Byrsonima Richard

Alves, Livia Lara, Francener, Augusto, Costa, Maria Tereza R. & Sobral, Marcos, 2021, Byrsonima (Malpighiaceae) from Espírito Santo state, Brazil, Phytotaxa 511 (1), pp. 1-19 : 2-10

Byrsonima Richard ex Kunth (1822: 147)

Trees, shrubs or subshrubs, branches aerial or underground. Stipules epipetiolar free or connate. Leaves opposite; petioles eglandular; blades glabrous or pilose, eglandular, with V-, Y-, T- or star-shaped trichomes. Thyrse sessile or pedunculate, cincinnus 1–2-flowered. Sepals all 2-glandular. Petals yellow to orange or red, pink or white, glabrous. Stamens 10, all fertile, free or connate at base, pilose at base; connectives glandular, surpassing or not the pollen sacs, anthers sericeous to glabrous. Ovary glabrous or sericeous, styles 3, glabrous, apex acute, 3 terminal stigmas. Drupes, exocarp fleshy, green when immature, yellow or red when mature.

Key to the species of Byrsonima in Espírito Santo state

1. Leaves glabrous to glabrescent on both sides, with few or no trichomes in the midrib or lateral ribs, abaxial side glabrous to glabrescent, with a few sericeous hairs in midrib, adaxial side glabrous...........................................................................................2

-. Leaves tomentose, velutine or sericeous on both sides or abaxially ................................................................................................13

2. Anthers glabrous to glabrescent .........................................................................................................................................................3

-. Anthers sericeous................................................................................................................................................................................7

3. Stipules up to 2.3 mm long; main axis of inflorescence up to 5.6 mm long ......................................................................................4

-. Stipules longer than 2.2 mm long; main axis of inflorescence longer than 5.6 mm long ..................................................................5

4. Floral rachis glabrous, bearing ca. 20 flowers; ovary glabrous to sericeous, leaves obovate, oblong to elliptical, all the petals white to pink becoming red with age ....................................................................................................................................... B. myricifolia

-. Floral rachis sericeous, bearing ca. 50 flowers; ovary glabrous; leaves narrowly elliptical to lanceolate; lateral petals white becoming pink in age and posterior petal or claw of posterior petal yellow, becoming red with age ......................... B. ligustrifolia

5. Leaves elliptical; apex acute, acuminate falcate to rounded .......................................................................................... B. perseifolia

-. Leaves obovate to widely elliptical; apex rounded to retuse..............................................................................................................6

6. Petiole ca. 13 mm; connectives slightly surpassing the anther sacs (ca. 0.1 mm length); accrescent sepals on the drupes ................ ............................................................................................................................................................................................ B. bahiana

-. Petiole ca. 7 mm; connectives visibly surpassing the anther sacs (ca. 0.7 mm length); sepals not accrescent on the drupes ............. .................................................................................................................................................................................... B. niedenzuiana

7. Bracts reflexed....................................................................................................................................................................................8

-. Bracts non-reflexed ............................................................................................................................................................................9

8. Petals yellow becoming red to orange in age; bracts and bracteoles reflexed,; sepals accrescent on the fruits (1.5–2 mm on the flower); drupes sericeous at the apex .................................................................................................................................... B. crispa

-. Petals white becoming pink in age; bracts reflexed and bracteoles non-reflexed; sepals not accrescent on the fruits; drupes glabrous ................................................................................................................................................................................ B. alvimii

9. Thyrses 1–2 flowered cincinnus; drupes larger than 12 mm diam................................................................................ B. cacaophila

-. Thyrses 1-flowered cincinnus; drupes less than 12 mm diam..........................................................................................................10

10. Shrubs ca. 1 m tall; petals yellow turning orange to red in age...................................................................................... B. intermedia

-. Small trees or trees higher than 1 m tall; petals white to pink .........................................................................................................11

11. Leaves widely elliptical to ovate; apex rounded; petiole absent (sessile) or up to 1.5 mm long ............................. B. coccolobifolia

-. Leaves elliptical, oblong, to obovate; petiole longer than 4 mm long .............................................................................................12

12. Leaves oblong; drupