Limnosipanea Spruceana Hook. BRÉSIL (Rio Janeiro) São João da Barra, dans les bourbiers [in swamps], 6 fevrier 1876, Fl. roses, n. 9887a" and the lower printed text "Herbier de A. Glaziou, donné par Mme SIMART, sa fille, en 1907." [Glaziou herbarium, donated by Mrs. Simart, his daughter, in 1907].

The specimens *Glaziou 9887a* at P and R are *Limnosipanea spruceana* var. spruceana. Schumann (1889) did not cite the gathering *Glaziou 9887a* in *Flora Brasiliensis*. This variety is a terrestrial, semi-aquatic or aquatic herb, usually found in seasonally or permanently inundated areas, along streams and in swamps. It ranges from Colombia, Venezuela, the Guianas, to north-central Brazil. In Brazil it has been collected in the states (arranged from North to South) of Roraima, Amapá, Amazonas, Pará, Tocantins, Rondônia, Mato Grosso, Goiás, and Minas Gerais. The closest collection to the state of Rio de Janeiro was made by Pohl in 1820 (*J.E. Pohl 3326* (F, W)) near Pirapora [ca. 17°20'S, 44°56'W], a town on the margin of the São Francisco River, in the northern portion of the state of Minas Gerais; this is the only collection reported from this state, and the southernmost collection of this species (see below). The town of Pirapora is about 630 km air distance from São João da Barra. It is well-known among botanists (Santos, 2016; Sleumer, 1954; Smith, 1966; Wurdack, 1970) that for numerous specimens Glaziou reported localities where he never was, and included Amazonian collections in his herbarium citing them as collected by him in Rio de Janeiro. Therefore, although there is no direct evidence that *Glaziou 9887a* has not been collected in São João da Barra, I assume that this specimen has a label reporting an erroneous collection locality, and I conclude that this taxon does not occur in the state of Rio de Janeiro. Schumann (1889, p. 253) under *Limnosipanea spruceana* cited the collection "ante Pirapora: Pohl n. 3326" without citing the Brazilian state. Most likely, this gathering was collected near the town of Pirapora, state of Minas Gerais, and, as far as I know, is the only collection of this species from that state, and is the southernmost record of this specie. The nearest collection, *B.M.T. Walter et al. 1956* (CEN), is from the Municipality of Uru-açu, 14°22'S, 49°1'W, state of Goiás, and was made in the area that is now inundated by the dam of Serra da Mesa. Therefore, particular attention is given to the gathering *Pohl 3326*. Pohl (1976, Portuguese translation), in his travel diary reports that he was at the "Fazenda Pirapora", now the town of Pirapora, state of Minas Gerais, on
20–21 Jul. 1820, however he did not report the collection of any specific plant.

There are three sheets of Pohl 3326 at W. Sheet with accession number W-0072339, has eight plants mounted on it and one label with the heading "HERB. MUSEI PALAT. VINDOB. (Botanische Abtheilung des K. K. Naturhist. Hofmuseums Wien.)" with the handwritten text "3326, Limnosipanea sp., Thurnheisera galioides, Brasilia. Pohl." Specimen with accession number W-0072340, has ten plants mounted on it and one label with the printed heading "HRB. MUSEI PALAT.

VINDOB. (Botanische Abtheilung des K. K. Naturhist. Hofmuseums Wien.)" and the handwritten text "3326, ante Pirapora, Pohl." Specimen with accession number W-0072337 has thirteen plants mounted on it and three labels, without the heading of the Wien Museum. The label at the bottom center has the text handwritten by Schumann "Limnosipanea Spruceana HKpt [? sic]" and the stamp "determ. C. Schumann". The label at the bottom right corner has the handwritten text "3326, Limnosipanea sp., Hb. Bras., Thurnheisera galioides, Ante Pirapora. Pohl." The third label, positioned above the other

<figure><img src="image_24.png" /><figcaption>**Figure 20.** Distribution of *Limnosipanea spruceana* var. *spruceana* (circles) and *L. spruceana* var. *macrantha* (star).</figcaption></figure>
two labels, also has the stamp "determ. C. Schumann" (with "determ." stroke through), and the text, handwritten by Schumann, "Hiervon wären einige Stengel für das Berliner Herbar sehr erwünscht", which means "From this one [the sheet] some stems [plants] would be very appreciated by the Berlin Herbarium" (English translation by Bruno Wallnöfer). In other words, Schumann asked to extract some of the plants from this sheet to be deposited in the herbarium in Berlin [now B]. But apparently he didn't receive any, because nothing was detached from the sheet, in which case this would have been annotated by the herbarium curators on that sheet (interpretation of the text and observations by Bruno Wallnöfer).

At F there is one specimen of *Pohl* 3326, Acc. No. 869868, with a single plant preserved in an envelope

glued on the sheet. This sheet has a label with the printed heading "HERB. MUSEI HIST. NATUR. VINDOBON. (Botanische Abtheilung des K. K. Naturhist. Hofmuseums Wien.)" indicating that this specimen was extracted from one of the sheets at W. Such label also has the handwritten text (author unknown) "Limnosipanea Spruceana, Pohl 3326, Brazil."

3-3b. *Limnosipanea spruceana* Hook.f. var. *macrantha* Delprete, var. nov. (Figures 20-21).

Type: BRAZIL. Mato Grosso: Mun. Nova Xavantina, Drainage of the Upper Rio Araguaia, periodically flooded campo, ca. 30 km S of Xavantina

<figure><img src="image_25.png" /><figcaption>**Figure 21.** Flower comparison of *Limnosipanea spruceana* var. spruceana and var. *macrantha*. **A–B.** *L. spruceana* var. spruceana. **A.** Flower in anthesis. **B.** Detail of hypanthium and calyx. **C–D.** *L. spruceana* var. *macrantha*. **C.** Flower in anthesis. **D.** Detail of hypanthium and calyx. **A–B:** drawn from *Marimon & Delprete BS-976* (CAY); **C–D:** drawn from Irwin et al. 17014 (NY). Illustration by Piero Delprete.</figcaption></figure>
[ca. 14°54'31"S, 52°16'36"W], ca. 400 m, herb with ascending stems to ca. 20 cm long, 12 Jun. 1966 (fl), H.S. Irwin, R. Souza, J.W. Grear & R. Reis dos Santos 17014 (holotype UB [Acc. No. 43389]; isotypes F [Acc. No. 1677438], IAN [Acc. No. 130279], MO [Acc. No. 3172561], NY, RB [Acc. No. 145574], US).

**Diagnosis:** *Limnosipanea spruceana* Hook.f. var. *macrantha* Delprete differs from the typical variety by the aerial portion of the stem glabrous (vs. appressed- strigose to appressed-pubescent or glabrous in var. *spruceana*); hypanthium ellipsoid, 1-1.1 x 0.5-0.7 mm (vs. narrowly ellipsoid to narrowly obovoid, 0.5-1 x 1-1.3 mm); corollas 8.5-11.2 mm long (vs. 3-6(-7) mm long), with tube 3.5-4.7 mm long (vs. 2-3(-3.5) mm long) and lobes oblong to oblong-ovate, 4.5-6.5 x 1.5- 2.2 mm (vs. ovate to oblong-ovate, 1-3(-3.5) x 0.5-0.7 mm); filaments 1.3-1.8 mm long (vs. 0.4-0.8 mm long); anthers narrowly ellipsoid, 1.0 x 0.4 mm (vs. oblong, 0.5-0.7 x 0.2-0.4 mm); and style 6.5-9 mm long (vs. 2.5-5 mm long). (Figure 21).

**Description:** *Herbs*, semi-aquatic (amphibious) or aquatic, erect or decumbent (when the submerged part of the stem laying on the ground after the water receded), single-stemmed; stem 35-50 cm long, submerged portion up to 34 cm long, glabrous (laying on the ground after the water has receded), aerial portion 10-15 cm long, glabrous. *Leaves* heterophyllous, dimorphic; *submerged leaves* 3-6(-8) per node, with blades linear to acicular, 5-14 x 0.2-0.5 mm, glabrous, without secondary veins; *aerial leaves* 3-5 per node, with blades linear, narrowly lanceolate to lanceolate, 4-6.5 x 1-2 mm, glabrous, with 1-2 secondary veins each side. *Inflorescence* 2-8.5 cm long, simple, not branched, with 1-3 flowers, or 2- or 3-chotomously branched, with 3 to many flowers, with one flower at the base of each bifurcation; bracteoles subtending flowers narrowly lanceolate to linear, 2.2-3.5 x 0.1-0.2 mm, glabrous, or absent. *Hypanthium* ellipsoid, 1-1.1 x 0.5-0.7 mm, glabrous. *Calyx* lobes narrowly lanceolate to linear, 2.3-3.2 x 0.3-0.6 mm, acute at apex, glabrous throughout, with 1, narrowly triangular, 0.1 mm long, colleter at each lobe sinus. *Corolla* hypocrateriform, 8.5-11.2 mm long, tube white, lobes pale reddish-violet, turning white after anthesis; tube narrowly cylindrical, flaring near the mouth, 3.5-4.7 mm long; lobes oblong to oblong-ovate, 4.5-6.5 x 1.5-2.2 mm, obtuse to acute at apex, glabrous on both sides. *Filaments* 1.3-1.8 mm long; anthers narrowly ellipsoid, 1.0 x 0.4 mm. *Style* exserted 3-5 mm beyond corolla mouth, 6.5-9 mm long, glabrous; style

branches oblong to narrowly oblanceolate, 0.6 mm long. *Capsules* and *seeds* unknown.

**Distribution and ecology:** Only known from the type collection from a locality ca. 30 km S of Nova Xavantina, Mato Grosso, Central Brazil (Figure 20), growing in seasonally inundated campos at ca. 400 m altitude (Brazilian Shield).

**Phenology:** Specimens with flowers were collected in June.

**Suggested conservation status:** Critically Endangered (CR) or Extinct (EX). Only known by the type collection made in 1966 at ca. 30 km S of Nova Xavantina, Mato Grosso, Central Brazil. The number of individuals of the population is unknown. The natural vegetation of this area has been destroyed by agriculture, mostly for establishment of soybean plantations. There is no recent field study to check if this variety is still extant or not. Considering that the area of occurrence is highly impacted by agricultural practices, this variety might be either Critically Endangered or Extinct following IUCN criteria (IUCN 2012, 2019).

## Excluded species

*Limnosipanea parviflora* Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 11(5): 220. 1936

**Type:** BRAZIL. Minas Gerais: Without locality, s.d. [1816-1821] (fl), *A. Saint-Hilaire B2-2141* (holotype P [barcode P02285103]; isotype fragment F; photo-P at BR).

(≡) *Perama parviflora* (Standl.) J.H. Kirkbr. & Steyerm. in Steyermark & Kirkbride in Brittonia 29(2): 195. 1977

This species was originally positioned in *Limnosipanea* by Standley (1936). In agreement with Steyermark and Kirkbride (1977), because of the bilobed calyx, included stamens, clavate stigma, and hemilenticular seeds, among other characters, it is correctly placed in the genus *Perama* Aubl.

## 4. MAGUIREOTHAMNUS

*Maguireothamnus* Steyerm., Mem. New York Bot. Gard. 10: 220. 1964.

Steyerm. in Lasser & Steyermark, Fl. Venezuela 9(1): 308- 318, fig. 50. 1974; Delprete & Steyerm. in Steyermark et
al., Fl. Venez. Guayana 8: 642–644, figs. 504–506. 2004.
Type: *Maguireothamnus speciosus* (N.E. Br.) Steyerm.

*Shrubs. Stipules* with a basal sheath adnate with the petioles, free portion narrowly triangular to linear, sometimes with an apical colleter, subcoriaceous to coriaceous, with a row of 3–7 colleters on each side, inserted just below the margin (internal side) or on the margin of the basal sheath. *Leaves* opposite, sessile, subsessile or short-petiolate; blades ovate, obovate, oblong, oblanceolate to lanceolate, thinly to thickly coriaceous; domatia absent. *Flowers* axillary, solitary, 5-merous, homostylous, subtended by two leaf-like bracts, with a fragrance reminiscent of gardenia. *Hypanthium* oblong-obconical, obconical, narrowly obconical to narrowly obovoid, glabrous, minutely puberulent, densely pubescent, antrorse-hispidulous, antrorse-hispid to densely spreading strigose. *Calyx* tube extremely reduced or absent; lobes large, subequal or unequal, elliptic, narrowly-elliptic, oblong, narrowly lanceolate to linear, persistent (eventually braking off in very old capsules), with 5–7 colleters inside each lobe sinus. *Calycophylls* absent. *Corolla* hypocrateriform, actinomorphic, white to cream-white; tube long, externally glabrous, internally puberulent, with a pubescent ring at orifice inside; lobes left-contorted, ovate, lanceolate to oblong, margin entire, obtuse, acute or short-acuminate at apex. *Stamens* included; filaments attached near the mouth of the corolla tube, inserted just below the mouth, short, equal; anthers sessile, elongate, dehiscing by longitudinal slits, dorsi-fixed near the base. *Pollen* 3(4)-colporate; exine smooth, not perforate, finely foveolate or finely and sparsely perforate. *Style* exerted well beyond the corolla mouth, terete, antrorse-papillose at distal portion; style branches elliptic, papillose on adaxial side. *Ovary* with a stalked placenta; stalk inserted at the middle of the septum, terminating with an oblong-ellipsoid extension; ovules inserted on the placental extension. *Fruit* capsular, dehiscing loculicidally, thinly woody, crowned by the persistent calyx lobes. *Seeds* many per locule, vertical, laterally compressed, reddish-brown, irregularly fusiform to rhomboid in outline, margin irregularly fringed; testa reticulate-foveolate.

**Distribution and ecology:** A genus endemic to the tepuis of the Guiana Shield, in Venezuela, Guyana and northern Brazil (Roraima); growing on exposed sandstone outcrops, open scrub formations, *Bonnetia roraimae* forests, shrubby savannas, and open wet savannas, on sandy soils, at 1300–2600 m altitude. Three species are here recognized in the genus. The flowers have a fragrance reminiscent of gardenias and are most likely polinated by hawk-moths.

Key to *Maguireothamnus* species

1. Leaf blades 1–3.5 × 0.3–2 cm; capsules hispidulous to hispid (Venezuela: Amazonas: Cerro Duida, Cerro Sipapo, Cerro Huachamacari, Cerro Marahuaca-Atuhua-Shiho).................... 3. *M. tatei*

1. Leaf blades 2.5–6.5 × 0.4–2.5 cm; capsules glabrous (vestiture of densely pubescent form of *M. jauaensis* unknown, probably densely pubescent).................................................... 2

2. Leaf blades glabrous throughout; hypanthium obconical, narrowly obovoid to oblong-obovoid, 5.5–6 × 3.5–5 mm, glabrous or minutely puberulent; corolla lobes oblong, oblong-elliptic, or lanceolate to liner-lanceolate, 2.3–5 × 0.7–1.5(–2) cm, acute to acuminate at apex (Guyana, and Venezuela: Bolívar: Auyan-tepui, Ptari-tepui, Chimantá Massif, Angasina-tepui, Mount Roraima, Carrao-tepui, Churi-tepui) .................................................................... 2. *M. speciosus*

2. Leaf blades densely antrorse-pubescent throughout, glabrous above and densely antrorse-pubescent below, or glabrous throughout; hypanthium oblong-obconical, 4–5 × 2.5–3 mm, densely pubescent or glabrous; corolla lobes oblong-ovate to obovate, 2–2.5 × 0.8–1.1 cm, round or obtuse at apex (Venezuela: Bolívar: Cerro Jáua) .................................................... 1. *M. jauaensis*

4-1. *Maguireothamnus jauaensis* (Steyerm.) Steyerm. in Lasser & Steyermark, Fl. Venez. 9(1): 311. 1974. (Figures 22–23).

(=) *Maguireothamnus speciosus* subsp. *jauaensis* Steyerm. in Mem. New York Bot. Gard. 23: 883. 1972.

Type: VENEZUELA. Bolívar: Cerro Jáua, summit of the Central-Occidental portion, 60 km NE of Sanidad Mission Camp of Río Canaracuni, 4°45′N, 64°26′W, 1922–2100 m, 22–27 Mar. 1967 (fl), *J.A. Steyermark* 97837 (holotype VEN [Acc. No. 71474]; isotypes NY [barcode 00132137], U [barcode 0006070]).

(=) *Maguireothamnus jauaensis* (Steyerm.) Steyerm. var. *breweri* Steyerm. in Lasser & Steyermark, Fl. Venez. 9(1): 311. 1974

Type: VENEZUELA. Bolívar: Meseta de Jáua, Cerro Jáua, selva de galería al borde del tributario del Río Marajano, in savanna north of campsite, 4°48′N, 64°34′W, 1750 m, 4 March 1974 (fl), *J.A. Steyermark*, *V. Carreño Espinoza & C. Brewer-Carías* 109765 (holotype VEN [Acc. No. 97817]).

*Shrubs*, 0.5–2 m tall; internodes of young branchlets densely antrorse-pubescent or glabrous. *Stipules* basal
<figure><img src="image_26.png" /><figcaption>**Figure 22.** *Maguireothamnus jauaensis*. A. Branch with one flower in anthesis. B. Stipule, inner side, with three colleters on each side. C. Longitudinal section of flower in anthesis. D. Longitudinal section of ovary. E. Capsule. A–D: drawn from *Huber 12981* (US). Illustration by Piero Delprete.</figcaption></figure>
sheath glabrous or puberulent outside, appressed puber-
ulent inside; free portion narrowly triangular, gradu-
ally tapering at apex, 6.3–10 × 2–4 mm, subcoriaceous,
densely antrorse-pubescent or glabrous outside, glabrous
inside; with a row of 3–4 colleters on each side, inserted
just below the margin (internal side) of the basal sheath.
Leaves erect, imbricate, sessile or subpetiolate; petioles
to 1.5 mm long, densely antrorse-pubescent; blades lan-
ceolate-elliptic or narrowly elliptic, 2.5–4.5 × 0.4–1 cm,
attenuate to acute at base, acute at apex, coriaceous, dark
green above, pale green below, drying olive-green (or
brown, when preserved in alcohol before drying), dense-
ly antrorse-pubescent throughout, or glabrous above and
densely antrorse-pubescent below, or glabrous through-
out; margins strongly revolute; secondary and tertiary
venation submerged within the lamina. *Bracts* subtend-
ing flowers lanceolate to oblong-lanceolate, 15–30 × 5–6

mm, acute at apex, subcoriaceous, glabrous or sparsely
pubescent, margins glabrous. *Flowers* with pedicels 2–4
mm long, sparsely pubescent or glabrous. *Hypanthium*
oblong-obconical, 4–5 × 2.5–3 mm, glabrous or densely
pubescent (fide Steyermark, 1974: 311). *Calyx* lobes 5,
pale green, oblong-lanceolate, 14–23 × 4–5 mm, acute at
apex, subcoriaceous, glabrous, margins glabrous, with
5–6, narrowly-lanceolate, 0.5–0.6 mm long, colleters at
each lobe sinus. *Corolla* 10–13 cm long; tube pale yel-
lowish-green, long-cylindrical, slightly wider at mouth
(anther portion), 9–10.5 cm long, 2–3 mm wide for
most length, 3.5–5 mm wide near the mouth, glabrous
outside, completely glabrous or basal portion puberu-
lent, the rest glabrous from base to 2–2.2 cm below the
mouth and villous above inside; lobes white, oblong-
ovate to obovate, 2–2.5 × 0.8–1.1 cm, round or obtuse at
apex, glabrous outside, glabrous (microscopically papil-

<figure><img src="image_27.png" /><figcaption>Figure 23. Distribution of *Maguireothamnus jauaensis* (star), *M. speciosus* (circles), and *M. tatei* (squares).</figcaption></figure>
lose) inside. *Anthers* subsessile, elongate, 6.5–7 × 0.8–1 mm, base round, apex acuminate. *Style* 10.5–12 cm long, exserted 10.5 mm beyond corolla mouth, glabrous for most of the length and papillose at distal 1.8–2 cm below style branches; style branches broadly ovate-spathulate, 2 mm long. *Capsules* obovoid, 1.5 × 1.1 cm, acute at base, round at apex, dark brown, glabrous (vestiture of densely pubescent form unknown). *Seeds* unknown.

**Taxonomic observations:** Steyermark (1974: 311–312) described *Maguireothamnus jauaensis* var. *breweri* Steyerm. and noted that most of its vegetative parts are densely pubescent while the calyx lobes are glabrous. He differentiated it from the typical variety by the leaves, branches, and stipules densely pubescent (vs. leaves glabrous above, branches and stipules glabrous), corolla lobes "0.7–1.3 × 0.3–0.4 cm" (vs. 2.5–3.5 × 0.7–1.6 cm; see comments below), round at apex (vs. acute to subobtuse at apex). On the other hand, he observed that (translated from Spanish) "All intermediary populations of *M. jauaensis* show the same characters that could be useful to separate this taxon from the other species of the genus, as the stipules narrowly triangular tapering towards the apex, the interior of the corolla throat glandular-puberulent at lower portion, and the well-developed peduncle." In the same page, he reported the presence of intermediary populations between the two varieties, with vegetative and reproductive parts less pubescent, and corolla throat either glandular-puberulent or glabrous at base inside.

Steyermark (1974: 311) described the corolla lobes of *Maguireothamnus jauaensis* var. *breweri* as 0.7–1.3 × 0.3–0.4 cm, and round at apex. This variety is known only from the holotype specimen at VEN. The specimen *Steyermark et al. 109765* has one flower at the tip of a small branch mounted on the sheet, and two loose flowers included in the attached envelop. All the corollas lobes of this specimen are completely wrinkled-up, and is impossible to know their real length when fully expanded. Most likely Steyermark's measurements were made on these wrinkled-up lobes; therefore, the corolla lobes dimensions supplied by Steyermark for this variety are unreliable.

Steyermark (1974: 312) indicated that *Maguireothamnus jauaensis* var. *breweri* is similar to *M. tatei* because of the pubescent hypanthium and midrib ending at leaf apex, the paler or silvery color of the leaves below and the expanded leafy portion of the branches; and is similar to *M. speciosus* because of the glabrous calyx lobes. Delprete and Steyermark (2004c: 643) treated *M. jauaensis* var. *breweri* as a synonym of *M. speciosus* ssp. *jauaensis*, without further comments; however, recent studies, with more material available,

allowed me to reach a different conclusion. As already pointed out by Steyermark, populations on Cerro Jáua are extremely variable in terms of vestiture of vegetative and reproductive parts. On the type specimens of var. *jauaensis* (*Steyermark* 97837), the leaves are glabrous throughout, while in the type specimen of var. *breweri* (*Steyermark et al. 109765*) the leaves are densely antrorse-pubescent throughout. However, two collections from the same locality (*Huber 12981* and *Steyermark et al. 109333*) have intermediate characters, as their leaves are glabrous above and densely antrorse-pubescent below, and their corolla throats are either glabrous (*Huber 12981*) or glandular-puberulent at lower portion (*Steyermark et al. 109333*). Therefore, all the above-cited collections and the names involved are here treated as synonymous and as part of a highly variable species endemic to Cerro Jáua.

**Distribution and ecology:** Known by four gatherings from Cerro Jáua (now part of the Jaua-Sarisariñama National Park), Bolívar, Venezuela, at 1750–2100 m altitude (Figure 23).

**Phenology:** Flowering specimens were collected in February, March and November; those collected in November have flowers in anthesis and mature capsules.

**Suggested conservation status:** Critically Endangered (CR). This species is endemic to Cerro Jáua, a tepui included in the Jaua-Sariñama National Park, Bolívar state, Venezuela. It is known from four collections made in 1967, 1974 and 1989, at 1750–2100 m altitude. The size of the populations is unknown. The label of the type of *Maguireothamnus speciosus* ssp. *jauaensis* (*Steyermark* 97837) reports that the specimen was collected on the summit of the Central-Occidental portion of Cerro Jáua, at 1922–2100 m, with coordinates "4°45′N, 64°26′W". However, these coordinates are erroneous, because they fall outside Cerro Jáua. Therefore, this locality is eliminated from the EOO calculation. Using the three collections localities within Cerro Jáua, the EOO of this species is of 1.89 km². Taking into account the extremely small area of occurrence, which is inside a national park, this species is treated as Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA. Bolívar: Dis- to. Cedeño, Meseta de Jáua, Cerro Jáua, sector centro- meridional, cabeceras del Río Marajano, afluente del Río Cácaro, 4°48′N, 64°32′W, 1750–1800 m, 20 Nov. 1989 (fl, imm fr), *O. Huber 12981* (US, VEN); Meseta de Jáua, Cerro Jáua, cumbre, porción SW, selva al borde del Río
Marajano, 4°48'50"N, 64°34'10"W, 1750-1800 m, 22 Feb. 1974 (fl), *J.A. Steyermark et al. 109333* (NY, US, VEN).

4-2. *Maguireothamnus speciosus* (N.E.Br.) Steyerm.,
Mem. New York Bot. Gard. 10: 223. 1964. (Figures
4G-H, 23-24).

(=) *Chalepophyllum speciosum* N.E.Br. in Trans. Linn.
Soc. London ser. 2, 6: 33, pl. 5, figs. 10-17. 1901.

Type: VENEZUELA. Bolívar: *F.V. McConnell & J.J. Quelch 653* (first-step lectotype, designated by Steyermark (1964: 223), **second-step lectotype** K [without barcode, with label "New York Botanical Garden Neg. No. N.S. 3240 (1953)"]**, here designated**; isotype BM [barcode BM000614299]; photo-K at NY).

(=) *Chalepophyllum coriaceum* Gleason in Gleason &
Killip, Brittonia 3: 192. 1939.

Type: VENEZUELA. Bolívar: Mount Auyan-Tepui
[ca. 5°54′N, 62°32′W], 2200 m, Dec. 1937 (fl, fr),
G.H.H. Tate 1179 (holotype NY [2 sheets, barcode
00131038]; isotype VEN [not found]).

Shrubs 0.5-3 m tall, erect or rarely partially pro-
strate; bark gray; internodes of young branchlets glabrous.
*Stipules* basal sheath glabrous or puberulent outside,
sparingly to densely sericeous inside; free portion narrow-
ly triangular, gradually tapering at apex or broadly trian-
gular at base and aristate at apex, 2-10 x 2-3 mm, some-
times with an apical colleter, subcoriaceous to coriaceous,
glabrous (rarely strigose) outside, sericeous throughout or
only at base inside; with a row of 5-7 colleters on each
side, inserted near the margin on internal side of the
basal sheath. *Leaves* erect and imbricate, sessile or short
petiolate; petioles (when present) to 4 mm long, glabrous;
blades oblong, obovate to oblanceolate, 2.5-6.5 x 0.8-2.5
cm, obtuse to acute at base, obtuse, round to acute at
apex, coriaceous, dark green and shiny above, pale green
below, drying olive-green, glabrous throughout; margin
often minutely revolute; secondary and tertiary venation
submerged within the lamina, sometimes with 2-3 sec-
ondary veins on each side of midrib, barely visible above.
*Bracts* subtending flowers round, or broadly to narrowly
obovate, (0.7-9.5-1.6 x (0.3-0.6-1 cm, round at apex,
coriaceous. *Flowers* with pedicels 2-5(-8) mm long. *Hyp-
anthium* obconical, obovoid to oblong-obovoid, 5.5-6 x
3.5-5 mm, glabrous or minutely puberulent. *Calyx* lobes
oblong-ovate, narrowly-elliptic to linear, 10-32 x 2-7
mm, acute to acuminate at apex, subcoriaceous, glabrous
or with sparse hairs at apex, with 5-7, narrowly-lanceo-

late to oblong-lanceolate, 0.5-0.7 mm long, colleters at
each lobe sinus. *Corolla* 9-17 cm long; tube pale green,
sometimes reddish at base, long-cylindrical and slightly
wider near the mouth (anther portion), (6.5-)8-13 cm
long, 2-4 mm wide for most length, 6-7 mm wide near
the mouth, glabrous outside, glabrous from base to 1-1.5
cm from the mouth and densely pubescent to villous
above inside; lobes white to cream-white, with one mar-
gin pink to brick-colored, oblong, oblong-elliptic, lan-
ceolate to liner-lanceolate, 2.3-5 x 0.7-1.5(-2) cm, acute
to acuminate at apex, glabrous outside, pubescent at base
inside. *Anthers* subsessile, elongate, 6-8 x 0.8-1.1 mm,
base round, apex acuminate. *Style* 7.5-15 cm long, exsert-
ed 15-20 mm beyond corolla mouth, glabrous for most
of the length and papillose on the distal portion; style
branches oblong-spathulate, 2.5-3 mm long. *Capsules*
ovoid to ellipsoid, 1.5-2.7 x 0.8-1.5 cm, obtuse to acute
at base, round at apex, dark brown, glabrous. *Seeds* 4-5 x
2-3 mm.

**Taxonomic observations:** Brown (1901: 33) described *Chalepophyllum speciosum* and cited three syntypes “*McConnell & Quelch 100, 305, 653.*” Steyermark (1964: 223) cited “*McConnell & Quelch 653* (photo of holotype of *Chalepophyllum speciosum*, from K at NY.” Although Steyermark cited this collection as holotype, according to According to Art. 7.11 and 9.10 of the Code (Turland et al. 2018) is a correctable error, and it is here treated as an inadvertent first-step lectotypification of *McConnell & Quelch 653* at K because there are two specimens of *McConnell & Quelch 653* at K. At the time they were studied, these specimens did not have barcodes. The specimen bearing the label “New York Botanical Garden Neg. No. N.S. 3240 [1953],” which has eight branchlets with flowers is here designated the second-step lectotype of this taxon. The other specimen with three branchlets with flowers, and without the NYBG label, is an isolectotype.

Leaf shapes, corolla dimensions, and vestiture are
variable within populations of *Maguireothamnus* species.
Steyermark (1972b) recognized two subspecies under *M.
speciosus*, and distinguished ssp. *jauaensis*, endemic to
Cerro Jáua, by the lower half of the corolla tube puberu-
lent inside (vs. lower half glabrous inside in the typi-
cal subspecies), among other characters. Two years later,
Steyermark (1974) elevated *M. speciosus* subsp. *jauaensis*
Steyerm. to species level. Delprete and Steyermark (2004c:
642-643) returned the taxon to subsp. *jauaensis*, recog-
nizing two subspecies in *M. speciosus*. However, recent
studies with more material available showed that the spec-
imens from Cerro Jáua are distinct from *M. speciosus* and
are here treated as a distinct species (see above).
<figure><img src="image_28.png" /><figcaption>**Figure 24.** *Maguireothamnus speciosus*. A. Branch with capsules, flower buds, and flowers in anthesis. B. Node with stipule and partially exserted colleters. C. Distal portion of style with partially opened lobes. D. Distal portion of corolla tube, base of corolla lobes, and included stamens. E. Seeds with detail of reticulated testa. F. Longitudinal section of hypanthium, ovary, and calyx, with internal colleters. Illustration by Bruno Manara (Reproduced with permission from Flora de Venezuela vol. 9(1), fig. 50. 1974).</figcaption></figure>
**Distribution and ecology:** Endemic to the tepuis of the Guayana Shield, in Guyana and Venezuela, state of Bolívar (Auyan-tepui, Ptari-tepui, Chimantá Massif, Angasina-tepui, Mount Roraima, Carrao-tepui, and Churi-tepui), and the Brazilian contiguous state of Roraima (Mount Roraima and Serra do Sol) (Figure 24). Growing on exposed sandstone outcrops, open scrub formations, *Bonnetia roraimae* forests, shrubby savannas, and open wet savannas, on sandy soils, at 1300–2600 m altitude.

**Phenology:** Flowering specimens were collected in January, February, May, and August; specimens with flowers and fruits in May and August.

**Suggested conservation status:** Least Concern (LC). This species occurs on numerous tepuis, at 1300–2600 m altitude, present in Guyana, Venezuela (state of Bolívar), and the contiguous Brazilian state of Roraima. In Venezuela, it has been collected on the Auyan-tepui, Ptari-tepui, Chimantá Massif, Angasina-tepui, Mount Roraima, Carrao-tepui, and Churi-tepui. Some of these tepuis are included within the Canaima National Park. On Mount Roraima, which also is within the Canaima National Park, where the triple border between Venezuela, Guyana and Brazil is located. This species has been collected in the three countries. Taking into account the numerous collection sites, and the large extent of occurrence, including many remote, preserved areas, this species is treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA: Bolívar: Altiplanicie of Auyan-tepui, SW Sector, along small river, headwaters of Río Churún, high tepui scrub and low tepui forest along river and among sandstone outcrops, 5°48′N, 62°33′W, 1850 m, 23–25 Apr. 1996 (fl), W.R. Andersson & O. Huber 13856 (MO, NY [2 sheets], VEN); Auyantepui, 2100 m, Jan. 1949 (fl), F. Cardona 2669 (US); Dto. Roscio, Mun. Dalla Costa Roraima [ca. 7°18′N, 61°30′W], 270–300 m, 22 Aug. 1982 (fl, fr), A. Castillo 1811 (U, VEN); Summit of Cerro Roraima, rock outcrops and pool, shrub in wet areas, 29–30 Sep. 1982 (fl, fr), S.E. Clemants 2459 (NY); parte central del plató de Auyantepui, 1900 m, Apr. 1956 (fl), E. Foldats 2603 (F); Valle Encantado, lado derecho del Salto Angel [ca. 5°58′N, 62°32′W], Auyantepuy, 15 Aug. 1968 (fl, imm fr), E. Foldats 7108 (FT); Disto. Piar, Auyan-tepui summit, E edge of massif, 5°53′N, 62°26′W, 1740 m, open area with large flat rocks and low vegetation, some patches of *Bonnetia roraimae*, 27 May 1986 (fl, fr), B.K. Holst 2994 (MO, NY, U, US); Disto. Piar, Ptari-tepui, summit,

5°46′N, 61°47′W, 2300 m, nearly flat summit with much exposed sandstone, patches of low vegetation and shrub islands with *Bonnetia roraimae* dominant, 27 May 1986 (fl, fr), B.K. Holst 3596 (MO, US); Disto. Piar, Macizo del Chimantá, sector centro-noroeste del Chimantá-tepui, cabeceras orientales del Caño Chimantá, 5°18′N, 62°9′W, 2000 m, 26–29 Jan. 1983 (fr), O. Huber & J.A. Steyermark 6863 (US, VEN); Disto. Piar, altiplanicie del Auyan-tepui, Sector SSW de la meseta, al W de la cumbre Sur, vegetación sobre rocas de arenisca y arbustales, 5°48′N, 62°32′W, 2070 m, 27 Aug. 1983 (fr), O. Huber et al. 8055 (NY, US, VEN); Disto. Piar, altiplanicie del Auyan-tepui, sector SE, ca. 5 km al ESE del Churún-merú (Salto Churún), 5°51′N, 62°30′W, 10 Dec. 1983 (fl, fr), O. Huber & E. Medina 8528 (MO, NY, VEN); Disto. Piar, Macizo del Chimantá, altiplanicie meridional del Abacápá-tepui, ubicada en el sector SW del Macizo, 5°10′N, 62°16′W, 2200 m, 31 Jan.–2 Feb. 1984 (fl), O. Huber & N. Dezzeo 8602 (MO, NY, VEN); Disto. Piar, Macizo del Chimantá, sección septentrional del Apacará-tepui, en el extrem N del macizo, 5°25′N, 62°11′W, 2350 m, 3–5 Feb. 1984 (fl), O. Huber & M. Colella 8703 (MO, NY, VEN); Disto. Roscio, Cumbre del Yuruaní-tepui, ca. 12 km al NNE del Kukenán-tepui, 5°19′N, 60°51′W, 2200 m, 29 Feb. 1984 (fl), O. Huber 9082 (NY, VEN); Distto. Roscio, Kukenán-Tepui, cumber del sector más septentrional algo separado del macizo principal, vegetación herbaceo-arbustiva sobre rocas de arenisca abiertas y alrededor de lagunas, 5°16′N, 60°48′W, 2500 m, 28 Apr. 1984 (fl), O. Huber 9436 (NY, US, VEN); Distto. Piar, altiplanicie ubicada en la sección más septentrional del Brazo Occidental del Auyantepui, ca. 25 km al SE de Canaima, 6°6′N, 62°43′W, 1650 m, 13 Nov. 1984 (fl), O. Huber 9720 (NY, US, VEN); Disto. Piar, Cumbre del Ptari-tepui, ca. 22 km al NNW de la Misión de Kavanayén, 5°547′N, 61°47′W, 2400 m, 19 Nov. 1984 (fl), O. Huber 9822 (NY, VEN); Disto. Piar, altiplanicie del Auyan-tepui, sector N del Brazo Oriental, a lo largo de un valle afluente (derecho) del Río Churún, 6°1′N, 62°26′W, 1900 m, 17 Jan. 1986 (fl), O. Huber 11181 (NY, VEN); Disto. Piar, cumbre del Angasina-tepui (Adanta-tepui), cabeceras altiplanicie poco inclinada hacia el NE con saban arbustiva en el sector Nor-occidental, ca. 40 km al WNW de la Misión de Wonkén, 5°3′N, 62°7′W, 2100 m, 4 Mar. 1986 (fl, fr), O. Huber 11362 (NY, US, VEN); Kukenan-tepui, summit, large open rocky areas, boggy areas, small patches of forest, 5°13′N, 60°18′W, 2550 m, 10 Apr. 1988 (fl), R. Liesner 23047 (MO, NY); Disto. Piar, Macizo del Chimantá, setor SSE, altiplanicie sur-oriental del Acopán-tepui, cabecera del Río Arauác, praderas húmedas y arbustales enanos sobre turberas, 5°11′N, 62°0′W, 1920 m, 14–16 Feb. 1984 (fl), J.L. Luteyn et al. 9475 (NY, U,
US, VEN); Disto. Piar, Macizo del Chimantá, sector SE, amplia altiplanicie en la sección nor-oriental del Acopán-tepui, levemente inclinada hacia el NE, en las cabeceras del Río Yunék, 5°12'N, 62°5'W, 1950 m, 8–11 Feb. 1985 (fl), *J.J. Pipoly et al.* 7210 (MO, NY, VEN); Disto. Piar, Macizo del Chimantá, sector centro-meridional, Amplio valle ubicado entre el borde nor-oriental del Torono-tepui y la sección central del Chimantá-tepui, drenando hacia el S, 5°16'N, 62°9'W, 2100 m, 11–15 Feb. 1985 (fl), *J.J. Pipoly et al.* 7245 (MO, NY, VEN); Plateau of Auyan-tepui, SW section of plateau, W of southern summit, 5°48'N, 62°32'W, 2070 m, 27 Aug. 1983 (fl, fr), *G.T. Prance & O. Huber* 28243 (MO, NY, US, VEN); Plateau of Auyan-tepui, Central Northern section of Eastern Branch, 5°57'N, 62°25'W, 1940 m, 27 Aug. 1983 (fl), *G.T. Prance & O. Huber* 28272 (FT [2 sheets], NY); Mun. Gran Sabana, Chimantá Massif, SSE sector, SE summit of Acopán-tepui, headwaters of Río Arauák, 5°11'N, 62°0'W, 1920 m, 6–8 Apr. 1989 (fl), *J. Pruski & O. Huber* 3556 (NY); Mun. Gran Sabana, Chimantá Massif, summit of central NE portion of Murey(Eruoda)-tepui, in the NE sector of the massif, 5°23'N, 62°3'W, 2550 m, 1–3 Apr. 1989 (fl), *J. Pruski & O. Huber* 3582 (MO, NY); Chimantá Massif, central section, rocky and swampy savanna between W and E branches of headwaters of Río Tirica, 2120–2210 m, 11 Feb. 1955 (fl), *J.A. Steyermark & J.J. Wurdack* 743 (F, NY, US); Chimantá Massif, vicinity of Summit Camp along Río Tirica, 1925 m, 20 Feb. 1955 (fl), *J.A. Steyermark & J.J. Wurdack* 1004 (NY [2 sheets], US, VEN); summit of Mount Roraima, on NW portion N and NW of Summit Camp, 2620–2740 m, 27 Sep. 1944 (fl), *J.A. Steyermark* 58860 (IAN [frag], F, NY [2 sheets], US); summit of Carrao-tepui, 2470–2500 m, 7 Dec. 1944 (fl), *J.A. Steyermark* 60896 (F); Ptari-tepui, *Bonnetia roraimae* forest on SW-facing shoulder, 2000–2200 m, 2 Nov. 1944 (fl), *J.A. Steyermark* 59740 (F, MO, NY); Ptari-tepui, S-facing slope, vicinity of Cave Rock, 1800–1900 m, 2 Nov. 1944 (fl), *J.A. Steyermark* 59801 (F, NY); Chimantá Massif, Bonnetia forest, NW part of Summit of Abácapa-tepui, 2125–2300 m, corolla tube pale green, corolla lobes white with one edge bordered rose, flowers very fragrant, 13 Apr. 1953 (fl), *J.A. Steyermark* 74881 (F, MO, NY [2 sheets]); Chimantá Massif, rocky plateau on SE-facing upper shoulder of Apacará-tepui, below summit, 2000–2100 m, 20 Jun. 1953 (fl bud), *J.A. Steyermark* 75781 (F, NY); Auyan-tepui, cumbre de la parte del brazo NE (division occidental del cerro), sabana de suelo de arenisca al S y alredor dela avioneta de Jimmy Angel, 1800 m, 7 May 1964 (fl), *J.A. Steyermark* 93501 (F, NY, P, U); Cerro Roraima, cumbre NE de Venezuela imediatamente al S del hito que marca los límites con Guyana, Brasil y Venezuela, El Valle, head-

waters of Río Arabapó, 5°12'N, 60°42'W, 2750–2800 m, 26 Aug-2 Sep. 1976 (fl), *J.A. Steyermark et al.* 112493 (F, NY, US, VEN); cumbre del Ptari-tepui, N de la Misión de Santa Teresita de Kavanayén, 5°45'N, 61°45'W, 2360–2420 m, 23 Feb. 1978 (fl), *J.A. Steyermark et al.* 115732 (F, MO); Dto. Sucre, Meseta de Jáua, cumber, sección oriental-central, afloramiento de piedra arenisca en sitios expuestos con vegetación herbácea y arbustos achaparrados, 4°35'N, 64°15'W, 2020 m, 14 Feb. 1981 (fl, fr), *J.A. Steyermark et al.* 124334 (NY); Dto. Piar, Macizo del Chimantá, sector centro-noreste del Chimantá-Tepui, cabeceras orientales del Caño Chimantá, 5°18'N, 62°9'W, 2000 m, 26–29 Jan. 1983 (fl), *J.A. Steyermark, O. Huber & V. Carreño E.* 128081-A (MO, NY, U, VEN); Dto. Piar, Macizo del Chimantá, altiplanicie en la base meridional de los farallomes superiors del Apacará-Tepui, sector N del macizo, 5°20'N, 62°12'W, 2200 m, 30 Jan–1 Feb. 1983 (fl), *J.A. Steyermark et al.* 128311 (MO, NY, U, VEN); Dto. Piar, Macizo del Chimantá, pequeñas altiplanicies en la base septentrional de los farallones superiors del Amurí-Tepui (Sector W del Acopán-Tepui), 5°10'N, 62°7'W, 1850 m, 2–5 Feb. 1983 (fl), *J.A. Steyermark et al.* 128668 (MO, NY, U, VEN); Dto. Piar, Macizo del Chimantá, cumbre sur-ocidental del Amurí-tepui (Sectow W del Acopán-tepui), small wooded area of dwarf forest bordering and below rochy open savanna, 2170 m, 6 Feb. 1983 (fl), *J.A. Steyermark et al.* 128765 (MO); Dto. Piar, cabeceras del affluent derecho superior del Río Tirica (Caño del Grillo), 5°18'N, 62°3'W, 2450 m, 7–9 Feb. 1983 (fl), *J.A. Steyermark et al.* 128916 (MO, NY, U, VEN); Dto. Piar, Macizo del Chimantá, sector SSE, altiplanicie sur-oriental del Acopán-tepui, cabeceras del Río Aruác, 1920 m, 14 Feb. 1984 (fl), *J.A. Steyermark et al.* 129900 (US, VEN); parte superior de Auyantepui, 2300 m, Apr. 1956 (fl), *V. Vareschi & E. Foldats* 4858 (F); Churi-tepui (Muru-tepui), NW cumbres, 2100–2200 m, 24 Jan. 1953 (fl), *J.J. Wurdack* 34178 (NY, P); Churi-tepui (Muru-tepui), NW cumbres, 2250–2300 m, 26 Jan. 1953 (fl), *J.J. Wurdack* 34209 (F, NY, U, US).

**GUYANA:** Cuyuni-Mazaruni Region, Mt. Maringma, summit, scrub forest and tepui bog on peat and sandstone, with *Bonnetia*, *Ilex*, and *Brochinia*, 5°13'N, 60°35'W, 2110 m, 15 Jun. 2004 (fl), *H.D. Clarke et al.* 11703 (MO, NY, US); Cuyuni-Mazaruni Region, 1 km N of tip of northern prow of Roraima, El Dorado Swamp, open scrub and low trees on white sand and boulders, 5°15'N, 60°35'W, 2000–2200 m, 25 Feb. 1989 (fl), *W. Hahn & D. Goupal* 5516 (BRG, MO, NY, US); Mazaruni-Potaro Region, Guyana-Venezuela border, Roraima summit, La Proa Camp, E of border, near Lake Gladys, mostly rocky areas with ravines and lake, 14 Apr. 1988 (fl), *R.L. Liesner* 23309 (MO); Upper Mazaruni District,
N slope of Mt. Roraima, 5°16′N, 60°43′W, 2300 m, 14 Feb. 1985 (fl), *J. Rentz* 14277 (U); Cuyuni-Mazaruni Region, Summit of Kamakusa Mountain (i.e., on top of 4th escarpment of four), impenetrable elfin forest to 3 m, extremely dense and wet, rich in epiphytes, with *Bonnettia* (2 spp.), *Brocchinia* cf. *tatei*, Malpighiaceae, Melastomataceae, Cyperaceae spp., *Weinmannia*, *Ilex* cf. *retusa*, 5°52′52″N, 6°6′10″W, 1686 m, 7 Jun. 2012 (fl), *E. Tripp* et al. 3186 (P, US); N ridge of Mt. Roraima, small swampy areas on gently sloping land, *Heliamphora*, Xyridaceae, Eriocaulaceae, soil humus-poor fine silty-sand, area windswept and often cloud-covered, with frequent rain, 5°15′40″N, 60°44′40″W, 2065–2430 m, 26 Mar. 1978 (fl), *J.F. Warrington* et al. 42 (E, K, U).

**BRAZIL. Roraima:** Summit of Serra do Sol (Ueitepui) [ca. 5°1′N, 60°36′W], 2300 m, 28 Dec. 1954 (fl, fr), *B. Maguire* & *C.K. Maguire* 40392 (IAN, NY [2 sheets], RB); Mt. Roraima (Guyana and N Brazil border), summit, 24 Nov. 1927 (fl), *G.H.H. Tate* 388 (NY); Mun. Boa Vista, Monte Roraima, substrato rochoso, 2600–2870 m, 5 Dec. 1983, *E.L. Sette* et al. 14 (MIRR); Monte Roraima, 9 Apr. 1973 (fl), *E.F. Terezo* 23 (IAN); Monte Roraima, 10 Apr. 1973 (fl), *E.F. Terezo* s.n. (INPA No. 60513); Felsenregion, 2500 m, Jan. 1910 (fl), *E. Ule* 8781 (L, MG).

4-3. *Maguireothamnus tatei* (Standl.) Steyerm., Mem. New York Bot. Gard. 10: 227. 1964. (Figures 23, 25).

(=) *Chalepophyllum tatei* Standl., Publ. Field Columbian Mus., Bot. Ser. 7 (4): 380. 1931.

Type: VENEZUELA. Amazonas: Mount Duida, summit of Peak N. 7, 2130 m, Aug. 1928–Apr. 1929 (fl, fr), *G.H.H. Tate* 623 (holotype NY [barcode 00131042]; isotype F [Acc. No. 603541, Negative No. 72096]).

(=) *Chalepophyllum latifolium* Standl., Publ. Field Columbian Mus., Bot. Ser. 7: 379. 1931.

(=) *Maguireothamnus tatei* (Standl.) Steyerm. var. *latifolium* (Standl.) Steyerm., Mem. New York Bot. Gard. 10: 228. 1964.

Type: VENEZUELA. Amazonas: Summit of Mount Duida (Cerro Yennamadi), [ca. 3°30′N, 65°37′W],

1320 m, Aug. 1928–Apr. 1929 (fl, fr), *G.H.H. Tate* 730 (holotype NY [barcode 00131040]; isotype F [Acc. No. 603528]).

*Shrubs* 0.5–3 m tall, erect; bark gray; internodes of young branchlets antrorse-hispidulous or glabrate on older internodes. *Stipules* basal sheath densely strigose outside, sparsely to densely sericeous at base and glabrous distally inside; free portion narrowly triangular to linear, gradually tapering at apex or broadly triangular at base and aristate at apex, 0.7–17.5 × 2–3 mm, subcoriaceous to coriaceous, sometimes with an apical collector, densely strigose outside, glabrous inside; with a row of 5–7 colleters on each side, inserted on the margin of the basal sheath. *Leaves* erect and imbricate, sessile or subpetiolate; petioles to 1 mm long, glabrous or sparsely hirtellous; blades very variable in shape and size (Figure 25E–L), lanceolate-elliptic, elliptic, oblanceolate-oblong to obovate, 1–3.5 × 0.3–2 cm, acute, obtuse to subcorporate at base, obtuse, round to acute at apex, coriaceous, dark green and shiny above, pale green below, drying olive-green or pale brown, glabrous throughout or lamina glabrous and midrib sparsely to densely strigillose above and below; margins narrowly to strongly revolute or planar; secondary and tertiary venation submerged within the lamina, sometimes with 2–3 secondary veins on each side of midrib, barely visible above. *Bracts* subtending flowers lanceolate to oblong-lanceolate, 7.5–17 × 2–3 mm, acute at apex, chartaceous to subcoriaceous, glabrous or sparsely hispidulous, margins glabrous or hispidulous-ciliate. *Flowers* with pedicels 1–3 mm long, densely strigose. *Hypanthium* obconical to narrowly obovoid, 2–3 × 1.5–2 mm, antrorse-hispidulous, antrorse-hispid to densely spreading strigose (hairs white). *Calyx* lobes pale green to whitish, oblong-lanceolate, linear-lanceolate to linear, 8–21 × 1.5–4 mm, attenuate, acute to acuminate at apex, glabrous or sparsely hispidulous, margins hispidulous-ciliate, and with 5–6, narrowly-lanceolate, 0.4–0.5 mm long, colleters at each lobe sinus. *Corolla* 8–14.5 cm long; tube pale green, greenish-yellow to cream-white, long-cylindrical and slightly wider near the mouth (anther portion), 6.5–11 cm long, 2–3 mm wide for most length, 6–7.5 mm wide near the mouth, glabrous outside, glabrous from base to 1.2–1.5 cm below the mouth and villous above inside; lobes white to cream-white, lanceolate to oblong-lanceolate, 1.5–3.6 × 0.7–1.6 cm, sub-obtuse, acute to acuminate at apex, gla-

**Figure 25.** *Maguireothamnus tatei.* A. Branch with flowers in anthesis and capsule. B. Nodes with permanent, overlapping stipules. C. Longitudinal section of flower in anthesis. D. Capsule. E-I. Leaf variation within the species. A, C, E: from *Tate* 730 (F); B, F: drawn from *Maguire & Politi* 28449 (US); D: from *Maguire & Politi* 27541 (F); G: drawn from *Steyermark* 58030 (F); H-I: from *Huber* 13287 (US). Illustration by Piero Delprete.
<figure><img src="image_29.png" /><figcaption>Figure 25.</figcaption></figure>
brous outside, glabrous (microscopically papillose) inside. *Anthers* subsessile, elongate, 6.5–7.5 × 0.8–1 mm, base round, apex acuminate. *Style* 7–12.7 cm long, exerted 10–17 mm beyond corolla mouth, glabrous for most of the length and papillose on distal 2.7–5 cm below style branches; style branches oblong-spathulate, 2–2.5 mm long. *Capsules* obovoid, ovoid to ellipsoid, 1–2.5 × 0.8–1 cm, obtuse to acute at base, round at apex, dark brown, hispidulous to hispid (hairs 0.7–1 mm long). *Seeds* 2.5–3 × 1.5–2.5 mm.

**Distribution and ecology:** Endemic to the tepuis of the Venezuelan state of Amazonas, on Cerro Duida, Cerro Sipapo, Cerro Huachamacari, and Cerro Marahuaca-Atuhua-Shiho (Figure 23). Growing on exposed sandstone outcrops, open scrub formations, tepui slope and summit scrubs, meadows, and open savannas, at 1300–2555 m altitude.

**Phenology:** Flowering from August to February, and fruiting from November to January.

**Suggested conservation status:** Least Concern (LC). This species is known from numerous collections from two main regions within the state of Amazonas, southern Venezuela, at altitudes of 1230–2650 m. The first region corresponds with the nearby tepuis Cerro Duida, Cerro Huachamacari, and Cerro Marahuaca-Atuhua-Shiho; most of them are part of the Duida-Marahuaca Massif, which is included within the Duida-Marahuaca National Park. The second region corresponds to the distant tepui Cerro Sipapo, ca. 4°57′N, 67°20′W, 1500 m, which is about 200 km NW of the other region, and is not within a protected area. The size of the populations is unknown. Therefore, considering its large extent of occurrence (EOO), and that most of the collections were made within a national park, which is without any imminent threat, this species is classified as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

**Specimens examined:** VENEZUELA: Amazonas: Depto. Atabapo, Cerro Duida (“Yennamadi”), sector meridional, herbazal arbustivo sobre ladera suavemente inclinada cerca del borde S de la meseta, 3°19′N, 65°38′W, 2100 m, 19 Nov. 1991 (fr), O. Huber 13281 (US, VEN); Depto. Atabapo, Cerro Duida (“Yennamadi”), sector meridional, herbazal abierto en un pequeño valle y arbustales densos sobre colinas bajas cerca del borde S de la meseta, 3°17′N, 65°38′W, 2060 m, 20 Nov. 1991 (fl), O. Huber 13287 (US, VEN); Depto. Atabapo, open rocky plateau of Cerro de Marahuaca, above Salto Los Monos on

tributary of headwaters of Río Iguapo, 3°37′N, 65°23′W, 2555 m, 27 Feb. 1985 (fl), R.L. Liesner 18040 (MO, NY, VEN); Depto. Atabapo, gallery forest and open area on Plateau of Huachamacari, 3°50′N, 65°43′W, 1720 m, 1 Mar. 1985 (fl), R.L. Liesner 18066 (MO, VEN); Cerro Sipapo (Paráque), [ca. 4°57′N, 67°20′W], near Savanna Camp, 1500 m, 6 Dec. 1948 (fr), B. Maguire & L. Politi 27541 (F, NY, RB); Cerro Sipapo (Paráque), between Savanna Camp and West Peak, 1500–1800 m, 20 Dec. 1948 (fl), B. Maguire & L. Politi 27801 (NY [2 sheets], VEN); Cerro Sipapo (Paráque), near Savanna Camp, 1500 m, 19 Jan. 1949 (fl), B. Maguire & L. Politi 28449 (FT, NY [2 sheets], US, VEN); Cerro Sipapo (Paráque) [ca. 4°57′N, 67°20′W], occasional on North Mountain, 1500 m, 25 Jan. 1949 (fr), B. Maguire & L. Politi 28582 (IAN, NY [2 sheets], P, U); Cerro Duida, Río Cunucunuma, N slopes and ridges of Caño Negro basin, 2000–2300 m, 23 Nov. 1950 (fr), B. Maguire et al. 29695 (NY [2 sheets]); Cerro Huachamacari, Río Cunucunuma, cumbre, 1700 m, 4 Dec. 1950 (fl), B. Maguire et al. 29810 (F, NY); Cerro Huachamacari, Río Cunucunuma, summit of E ridge [ca. 3°55′N, 65°30′W], 1820 m, 8 Dec. 1950 (fr), B. Maguire et al. 30058 (NY [2 sheets], P, U, US); Cerro Huachamacari, Río Cunucunuma, W rim of ridge N. 1, summit camp, [ca. 3°55′N, 65°30′W], 1900 m, 11 Dec. 1950 (fr), B. Maguire et al. 30129 (NY, P, U, US); Cerro Huachamacari, Río Cunucunuma, along SE escarpment, [ca. 3°55′N, 65°30′W], 1900 m, 11 Dec. 1950 (fl buds), B. Maguire et al. 30142 (FT, NY, US, VEN); Cerro Huachamacari, Río Cunucunuma, between Summit Camp and E escarpment, [ca. 3°55′N, 65°30′W], 1800–1950 m, 16 Dec. 1950 (fl), B. Maguire et al. 30263 (NY [2 sheets], US, VEN); Parque Nacional Duida-Marahuaca, cumber del Cerro Duida, esquina SE, 2200 m, Mar. 1993 (fl), B. Stergios et al. 15768 (NY, PORT, VEN); Cerro Duida, SE-facing slopes along Caño Negro (tributary of Caño Iguapo), 1700 m, 25–26 Aug. 1944 (fl), J.A. Steyermark 58030 (IAN [frag], NY, US, VEN); summit of Cerro Duida, Savana Hills, 1025–1200 m, 2 Sep. 1944 (fl), J.A. Steyermark 58240 (F, NY, US); summit of Cerro Duida, on high moist ridge top, 1820–2075 m, 4 Sep. 1944 (fl), J.A. Steyermark 58335 (F, NY, IAN [frag]), 58348 (F); Depto. Atures, Serranía Sipapo, cumbre, seción septentrional, vegetación tipo sabana a lo largo de un riachuelo, 5°0′N, 67°30′W, 1500 m, 17 Feb. 1981 (fl), J.A. Steyermark et al. 124538 (NY, VEN); Depto. Atabapo, Cerro Marahuaca-Huha, cumbre altiplanicie no arbolada, mas o menos plana, con piedras expuestas, río abajo, 3°40′N, 65°30′W, 2580 m, 31 Jan. 1982 (fr), J.A. Steyermark et al. 125914 (MO, NY, US, VEN); Depto. Atabapo, Cerro Marahuaca-Fhuif, cumbre, zona pantanosa no arbolada en la altiplanicie, río arriba, 3°35′N, 65°20′W, 2480–2500 m, 2 Feb. 1982 (fl), J.A. Steyermark et al. 126073 (MO,
NY, VEN); Depto. Atabapo, Cerro Marahuaca-Atuhua-Shiho, cumbre, parte aislada al SW del cerro, vegetación no arbolada en terreno pendiente, 3°30'N, 65°20'W, 2450-2480 m, 9-10 Feb. 1982 (fl), *J.A. Steyermark et al. 126338* (F, MO, NY, US, VEN); Depto. Atabapo, Cerro Duida, cumbre, cerca de escarpa oriental, sabana pantanosa dominada por *Eliamphora y Brewcaria*, 3°25'N, 65°40'W, 1230 m, 10 Feb. 1982 (fl), *J.A. Steyermark et al. 126399* (NY, VEN); Depto. Atabapo, Cerro Marahuaca, summit on undulating plateau with islands of shrubs and *Mara-huacaea*, with stream branch leading to Salto de Monos and SE edge to Caño Iguapo, S and SE of Summit Camp, 3°37'N, 65°23'W, 2520-2650 m, 26-27 Feb. 1985 (fl), *J.A. Steyermark & B. Holst 130685* (MO, NY, U, VEN), 130731 (MO, NY); Mount Duida, slopes of Rudge 25, 1830-2000 m, 1928 (fl), *Tate G. H. H. 427* (NY, frag-F [ex NY]); Mount Duida, 1928-1929 (fl), *G.H.H. Tate 1016* (NY, frag-F [ex NY]); Estación D-21, NNE of Savana Hills Camp of Tate, Cerro Duida, inmediatamente al N de La Esmeralda, en el Río Orinoco, 3°10'N, 65°31'W, 1500 m, 2 Feb. 1975 (fl), *S.S. Tillett et al. 751-49* (NY, U, VEN), 751-49a (NY, U, VEN).

## 5. NEBLINATHAMNUS

*Neblinathamnus* Steyerm., Mem. New York Bot. Gard. 10(5): 229. 1964.

Steyerm. in Mem. New York Bot. Gard. 23: 309. 1972; Steyerm. in Lasser and Steyermark, Fl. Venezuela 9(1): 318-322, fig. 51. 1974; Steyerm. in Ann. Missouri Bot. Gard. 76: 968. 1989; Delprete and Steyerm. in Steyermark et al., Fl. Venez. Guayana 8: 659-660, fig. 520. 2004.

Type: *Neblinathamnus argyreus* Steyerm.

*Shrubs. Stipules* basally adnate to the petioles, minute, broadly to shallowly triangular to truncate, sometimes shallowly bilobed, often bilobed at apex, or rarely constituted of two free lobes on each side of the stem; with a row of 7-9 sparse colleters on margin in shallowly triangular to truncate stipules, or with a row of 4-5 colleters on margin of each lobe in stipules bilobed or constituted of two free lobes. *Leaves* opposite or ternate, short-petiolate or subsessil; blades elliptic, obovate or oblong, thinly coriaceous; domatia absent. *Inflorescence* axillary, sometimes appearing terminal when on apical node, with 1-2 flowers, or cymose, (2-)3-5-flowered. *Flowers* heterostylous, 5-merous, short-pedicellate, subtended by two small, setaceous bracts. *Hypanthium* obovoid, oblong-ovoid, turbinate or narrowly obovoid, sparsely to densely hirtellous. *Calyx* lobes erect, equal

to subequal, spathulate, lanceolate, linear-acuminate, or linear-spathulate, with 2-3 colleters at each lobe sinus. *Calycophylls* absent. *Corolla* hypocrateriform, actinomorphic; corolla green to yellowish-green, or tube red and lobes yellowish or greenish; tube externally glabrous or hirtellous, internally puberulent throughout or pubescent at mouth, with a pubescent ring at orifice inside; orifice annular thickening absent; lobes left-contorted, narrowly ovate, ovate, elliptic or oblong, margin entire, obtuse or rounded at apex. *Pollen* 3-colporate; exine foveolate-reticulate, perforate. *Short-styled flowers*: stamens included, inserted at distal portion of corolla throat; filaments short; anthers linear; style included, style branches linear. *Long-styled flowers*: stamens included, inserted at medio-basal or median portion of corolla throat; filaments short; anthers narrowly oblong or linear; style as long as corolla tube, style branches barely exserted beyond corolla mouth or exserted, style branches narrowly ovate, narrowly lanceolate or linear. *Ovary* with a stalked placenta; stalk inserted at the middle of the septum, terminating with an oblong-ellipsoid extension; ovules inserted on the placental extension. *Fruit* capsular, dehiscing loculicidally, thinly woody. *Seeds* horizontal or vertical, minute, 3-5-angular, laterally compressed; testa reticulate-foveolate.

**Geographic distribution:** A genus of two species restricted to the Venezuelan and Brazilian sides of the Cerro de la Neblina Massif, hence the generic name.

**Morphological notes:** Steyermark (1964, 1974) wrote that the stipules of *Neblinathamnus* are obsolete and caducous, but they are instead shallowly triangular to truncate, shallowly bilobed, or constituted of two free lobes, and persistent.

### Key to Neblinathamnus Species

1. Leaf blades obovate, obovate-oblong, narrowly oblong to linear-oblong, 1-2.5(-3.3) × 0.5-1.3(-1.5) cm, round, sometimes mucronate at apex; flowers 1-2 per axil; corolla tube 10.5-14.5(-16) mm long.................... 1. *N. argyreus*

1. Leaf blades oblong-elliptic to ovate-lanceolate, (2.5-)3-6 × (0.8-)1-3 cm, acute to obtuse at apex; inflorescence cymose, (2-)3-5-flowered; corolla tube 9-12 mm long...... ......................................................................................... 2. *N. brasiliensis*

5-1. *Neblinathamnus argyreus* Steyerm., Mem. New York Bot. Gard. 10(5): 229, fig. 77. 1964. (Figures 26-27).

Type: VENEZUELA. Amazonas: Cerro de la Neblina [ca. 0°53'N, 65°59'W], Río Yatua, along escarpment,
1200 m, shrub 2 m tall, corolla greenish, 16 Nov. 1957 (fl, fr), *B. Maguire, J.J. Wurdack, Keith & C.K. Maguire* 42120 (holotype NY [barcode 00132332]; isotypes P [barcode P00729462], US [barcode 00137734], VEN [Acc. No. 49441]).

(=) *Neblinathamnus glabratus* Steyerm., Mem. New York Bot. Gard. 10(5): 231. 1964.

Type: VENEZUELA. Amazonas: Cerro de la Nebli-
na, Río Yatua, occasional along W escarpment
edge, summit, 1700–2000 m, 30 Dec. 1953 (fl, fr), B.
*Maguire, J.J. Wurdack & G.S. Bunting* 37011 (holo-
type NY [barcode 00132335]; isotypes F [Acc. No.
1589377], VEN [Acc. No. 49435]).

Shrub 0.3-2 m tall, erect; internodes of young
branchlets hispidulous-hirtellous or densely hispidu-
lous. *Stipules* shallowly triangular to truncate, 0.3-0.6 x
1-1.5 mm, sometimes shallowly bilobed, subcoriaceous,
hispidulous-hirtellous or sparsely to densely hispidulous
outside, sericeous-pubescent inside, with a row of 7-9
sparse colleters on margin. *Leaves* shortly petiolate; pet-
ioles 2-3 mm long, glabrous or hispidulous; blades obo-
vate, obovate-oblong, narrowly oblong to linear oblong,
1-2.5(-3.3) x 0.5-1.3(-1.5) cm, cuneate to acute at base,
round and sometimes mucronate at apex, coriaceous,
dark green and shiny above, silvery to whitish below,
drying brown above and pale to dark olive green below,
glabrous and rugose above, tessellate and foveolate, with
a silvery appearance, foveolate with tufts of vermiform
hairs and margins hispidous-ciliate, below; midrib
impressed above, prominent and appressed-hispidulous
below; margins narrowly revolute; secondary veins 5-7
on each side of midrib, impressed above and prominent
below; tertiary venation obsolete above and finely retic-
ulate below. *Flowers* axillary, 1-2 per axil. *Bracteoles*
subtending flowers setaceous, linear to very narrowly
lanceolate, 1-3 x 0.1-0.4 mm, hispidulous. *Flowers* pedi-
cellate; pedicels 1.5-2.5 mm long during anthesis, his-
pidulous-hirtellous or densely hirtellous. *Hypanthium*
obovoid to oblong-ovoid, 2-2.5 x 1.7-2 mm, sparsely to
densely hirtellous. *Calyx* lobes subequal, linear, linear-
lanceolate to linear-spathulate, 4-7 x 0.3-1 mm, acute
to acuminate at apex, sparsely to moderately hirtellous
or hispidulous, with 2 or 3 linear, 0.2-0.3 mm long,
colleters at each lobe sinus. *Corolla* 15-22 mm long,

green to yellowish-green, or tube red and lobes green to
yellowish-green; tube narrowly cylindrical and slightly
wider at mouth, 10.5-14.5(-16) mm long, 1-2 mm wide
at lower third, 2-3 mm wide at mouth, glabrous or
sparsely hirtellous outside, glabrous from base to 3-4
mm below the mouth and puberulent above, portion
corresponding with the anthers with vermiform, multi-
cellular hairs, inside; lobes ovate to oblong-ovate, 4.5-6
× 3.5-5 cm, round, obtuse or acute at apex, glabrous
outside, glabrous (microscopically papillose) inside.
*Short-styled flowers* (Stein et al. 1636; corolla tube 14
mm long, lobes 5.5-6 mm long): stamens included,
inserted at 3 mm below corolla mouth; filaments 0.7
mm long; anthers narrowly oblong, 2.5 × 0.25-0.3 mm,
acute at both ends, tips reaching corolla mouth; style
included, 1010.5 mm long, 5/7 of corolla tube, glabrous,
style branches linear, 2.5-2.7 mm long, acuminate at
tip. *Long-styled flowers* (Luteyn & Steyermark 9437;
corolla tube 10.5 mm long, lobes 4.5 mm long): sta-
mens included, inserted at 4 mm below corolla mouth;
filaments 0.8-1 mm long; anthers narrowly oblong, 2 ×
0.15-0.2 mm, acute at base, apiculate at apex; style as
long as the corolla tube, 10.5 mm long, style branches
narrowly ovate, 0.7 mm long, round at tip. *Capsules*
with pedicels to 4 mm long, subturbinate to obovoid-
subglobose, 4-5 × 3.5-4 mm, acute to obtuse at base,
round to truncate at apex, brown, densely hirtellous or
hispidulous. *Seeds* oblong-elliptic, 1-1.5 × 0.6-0.9 mm,
yellowish rust brown.

Notes: Steyermark (1964: 231-232; 1974: 319) differ-
entiated *Neblinathamnus argyreus* from *N. glabratus* by
the corolla tube sparsely hirtellous outside (vs. complete-
ly glabrous outside in *N. glabratus*), leaf blades obovate
or oblong-obovate, foveolate-tessellate and with tufts of
vermiform papillae, giving a whitish appearance below
(vs. narrowly oblong or linear-oblong, foveolate-tessel-
late but without tufts of white hairs below), corolla green
to yellowish-green (vs. red or tube red with lime-yel-
low lobes), and flowers mostly solitary or sometimes in
pairs (vs. flowers in pairs). At the same time, Steyermark
(1964: 231-232) stated "That there are pronounced dif-
ferences between *N. argyreus* and *N. glabratus* is appar-
ent. Although it is quite possible that additional collec-
tions may break down the differences which appear to
exist on the basis of the present material at hand, it is
also evident that the differences now manifested can be

**Figure 26.** *Neblinathamnus argyreus.* *A.* branch with flowers in anthesis and flower buds. *B–C.* Stipule variation within the same branch. *B.* Truncate stipule. *C.* Shallowly bilobed stipule. *D.* Dissected long-styled flower; style as long as corolla tube with narrowly ovate branches. *E.* Dissected short-styled flower; style included with linear branches. *F.* Dehiscent capsule. *A–D:* drawn from *Luteyn & Steyermark 9437* (F); *E:* drawn from *Stein et al. 1636* (F); *F:* drawn from *Liesner et al. 16102* (US). Illustration by Piero Delprete.
<figure><img src="image_30.png" /><figcaption>Figure 26.</figcaption></figure>
best be expressed as specific taxa.” Delprete and Steyermark (2004d: 660) synonymized the two taxa and wrote “Seyermark distinguished *N. glabratus* by details of leaf shape and pubescence and by the absence of vermicomphiles on its lower leaf surface. However, as he predicted in his description of *N. glabratus*, recent collections have shown that these characters intergrade. Consequently these names are here considered as synonymous.” In addition, the types of these two taxa were collected in two nearby localities near Río Yatua, Cerro da Neblina, Venezuela. After a detailed analysis complemented by additional material, the conclusion reached by Delprete and Steyermark is here confirmed, and the two taxa are maintained as synonymous.

**Distribution and ecology:** Endemic to Serranía de la Neblina (Figure 27), state of Amazonas, southernmost Venezuela, in open scrub forests, escarpments and summits of tepuis. The Neblina Massif divides northern Brazil from southern Venezuela, and is present on both countries, with most part in Venezuela. This portion of the Guiana Shield highlands is known as “Tepuis”, includes numerous unique abitats, and is a region rich in

endemic species. So far, this species has been found only in Venezuela. On the Brazilian side, where the highest point of the mountain (3014 m), as well as for Brazil, is located, remains poorly explored due to the difficult access. This species grows on exposed sandstone outcrops, open scrub formations and escarpments, at 1600–1900 m altitude.

**Phenology:** Flowering specimens were collected from November to February and April, and fruiting specimens in December and February.

**Suggested conservation status:** Least Concern (LC). Cerro de la Neblina is a protected area, with the Serranía de la Neblina National Park on the Venezuelan side, and the Pico da Neblina National Park on the Brazilian side. This species has been collected only on the Venezuelan side, at altitudes of 1600–1900 m. The size of its populations is unknown. As all the collections were made within a remote, well-preserved national park, which is without any imminent threat, this species is treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

<figure><img src="image_31.png" /><figcaption>**Figure 27.** Distribution of *Neblinathamnus argyreus* (circles) and *N. brasiliensis* (squares).</figcaption></figure>
**Specimens examined:** VENEZUELA: Amazonas:
Río Negro, Camp III, Neblina Massif, NW Plateau, 13.5 km ENE of Cerro de la Neblina Base Camp, 0°54'N, 66°4'W, 1750-1850 m, 16-18 Feb. 1984 (fr), R.L. Liesner 16102 (MO, NY, US); Cerro de la Neblina, altiplanicie en la cumbre del brazo nor-occidental, al N del campamento base a lo largo del Río Mawarinuma, afluente del Río Baria, 0°52'N, 66°5'W, 1880 m, 7-8 Feb. 1984 (fl), J.L. Luteyn & J.A. Steyermark 9437 (F, MO, NY, US); Cerro de la Neblina, uppermost escarpment slopes S of Camp 3, 1700 m, 27 Dec. 1953 (fl), B. Maguire et al. 36960 (NY, SP, paratype); Cerro de la Neblina, Río Yatua, occasional, NW Cumbre, 1900 m, 20 Dec. 1953 (fl), B. Maguire et al. 37011-A (IAN, NY, RB, paratype); Cerro de la Neblina, frequent in open areas 6-8 km N of Cumbre Camp, 1800-1900 m, 10 Jan. 1954 (fl), B. Maguire et al. 37175 (NY [2 sheets], paratype); Cerro de la Neblina, escarpment edge above Cañon Grande, E of Cumbre Camp, locally abundant, 1800 m, 22 Nov. 1957 (fl), B. Maguire et al. 42171 (COL, F, NY, P, S, US, VEN, paratype); Cerro de la Neblina, 6.5 km SSW of base camp, southern extension of range, ridge top with Bromeliads, *Heliamphora* and low shrubs, 0°47'N, 66°11'W, 1600 m, 18 Apr. 1984 (fl), B.A. Stein et al. 1636 (F, MO, NY, US); Cerro de la Neblina, altiplanicie en la cumbre del brazo nor-occidental, al N del campamento base a lo largo del Río Mawarinuma, afluente del Río Baria, rocky margins, 0°52'-53'N, 66°5'W, 1880 m, 7-8 Feb. 1984 (fl), J.A. Steyermark & J.L. Luteyn 129788 (F, MO, NY, U; same as Luteyn & Steyermark 9437).

### 5-2. Neblinathamnus brasiliensis Steyerm., Mem. New York Bot. Gard. 23: 309. 1972. (Figures 4I, 27-28).

Type: BRAZIL. Amazonas: Rio Negro, Rio Cauaburí, Rio Maturacá, on slopes of Serra Pirapucú [Serra da Neblina, ca. 0°48'24"N, 66°0'19"W], plant 1 m tall, calyx green, corolla with red base and green lobules, 1300 m, 26 Jan. 1966 (fl, fr), N.T. Silva & U. Brazão 60907 (holotype NY [barcode 00132333]; isotypes COL [Acc. No. 129225, barcode COL000004628], F [Acc. No. 1713895], S [Acc. No. S05-1759], US [barcode 00137735]).

(=) *Neblinathamnus aracamunianus* Steyerm., Ann. Missouri Bot. Gard. 76: 968. 1989.

Type: VENEZUELA. Amazonas: Depto. Río Negro, [Neblina-Aracamuni Massif], Cerro Aracamuni, summit, Popa Camp, savanna at edge of tepui, shrub 0.5 m tall, corolla tube red with lobes greenish-

white, 1°26'N, 65°47'W, 1550 m, 19 Oct. 1987 (fl), R. Liesner & F. Delascio 22153 (holotype MO [Acc. No. 3663774]; isotypes NY [barcode 00007410], VEN [Acc. No. 82233]).

Shrub 0.5-1 m tall, erect; internodes of young branchlets strigillose-hirtellous or appressed hirtellous. *Stipules* bilobed or constituted of two free lobes, lobes ovate, deltoid to broadly triangular, 1.2-1.5 × 0.7-0.8 mm, subcoriaceous, hispidulous-hirtellous or sparsely to densely hispidulous outside, appressed-hispidulous inside, with a row of 4-5 colleters on margin of each lobe. *Leaves* shortly petiolate; petioles 1.5-3 mm long, densely appressed strigillose-hirtellous or moderately to densely appressed hirtellous; blades oblong-elliptic to ovate-lanceolate, (2.5-)3-6 × (0.8-)1-3 cm, acute to obtuse at base, obtuse to acute at apex, coriaceous, dark green and shiny above, whitish pale green below, drying brown above and pale to dark olive green below, lamina glabrous, midrib sulcate and strigillose or with sparse appressed hairs above, tessellate and foveolate, foveolate with tufts of short vermiform hairs and margins glabrous, below; midrib impressed above, prominent and appressed-hispidulous below; margins narrowly revolute; secondary veins (4-)5-7 on each side of midrib, obscure, inconspicuous to obsolete above, inconspicuous to slightly prominent, appressed-hispidulous or glabrescent in older leaves) below; tertiary venation obsolete above, finely reticulate below and impressed, the areoles in depressed foveolate areas with circles of minute white trichomes. *Inflorescences* axillary, sometimes appearing terminal when axillary on apical node, cymose, (2-)3-5-flowered. *Bracteoles* subtending flowers subulate, setaceous, 1.5-3 × 0.1-0.4 mm, sparsely hispidulous. *Flowers* pedicellate; pedicels 1-3 mm long during anthesis, densely tomentose-strigillose. *Hypanthium* turbinate or narrowly obovoid, 1.8-2 × 1.4-1.5 mm, densely substrigillose-hirtellous. *Calyx* lobes green, equal to subequal, linear, linear-acuminate or linear-spathulate, sometimes wider near the apex, (4-)5-7 × 0.3-0.7 mm, acuminate to subacute at apex, sparsely hispidulous to hispidulous outside, glabrous inside, margins sparsely hispid-ciliate, with 2 or 3 linear, 0.2-0.3 mm long, colleters at each lobe sinus. *Corolla* 15-18 mm long, tube red and lobes green to yellowish- to whitish-green; tube narrowly cylindrical and slightly wider at mouth, 9-12 mm long, 1.4-1.8 mm wide at lower third, 2.2-2.5 mm wide at mouth, glabrous outside, glabrous from base to 3-4 mm below the mouth and puberulent above, portion corresponding with the anthers with vermiform/multicellular hairs inside; lobes 5, elliptic-oblong or ovate-oblong, 5-6 × 2.5-3.7 cm, round or obtuse at
apex, glabrous outside, glabrous (microscopically papillose) inside. *Short-styled flowers*: unknown. *Long-styled flowers*: stamens inserted at 1 or 3-4 mm from corolla tube base, filaments 1-3 mm long; anthers linear, 2-2.5 × 0.2 mm, round at base, acute at apex; style exserted, 10-13 mm long (style branches exserted beyond corolla mouth), glabrous, branches narrowly lanceolate to linear, 2-3 mm long, acuminate at apex. *Capsules* broadly obconical to obovoid, 4-5 × 3-4 mm, acute to obtuse at base, round to truncate at apex, brown, densely subhirtellous. *Seeds* unknown.

**Notes:** Steyermark (1972a: 309) described *Neblinathamnus brasiliensis* Steyerm. as having “3-4 leaves per node” [foliis 3-4-verticillatis] but the leaves of type specimens of this species are opposite or ternate. In the same description he also wrote that the corolla throat is densely villous-puberulent above 5 mm from the base [“intus supra basin 5 mm dense villosulo-puberulenti”], but in all the specimens examined the corolla throat is glabrous from base to 3-4 mm below the mouth and puberulent above, and the portion corresponding with the anthers is covered with vermiform/multicellular hairs.

Steyermark (1989: 968) along with the description of *Neblinathamnus aracamunianus* Steyerm. added “*Neblinathamnus* was originally described by Steyermark (1964) with two species known, *N. argyreus* from Venezuela on Cerro de la Neblina and *N. glabratus* [= *N. argyreus*] from adjacent northern Brazil on Serra Pirapucú [sic! Type material of *N. glabratus* was collected on Serra da Neblina, as *N. argyreus*]. This third species differs from the other two in its shorter corolla; larger leaf blades with subacute or obtuse apices; and shorter, spathulate-linear, subacute calyx lobes, which are more dilated at the apex, dorsally pubescent, and more abundantly ciliolate.” However, with this statement it is clear that Steyermark confused his new species with a third species already described by him, *N. brasiliensis*, which is from Serra Pirapucú, near Pico da Neblina, at the Venezuelan-Brazilian border, and with which he did not make a comparison with his fourth new species. Delprete and Steyermark (2004d: 660) synonymized the two taxa without any further comment. A comparative analysis of the type specimens of both taxa confirmed that the morphological features *N. brasiliensis* and *N. aracamunianus* are widely overlapping. In addition, although located in different countries, both Cerro

Aracamuni (Venezuela) and Serra Pirapucú (Brazil) are parts of the Cerro da Neblina Massif, therefore geographically close (Figure 27), and the two taxa are here confirmed to be synonymous.

**Distribution and ecology:** Only known from the type specimens. Endemic to Cerro da Neblina Massif, occurring both on the Venezuelan (Cerro Aracamuni) and on the Brazilian side (Serra Pirapucú) (Figure 27). Growing on meadows along escarpments on tepui summits, at 1300 and 1550 m altitude.

**Phenology:** The gathering collected in January has flowers and mature capsules, and that collected in October only has flowers.

**Suggested conservation status:** Least Concern (LC). As explained for the other species of this genus, Cerro de la Neblina is protected by two national parks on the Venezuelan and on Brazilian sides. This is a rare species, only known by the two type gatherings, at 1550 m on the Venezuelan side, and at 1300 m on the Brazilian side. The size of the populations is unknown. As the collections were made within remote, well-preserved national parks, which are without any imminent threat, this species is treated as Least Concern (LC) following IUCN criteria (IUCN 2012, 2019).

## 6. NEOBERTIERA

*Neobertiera* Wernham, J. Bot. 55: 169. 1917.

Emend. Sandwith, Kew Bull. Misc. Inf. 1931: 470. 1931; emend. Steyermark, Mem. New York Bot. Gard. 17: 289-290. 1967; emend. Delprete, Phytotaxa 206: 118-132. 2015; Delprete, Phytotaxa 392: 225-231. 2019.

Type: *Neobertiera gracilis* Wernham.

*Perennial herbs*, subshrubs or shrubs. *Stipules* basally adnate with the petioles, broadly ovate, ovate, oblong-ovate or narrowly triangular, acuminate or bifid at apex; pubescent, and with numerous colleters either in a row or as two groups on each side at medio-basal portion inside. *Leaves* opposite, petiolate; blades elliptic, oblong-elliptic to oblanceolate, chartaceous; domatia absent. *Inflorescences* terminal, many-flowered, laxly cymose to densely glomerulate or subcapitate (extremely reduced cyme),

**Figure 28.** *Neblinathamnus brasiliensis*. A. Branch with flowers in anthesis, flower buds and capsule. B. Node with bilobed stipule. C. Dissected long-styled flower; style exserted with linear branches. D. Capsule, before dehiscence. A–D: drawn from *Silva & Brazão 60907* (F). Illustration by Piero Delprete.
<figure><img src="image_32.png" /><figcaption>Figure 28.</figcaption></figure>
when laxly cymose with ultimate branches scorpiod, (more evident when fully expanded), when glomerulate to densely cymose with lateral branches terminating in flowering glomerules. *Flowers* commonly distylous, rarely tristylous, 4-5-merous. *Hypanthium* globose, broadly obovoid to obovoid, appressed strigose, trichomes simple or basally tuberculate (tubercules multicellular in *N. montedouradensis*). *Calyx* tube extremely reduced or absent; lobes free, frequently unequal, erect, narrowly-lanceolate, linear-lanceolate to narrowly lanceolate, acute at apex, with 1 or 2 colleters in each calycine sinus. *Calycophylls* absent. *Corolla* hypocrateriform, white ("orange" [?] in *N. montedouradensis*), tube appressed strigose-sericeous outside, puberulent inside, with a narrow line of sparse yellow moniliform hairs at mouth; lobes left-contorted, thickened at base, narrowly oblong or oblongovate, margin entire, round or acute at apex. *Pollen* 3-colporate; exine foveolate, perforate. *Short-styled flowers:* stamens included, filaments inserted around the middle or at distal portion of corolla tube; anthers linear; style included, much shorter than corolla tube, style branches narrowly oblong or elliptic. *Long-styled flowers:* stamens included, filaments inserted at the middle or near the base of corolla tube; anthers linear or narrowly sagitate; style about the same length as the corolla tube (tip of branches barely exserted) or exserted beyond corolla mouth, style branches narrowly oblong, narrowly lanceolate to linear. *Ovary* with a stalked placenta; stalk inserted at the middle or medio-basal portion (*N. montedouradensis*) of the septum, with a hemi-elliptic extension; ovules inserted on the placental extension. *Fruits* independent, globose, falling off as dispersal units, leaving a tiny round scar on the infructescence rachis, crustaceous, appressed strigose, trichomes simple or basally tuberculate in *N. montedouradensis*. *Seeds* many, inserted on the stalked placenta, small, tetrahedral.

**Notes about generic emendations:** Wernham (1917) described *Neobertiera* with a single species, *N. gracilis*, and positioned it in the tribe Hamelieae. His description and discussion are erroneous because he stated that its fruits are berries with a few globose seeds. He also stated that the inflorescence of *Neobertiera* is similar to those of a section of *Bertiera*, hence the generic name. In addition, he described the sole species as having “Stamina 4, infra corollae fauce inserta” (although the stamens of the

holotype specimen are inserted at the distal portion of the corolla, and not at the corolla mouth) and “Anthers 1.5 mm, filaments slightly shorter, [...] style 1.6 mm long, stigmatic branches 0.7 mm.” He also wrote, with a pencil, on the holotype specimen “filaments inserted at mid-tube”, contrasting with what he wrote in the publication. Therefore, Wernham’s original description of this species was made using short-styled specimens (Jenman 2388).

Both Sandwith (1931: 470) and Steyermark (1967: 289-290) did not notice the disty in *Neobertiera gracilis*, although they had ample material at hand. Sandwith (1931: 470) saw several collections of this species at K and described it as “a very pretty herb or low shrub, 1-5 ft. [30-150 cm] high, with pure white flowers. The flowers were described by Wernham as tetramerous, but even on his own type specimen at least two of the calyces are pentamerous, while pentamery is obvious on the material at Kew.” Therefore, Sandwith correctly emended the merosity of the genus. In fact all the species successively described in this genus are pentamerous.

Steyermark (1967: 234-235) in the key to the Rondeletieae genera included: *Gleasonia*, *Sipaneopsis*, *Cuatrecasasiiodendron* [= *Arachnothryx*], *Rondeletia*, *Arachnothryx*, *Pteridocalyx*, *Neobertiera*, *Maguireothamnus*, *Neblinathamnus* [as “Neblinanthus”], *Sipanea*, *Limnosipanea*, *Chalepophyllum*, and *Dendrosipanea*. He did not describe the fruits of *Neobertiera*, but considering all the other genera of his widely delimited Rondeletieae are capsular, he probably assumed that its fruits are not berries.

*Neobertiera* has been traditionally reported as having homostylous flowers, with filaments inserted at the distal portion of the corolla tube (Wernham 1917; Sandwith 1931; Steyermark 1967). However, Delprete (2015a) after a detailed observations showed that all the species of *Neobertiera* are distylous and that the filaments are not always inserted at the distal portion of the corolla tube. In long-styled flowers the stamens are inserted near the base of the corolla tube, and in short-styled flowers the filaments are inserted at median or in distal portions of the corolla tube, and their vascular tissue could be easily traced all the way to the base of the corolla. Delprete (2015a) therefore emended the description of *Neobertiera*, by reporting for the first time that the genus is heterostylous, and described three additional species. Delprete (2019b) described a new species, *N. montedouradensis*, from the state of Pará, Amazon Basin, which represents

**Figure 29.** *Neobertiera gracilis.* A. Branch with inflorescences. B. Node with bifid stipule. C. Detail of a lateral branch of a fully developed inflorescence. D. Dissected short-styled flower, with stamens inserted at the upper portion of the corolla tube. E. Dissected long-styled flower, with stamens inserted at the middle of the corolla tube. F. Fruit. A-B: drawn from *Maguire & Fanshawe 22964* (F); C: drawn from *Gleason 6* (US) D: drawn from *Grewal & Persaud 110* (L); E: drawn from *Schultz & Wessels Boer LBB 10223* (L); F: drawn from *Smith 2106* (US). Illustration by Piero Delprete (Reproduced from fig. 6 of Delprete (2015a) with permission of copyright holder Magnolia Press).
<figure><img src="image_33.png" /><figcaption>Figure 29.</figcaption></figure>
a significant range extension and the first record of this genus for Brazil. The new species is unique within the genus by having hypanthia and fruits with trichomes conspicuously tuberculate at base, requiring an expansion in the generic description. In addition, he published a new combination for a species occurring in French Guiana.

**Geographic distribution:** A genus of five species endemic to the Guiana Shield (Funk & Berry, 2005), with two species probably restricted to Guyana, two species probably restricted to French Guiana, and one species from the Brazilian states of Pará and Amapá, at the southeastern edge of the Guiana Shield.

Key to *Neobertiera* species

1. Inflorescence subcapitate, an extremely reduced cyme; corolla tube 3–3.2 mm long, lobes 0.9–1 × 0.5–0.6 mm.................. 2. *N. micrantha*

1. Inflorescence densely to laxly cymose or glomerulate; corolla tube 4.5–10.5 mm long, lobes 3–7 × 1–2.7 mm..................2

2. Inflorescence glomerulate to densely cymose during anthesis, lateral branches cymose, extremely reduced, terminating with glomerules .................................................. 5. *N. palustris*

2. Inflorescence laxly cymose during anthesis, lateral branches dichotomously divided, with one flower at the base of each bifurcation, ultimate branches scorpiod, more evident when fully expanded..................................................................................3

3. Stipules narrowly triangular, acute at apex; leaves broadly ovate to elliptic, round to obtuse at base; hypanthium with hairs basally tuberculate; corolla 14–15.5 mm long, tube 9.5–10.5 mm long, lobes ovate to oblong-ovate.................................................. 3. *N. montedouradensis*

3. Stipules ovate to broadly ovate, acuminate or bifid at apex; leaf blades narrowly elliptic, elliptic, lanceolate or oblong-elliptic to oblanceolate, acute-decurrent at base; hypanthium with hairs not basally tuberculate; corolla 7–7.7 or 9.5–13.5 mm long, tube 4–7.5 mm long, lobes oblong or narrowly oblong ..................................................................................4

4. Rachis slender to filiform; corolla 9.5–13.5 mm long, tube 4.5–7.5 mm long, lobes narrowly oblong, 5–6 × 1.2 mm .................... 1. *N. gracilis*

4. Rachis stout; corolla 7–7.7 mm long, tube 4–4.5 mm long, lobes oblong, 3–3.2 × 1.1–1.2 mm............ 4. *N. pakaraimensis*

6-1. *Neobertiera gracilis* Wernham, J. Bot. 55: 169. 1917. (Figures 4J, 29–30).

Emend. Sandwith, Kew Bull. Misc. Inf. 1931: 470. 1931; emend. Steyermark, Mem. New York Bot. Gard. 17: 289–290. 1967; emend. Delprete, Phytotaxa 206. 127, fig. 2, 6. 2015.

Type: GUYANA. Macouria River, Nov. 1886 (fl), G.S. Jenman 2388 (holotype BM [barcode 000543796]; isotype K? n.v.).

*Perennial herbs or shrubs, 0.5–4 m tall, erect, sparsely branched; branches densely appressed strigose. Stipules* broadly to narrowly ovate, 3–4.5 × 2.5–3.5 mm, acuminate or bifid at apex, lobes narrowly triangular to linear, sometimes with an apical colleter, appressed strigose-pubescent outside, appressed antrorse-pubescent at medio-basal portion and intermixed with 5–6 colleters on each side inside, margins strigose. *Leaves* petiolate; petioles 3–9 mm long, appressed-strigose-pubescent; blades elliptic, oblong-elliptic to oblanceolate, 4.5–13.5 × 2–5 cm, acute-decurrent at base, acute at apex, chartaceous, drying pale brown above and olive-green below, sparsely appressed strigose above and below, midvein densely strigose above and below; secondary veins 9–14 on each side of midrib. *Inflorescences* laxly cymose during anthesis; peduncles 2–3.5 cm long; rachis slender to filiform, appressed white-pubescent, 5.5–7.5 cm long when inflorescences fully expanded, with 2–3 pairs of lateral branches; lower lateral branches 1.5–4 cm long (excluding corollas); lateral branches dichotomously divided, with one pedicellate flower at the base of each bifurcation, ultimate branches scorpiod when fully expanded; bracteoles opposite to the insertion of each flower, linear-lanceolate, 1.5–2 × 0.2–0.4 mm, appressed-pubescent. *Flowers* 4-5-merous, pedicellate; pedicels filiform, 0.5–1(–1.5) mm long. *Hypanthium* obovoid to broadly obovoid, 0.7–1.1 × 0.7–1.1 mm, appressed strigose. *Calyx* lobes frequently unequal, narrowly lanceolate, 2.5–5 × 0.2–0.3 mm, appressed strigose outside, and with 2, oblong, 0.1 mm long, colleters at each lobe sinus. *Corolla* white, 9.5–13.5 mm long; tube narrowly cylindrical, slightly wider at mouth, 4.5–7.5 mm long, appressed strigose-sericeous outside, puberulent inside, with a narrow line of yellow, moniform hairs at mouth inside; lobes narrowly oblong, 5–6 × 1–2 mm, acute at apex, glabrous on both sides. *Long-styled flowers* (Gleason 6 (US) measured on specimen): stamens inserted at 0.6–1 mm from base of corolla tube; filaments 0.3–0.5 mm long; anthers linear, 0.9–1.5 × 0.1–0.2 mm; style about the same length or slightly longer than the corolla tube, 5–7.5 mm long, glabrous; style branches narrowly oblong to linear, 0.8–1.5 mm long, acute at apex, densely papillose when receptive. *Short-styled flowers* (Grewal & Persaud 110 (U)): stamens inserted at distal portion of corolla, at 5.5–6 mm from base of corolla tube in flowers with corolla tube 7.5–8.5 mm long; filaments ca. 1 mm long; anthers linear, 1.5 × 0.2 mm, acute at both ends; style much short-
er than corolla tube, 4-4.5 mm long, glabrous; style branches narrowly oblong, 1.4 mm long, densely papillose when receptive. *Fruits* 2.5-3 mm diam., sparsely strigose-sericeous. *Seeds* ca. 0.3 mm long, microscopically foveolate (barely visible at 50× magnification).

**Notes:** Wernham (1917) in the introduction of the article where he described *Neobertiera gracilis* wrote "The types of all the novelties described are in the National Herbarium [BM]." Therefore the specimen *Jenman 2388* at BM, barcode 000543796, is the holotype of this name. He described the locality of the type specimen as "British Guiana, Macouria River (not far west of Cayenne), *Jenman 2388*." The Macouria River is in Guyana, but he confused this river with the village of Macouria, which is located, as he stated, "not far west of

Cayenne" and is in French Guiana. Therefore, the type specimen of *Neobertiera gracilis* was collected in Guyana, not French Guiana.

Delprete (2015a) cited the collections *Irwin et al. 54918, Schultz & Wessels Boer s.n.* (LBB 10223) and *Schultz J.P. s.n.* (LBB 10471) under *Neobertiera gracilis*; however, a further analysis of additional duplicates revealed that these collections have apically dehiscent capsules (which can be observed only in very advanced stage of maturity) and hypanthium and capsules with hairs basally tuberculate. Therefore, these three gatherings belong to *Sipanea stahelii*, which, although it belongs to another genus, is a very similar species. In addition, as these collections were studied for the description of *N. gracilis*, the above species description differs from that published by Delprete (2015a).

<figure><img src="image_34.png" /><figcaption>**Figure 30.** Distribution of *Neobertiera gracilis* (circles), *N. micrantha* (star), *N. montedouradensis* (triangles), *N. pakaraimensis* (squares), and *N. palustris* (inverted triangles).</figcaption></figure>
**Distribution and ecology:** Endemic to the Potaro-Siparuni Region, Guyana (Figure 30). It has been collected at the edge or inside wet forests, often near waterfalls, in alluvial or white-sand soils; at 25-120 m elevation. Delprete (2015a) stated that this species occurs in Guyana and Suriname; however, because of the re-identifications of the Suriname gatherings Irwin et al. 54918, Schultz & Wessels Boer LBB 10223 and Schultz LBB 10471 as Sipanea stahelii Bremek., this species is confirmed to be endemic to Guyana.

**Phenology:** Most specimens examined have flower buds, flowers in anthesis and mature fruits on the same inflorescence or at least on the same individual; these specimens were collected in April, June, July, August, September and November. The two specimens collected in June and July have flowers in full anthesis and some flower buds, but no fruits.

**Suggested conservation status:** Vulnerable (VU). This species is known from 19 collections from several localities in Guyana. Most of them are historical collections made from 1881 to the first half of 1990s. None of these localities are within protected areas, except one collection made in 1881 just below Kaieteur, which is now within the Kaieteur National Park. The size of the populations is unknown, and no recent observations in its natural environment are available. Because of this, Delprete (2015a) assigned this species to the category of Data Deficient (DD).

The extent of occurrence (EOO) of this species, using the collection localities with available or estimated coordinates is of ca. 19,700 km², and its most recent collection was made in 1990. Also, recent information (Delprete, pers. obs.) confirmed that within the EOO are present indiscriminate, often illegal, gold hunters' activities, either by mining, panning, or by dragging the bottom of rivers with dredgers. Because the localities where it has been collected are threatened by these human activities, this species is here assigned to the Vulnerable (VU) category, following IUCN criteria (IUCN 2012, 2019); however, future field studies might show that it is even more endangered.

**Specimens examined:** GUYANA: Rockstone, Jun. 1904 (fl, fr), A.B. Bartlett 8547 (K, U); Moraballi Creek [ca. 6°11'N, 58°50'W], Essequibo River, 3 Oct. 1938 (fl-fr), Forest Department of British Guiana F120 (Record N. 2729) (K [2 sheets]); Tumatumari [ca. 5°22'N, 59°0'W], 18-20 Jun. 1921 (fl), H.A. Gleason 6 [= Appun 331] (NY, US); Rockstone, 13-30 Jul. 1921 (fl), H.A. Gleason 483 (NY, US); Potaro River, Amatuk Mountain [ca. 5°18'N,

59°18'W], 1-2 Apr. 1958 (fl, fr), V. Graham 193 (K); Rockstone, Essequibo River, 25 m, 20 Apr. 1976 (fl), M.S. Grewal & R. Persaud 110 (U); Potaro River, below Kaieteur [ca. 5°10'N, 59°28'W], Sep.-Oct. 1881, G.S. Jenman 945 (K); Demerara River, Mar. 1898 (fl), G.S. Jenman 7338 (K); Potaro River, Garraway Stream [ca. 5°16'N, 59°9'W], 25 Apr. 1944 (fl, fr), B. Maguire & D.B. Fanshawe 22964 (F, MO, NY, S, U, US); Moraballi Creek [ca. 6°11'N, 58°50'W], Essequibo River, May 1932 (fl), E.B. Martyn 381 (K); Potaro-Siparuni Region, from camp on Elizabeth Creek, ca. 1 km along rd to NE and along creek, 5°18'N, 59°5'W, 120 m, 16 Oct. 1990 (fl, fr), T. McDowell & D. Goupal 3571 (MO, NY, US); Northwest District, vicinity of Matthews Ridge [ca. 7°30'N, 60°10'W], Forest Dept. Camp at White Creek, 8 km SSW of Matthews Ridge, weed at edge of recently cleared field, 31 Aug. 1976 (fl), S. Mori et al. 8261 (NY); Essequibo River, Moraballi Creek, near Bartica [ca. 6°24'N, 58°37'W], near sea level, 14 Sep. 1929 (fl), N.Y. Sandwith 256 (K, NY, RB, U); Basin of the Essequibo River, Head Falls, 6°7'N, 20 Sep. 1937 (fl), A.C. Smith 2106 (F, NY, U, US); Potaro road, Jun. 1910 (fl, fr), F.A. Stockdale 8775 (K); ca. 83 miles, Bartica-Potaro road, Amatuk Falls [ca. 5°18'N, 59°18'W], 26 Aug. 1933 (fl, fr), T.G. Tutin 602 (BM, K, US); Potaro River, Amatuk Mountain [ca. 5°18'N, 59°18'W], ca. 67 m, 23 Jul. 1959 (fl), B.A. Whitton 5 (K).

## 6-2. Neobertiera micrantha Delprete, Phytotaxa 206: 123, figs. 1, 2. 2015. (Figures 30-31).

**Type:** FRENCH GUIANA. Commune de Camopi, Upper Oyapock River Basin, Roche Touatou, small creek in forest, S slope of Roche Touatou (inselbergs), herb 0.5 m tall, stem hirsute, reddish-brown, erect, with age becoming decumbent and rooting at basal nodes, inflorescence terminal, corolla white, 2°57'N, 52°32'W, 120 m, 22 May 1995 (fl, fr), G. Cremers & J.J. de Granville 14097 (holotype CAY [barcode 079885]; isotypes CAY [barcode 079886], MO [Acc. No. 05000971, barcode MO-1508681], NY [barcode 03212820], P [barcode P007229247], U [barcode U010672], US [Acc. No. 3349483]).

*Perennial herbs*, 0.5 m tall, erect or basally decumbent, sparsely branched; branches densely appressed strigose; horizontal portion of stem rooting at basal nodes and internodes. *Stipules* oblong-ovate to broadly oblong-ovate, 4.5-6 × 4-5 mm, bifid at apex; each lobe narrowly triangular, ca. 2 mm long, appressed-strigose outside, basal portion appressed antrorse-pubescent intermixed with 5-6 colleters on each side and glabrous at distal por-
<figure><img src="image_35.png" /><figcaption>**Figure 31.** *Neobertiera micrantha*. A. Habit. B. Node with bifid stipule. C. Flower in anthesis. D. Dissected long-styled flower, with stamens inserted at lower portion of the corolla tube. Drawn from *Cremers & Granville 14097* (CAY). Illustration by Piero Delprete (Reproduced from fig. 1 of Delprete (2015a) with permission of copyright holder Magnolia Press).</figcaption></figure>
tion inside, margins strigose. *Leaves* petiolate; petioles 4–10(–1.5) mm long, sparsely appressed-strigose; blades narrowly elliptic, elliptic to ovate, 4.5–9 × 1.5–3.5 cm, acute-decurrent at base, acute to obtuse at apex, chartaceous, drying dark olive-green to brown above, pale olive-green to olive-green below, sparsely appressed short-strigose above, glabrous to sparsely sericeous below, midvein and secondary veins densely strigose above, densely strigose-sericeous below; secondary veins 8–11 on each side of midrib. *Inflorescences* always subcapitate during and after anthesis; peduncles 1.5–2 cm long; rachis thin, appressed white-pubescent; flowering portion capitate (extremely reduced cyme); bracts subtending the head lanceolate, 2–3 × 1–2 mm; bracteoles subtending each flower, linear-lanceolate, 1–1.5 × 0.2–0.5 mm, glabrous, margins appressed-strigose. *Flowers* 5-merous, pedicellate; pedicels thin, 0.5–1.3 mm long. *Hypanthium* globose to broadly obovoid, 0.5–0.7 × 0.5–0.7 mm, appressed strigose. *Calyx* lobes slightly unequal, lanceolate, 1.5–2.2 × 0.3–0.5 mm, appressed strigose outside, with 1, oblong, 0.1 mm long, colletter at each lobe sinus. *Corolla* white, 3.9–4.2 mm long; tube narrowly cylindrical, 3–3.2 mm long, sparsely appressed-sericeous outside, densely pubescent inside (hairs microscopically moniliform), with a narrow line of sparse, yellow, moniliform hairs at mouth inside; lobes 5, ovate, 0.9–1.0 × 0.5–0.6 mm, round at apex, glabrous on both sides. *Long-styled flowers:* stamens inserted at 0.9 mm from the base of corolla tube; filaments 0.4 mm long; anthers narrowly sagittate, 0.7 × 0.2 mm; style about the same length as the corolla tube (with branch tips barely exserted), 3–3.2 mm long, glabrous; style branches elliptic, 0.3 mm long, densely long-papilllose when receptive. *Short-styled flowers* unknown. *Fruits* 1.2–1.5 mm diam., sparsely strigose-sericeous. *Seeds* ca. 0.3–0.4 mm long, pale brown, microscopically foveolate (barely visible at 50× magnification).

**Distribution and ecology:** Known only from the type gathering, collected in 1995 near Roche Touatou (Figure 30), a group of small inselbergs located on the Upper Oyapock River Basin, French Guiana, near a small creek, in the undercanopy of primary forest, at 120 m altitude.

**Phenology:** Specimens with flowers and fruits were collected in May.

**Suggested conservation status:** Critically Endangered (CR). Known by a single collection near Roche Touatou, Upper Oyapock River Basin, French Guiana. The number of individuals of this population was not reported by the collectors. Due to the remote locality and the unknown number of individuals of the population, Delprete (2015) treated the conservation status of this species as Data Deficient (DD).

Roche Touatou is a group of granitic inselbergs, ca. 2°57′N, 52°32′W, situated at about 35 km SW of the town of Camopi, a community of about 1800 inhabitants. The town is within the zone of free adhesion of the French Guiana Amazonian Park. The region surrounding Camopi is mostly covered by tropical rainforest, with deforested patches of land cultivated by the local population. In addition, such a region is subject to illegal gold mining, which, aside from the destruction of natural vegetation, also cause mercury pollution of fresh water and aquatic fauna, both of them vital sources of the local population. As this species is known by a single collection in forest undercanopy, and in view of the threat of illegal gold mining, this species is Critically Endangered (CR) following IUCN criteria (IUCN 2012, 2019).

6-3. *Neobertiera montedouradensis* Delprete, Phytotaxa 392: 226, fig. 1. 2019. (Figures 30, 32).

Type: BRAZIL. Pará: Mun. Almeirim, Monte Dourado, Estrada Perimetral, mata de terra firme, solo argiloso, [ca. 0°52'S, 52°33'W], 27 Nov. 1978 (fl), *M.R. Santos 414* (holotype MG [Acc. No. 72269]).

*Shrub*, 0.4–3.5 m tall; branches sparsely appressed golden-pubescent. *Stipules* narrowly triangular, 2.5–4.5 × 1.7–2 mm, entire (not lobed), acute at apex, appressed-strigose outside, glabrous, with a row of golden hairs and a row of colleters at base inside, margins golden-pubescent ciliate. *Leaves* petiolate; petioles 7–17 mm long, densely appressed golden-pubescent; blades broadly ovate to elliptic, 3.5–8.7 × 1.9–3.5 cm, round, obtuse to acute-decurrent at base, acuminate at apex, chartaceous, drying dark brown above and rust-brown below, sparsely appressed strigose above and below, midvein and secondary veins densely appressed golden-pubescent below;

**Figure 32.** *Neobertiera montedouradensis.* A. Branch with a young, unexpanded inflorescence. B. Branch with a fully expanded inflorescence. C. Node with entire stipule. D. Dissected long-styled flower, with stamens inserted at lower portion of the corolla tube, and exserted style. E. Longitudinal section of flower, with corolla removed; note stalked placenta and hypanthium with basally tuberculate hairs. F. Basal portion of one hypanthium hair, longitudinal section. A–F: drawn from *M.R. Santos 414* (MG). Illustration by Piero Delprete (Reproduced from fig. 1 of Delprete (2019b) with permission of copyright holder Magnolia Press).
<figure><img src="image_36.png" /><figcaption>Figure 32.</figcaption></figure>
secondary veins 8-9 each side of midrib, complanar or slightly depressed above, prominent below. *Inflorescence* densely cymose at initial stage of anthesis, expanding to compound laxly cymose during anthesis; peduncles initially 2-3 cm long and expanding to 4.7 cm long during final stage of anthesis; rachis stout to filiform, sparsely appressed golden-pubescent, 10-15.5 cm long when fully expanded, lateral branches 3 times dichomously divided, with one sessile to subsessile flower at the base of each bifurcation, ultimate branchlets scorpiod when fully expanded, 7-10-flowered; bracts subtending secondary and tertiary branches linear-lanceolate, 2.7-4 × 0.3-0.4 mm, sparsely appressed-pubescent; bracteoles opposite to the insertion of each flower, linear-lanceolate, 1.7-2.2 × 0.2-0.3 mm, sparsely appressed-pubescent. *Flowers* 5-merous, subpedicellate; pedicels 0.3-0.4 mm long, appressed strigose. *Hypanthium* globose to obovoid, 0.8-0.9 × 0.8-0.9 mm, densely appressed strigose; hairs basally tuberculate, basal tubercules multicellular. *Calyx* lobes frequently unequal, linear-lanceolate to narrowly lanceolate, 2.2-2.6 × 0.3-0.5 mm, sparsely appressed strigose outside, and with 1, oblong, 0.2 mm long, collector at each lobe sinus. *Corolla* pink (see Notes, below), 14-15.5 mm long; tube narrowly cylindrical, slightly wider at medio-distal portion, 9.5-10.5 mm long, sparsely appressed pubescent outside, glabrous at base, densely spreading pubescent at portion surrounding stamens and sparsely appressed pubescent at distal portion inside, glabrous at mouth; lobes 5, ovate to oblong-ovate, 4.5-5.2 × 2.2-2.7 mm, acute at apex, glabrous outside, papillose inside. *Long-styled flowers*: stamens inserted 3 mm from the base of corolla tube; filaments 0.6 mm long; anthers linear, 1.8 × 0.2 mm, acute at both ends; ovary 2-locular, placenta stalked, inserted at medio-basal portion of the septum; ovules many, inserted on the placental extension; style slightly longer than the corolla tube, ca. 13 mm long, glabrous, style branches almost completely exerted, linear, 1.5-1.8 mm long, acute at apex, densely papillose when receptive. *Short-styled flowers* (Pires & Silva 110 (HAMAB)): stamens inserted at distal portion of corolla, at 3.5 mm below corolla mouth; filaments 0.5 mm long; anthers linear, 1.7 × 0.2 mm, round at both ends; style much shorter than corolla tube, ca. 5 mm long, glabrous; style branches narrowly oblong-lanceolate, 1.5 mm long, densely papillose when receptive. *Fruits* 2.5-3 mm diam., densely appressed strigose; hairs basally tuberculate, basal tubercules multicellular. *Seeds* unknown.

**Notes:** *Neobertiera montedouradensis* Delprete is unique within the genus by having hypanthium and fruit with hairs basally tuberculate (Fig. 32E-F). Hypanthia and fruits with basally tuberculate hairs are present

in a few species of *Sipanea*, e.g., *S. stahelii* and *S. wilsonbrownei*. While the fruits of *Neobertiera* are indehiscent, those of *Sipanea* are loculicidal capsules.

On the specimen label of *M.R. Santos 414*, the flowers of *N. montedouradensis* are described as orange ["flor alaranjada"]; however, this is probably erroneous, as the labels of *Sarquis et al. 62* and *Forza & Leal 5928* indicate that the corollas of this species are pink.

*Neobertiera montedouradensis* is most similar to *N. pakaraimensis* because of its leaf blades chartaceous, with 8-10 secondary veins on each side of midvein, and inflorescences compoundly cymose during anthesis; the former differs from the latter by the narrowly triangular, entire stipules (vs. ovate to broadly ovate, bifid in *N. pakaraimensis*), leaf blades broadly ovate to elliptic, 3.5-8.7 × 1.9-3.5 cm, round, obtuse to acute-decurrent at base, acuminate at apex (vs. narrowly elliptic, lanceolate to narrowly oblong-elliptic, 4-10 × 0.7-2.5 cm, acute-decurrent at base, acute and acuminate at apex), hypanthium with hairs basally tuberculate (vs. with hairs not basally tuberculate), corollas 14-15.5 mm long, with tube 9.5-10.5 mm long, and lobes ovate to oblong-ovate, 4.5-5.2 × 2.2-2.7 mm, acute at apex (vs. 7-7.7 mm long, with tube 4-4.5 mm long, lobes oblong, 3-3.2 × 1.1-1.2 mm, round to obtuse at apex).

**Distribution and ecology:** Originally described from a single specimen collected near the town of Monte Dourado, state of Pará, in terra firme forest on clay soil, near the Jari River, a river running between the Brazilian states of Pará and Amapá. Three additional gatherings have recently been found in HAMAB and MG, one collected near the type locality, and the other two from the contiguous state of Amapá, near the Jari River, and in the Serra do Navio, in the center of the state (Figure 30); all these collections were also made at the edge of a terra firme forest.

**Phenology:** Three gatherings with flowers and fruits were collected in June and August, and one with only flowers was collected in November.

**Suggested conservation status:** Critically Endangered (CR). This species was originally described from a single specimen collected near the town of Monte Dourado [ca. 0°53'S, 52°36'W], near the Jari River, state of Pará. An expedition to the town of Monte Dourado was made during the first week of November 2018 to search for this species. Although a considerable portion of the forests surroundings this town was destroyed and replaced by eucalyptus plantations, several patches of well-preserved forests are still present in a radius of sev-
eral kilometers from the town; nevertheless, after three
days of extensive searches in numerous primary and sec-
ondary forests of this region, the species was not found.
Along with its original description Delprete (2019b) sug-
gested a conservation status of Critically Endangered
(CR) following IUCN criteria (IUCN 2012, 2017).

Three additional gatherings of this species have
recently been found in HAMAB and MG, one from
a locality between Monte Dourado and Almerim, at
1°15'6"S, 52°38'10"W, state of Pará, and the other two
from the contiguous state of Amapá, at ca. 0°53'S,
52°22'W and 0°55'N, 51°52'W, extending the extent of
occurrence of the species (EOO) to ca. 5,600 km². The
number of individuals are not reported in the specimen
labels. Taking into account the aggressive forest destruc-
tion still ongoing around the town of Monte Dourado
and in the other localities in the state of Amapá, it is dif-
ficult to predict how long these forest patches will last.
Therefore, this rare species is maintained as Critically
Endangered (CR) following IUCN criteria (IUCN 2012,
2019).

**Specimens examined:** **BRAZIL.** *Amapá:* Mun. Mazagão, margem esquerda do Rio Jari, Morro do Felipe IV [ca. 0°53'S, 52°22'W], beira da floresta mis- ta primária-secundária, 16 Aug. 1985 (fl), *M.J.P. Pires & N.T. Silva* 546 (HAMAB); Serra do Navio, Miner- ação Pedra Branca do Amapari, Ramal Serra da Canga, 0°55'N, 51°52'W, arbusto aproximadamente de 2 m de altura, flores roxas claras, 29 Aug. 2003 (fl, fr), *R.S.F.R. Sarquis et al.* 62 (HAMAB [2 sheets], IAN, MG). **Pará:** Mun. Almeirim, Distrito de Monte Alegre, Área de Manejo da Orsa Florestal, Platô 2, floresta de terra firme, borda de mata, 1°15'6"S, 52°38'10"W, 250 m, arbusto 40-60 cm, ramos e nervuras vináceos, cálice verde, corola rósea, frutos verdes, 31 Jun. 2010 (fl, imm fr), *R.C. Forzza & E.S. Leal* 5928 (K n.v., MG, RB).

6-4. *Neobertiera pakaraimensis* Delprete, Phytotaxa 206: 126, figs. 2, 5. 2015. (Figures 30, 33).

Type: GUYANA. Potaro-Siparuni Region, Pakarai-
ma Mountains, Mount Wokomung, across slope to
SW 0.5 km from headwaters of Wusupubaru Creek,
montane slope forest, W aspect, upper canopy 20-35
m, 2-storied, epiphytes abundant, incl. 0.5 hectare
tree fall, shrub 0.5 m [tall], filaments white, 5°3'N,
59°53'W, 975-1125 m, 17 Feb. 1993 (fl), *T.W. Henkel*,
*M. Chin & W. Ryan* 1449 (holotype CAY [barcode
079485]; isotypes BBS [barcode BBS0032627], CAY
[barcode 079486], F [Acc. No. 2232663], INPA [Acc.

No. 214891], MO [Acc. No. 04926765], NY [bar-
code 03212819], U [barcode 0248751], US [barcode
00699048]].

Shrubs, 0.5-4 m tall, erect or basally decum-
bent, sparsely branched; branches sparsely to densely
appressed strigose; rooting at basal nodes and inter-
nodes. *Stipules* ovate to broadly ovate, 4-5.5 x 2.5-3
mm, bifid at medio-distal portion, appressed-strigose
outside, basal portion antrorse-pubescent intermixed
with 2-3 colleters on each side and glabrous at dis-
tal portion inside, margins pubescent. *Leaves* petiolate;
petioles 6.5-11.5 mm long, densely appressed-strigose;
blades narrowly elliptic, lanceolate to narrowly oblong-
elliptic, 4-10.5 x 0.7-2.5 cm, acute-decurrent at base,
acute or acuminate at apex, chartaceous, drying dark
brown above and olive-green below, sparsely appressed
strigose above, sparsely short- to long-strigose below,
midvein and secondary veins densely strigose below;
secondary veins 8-10 each side of midrib, evident only
below, immersed in the lamina above. *Inflorescence*
laxly cymose during anthesis; peduncles 1.7-4 cm long;
rachis stout, appressed white-pubescent, 5.5-6.5 cm long
when fully expanded, with 1-3 pairs of lateral branches;
lower lateral branches 2.5-3.5 cm long (excluding corol-
las); lateral branches dichotomously divided, with one
pedicellate flower at the base of each bifurcation, ulti-
mate branches scorpiod when fully expanded; bracteoles
opposite to the insertion of each flower, linear-lanceo-
late, 1.5-2 x 0.2-0.4 mm, appressed-pubescent. *Flowers*
4-5-merous, pedicellate; pedicels 1-1.5 mm long, densely
appressed strigose. *Hypanthium* globose to obovoid, ca.
1 x 1 mm, appressed strigose. *Calyx* lobes frequently
unequal, narrowly lanceolate to narrowly oblong-lanceo-
late, 2-3 x 0.3-0.4 mm, appressed strigose outside, and
with 1 oblong, 0.1 mm long, colleter at each lobe sinus.
*Corolla* white, 7-7.7 mm long; tube narrowly cylindri-
cal, slightly wider at mouth, 4-4.5 mm long, appressed
strigose-sericeous outside, puberulent inside, with a nar-
row line of yellow, moniliform hairs at mouth inside;
lobes 5, oblong, 3-3.2 x 1.1-1.2 mm, round to obtuse at
apex, glabrous on both sides. *Long-styled flowers:* sta-
mens inserted 1.8 mm from the base of corolla tube;
filaments 0.7-1 mm long; anthers linear, 1.2 x 0.2 mm,
acute at both ends; style slightly longer than the corolla
tube, 4.5-5 mm long, glabrous; style branches partially
exserted, narrowly oblong, 0.9-1 mm long, acute at apex,
densely papillose when receptive. *Short-styled flowers:* unknown. *Fruits and seeds* unknown.

**Notes:** Specimens of this taxon have incorrectly been
identified as *Sipaneopsis cururuensis* (Henkel et al. 1449,
<figure><img src="image_37.png" /><figcaption>**Figure 33.** *Neobertiera pakaraimensis*. A. Branch with a young inflorescence. B. Branch with a fully developed inflorescence. C. Node with bifid stipule. D. Dissected long-styled flower, with stamens inserted at the middle of the corolla tube. A–D: drawn from *Henkel et al. 1449* (CAY). Illustration by Piero Delprete (Reproduced from fig. 5 of Delprete (2015a) with permission of copyright holder Magnolia Press).</figcaption></figure>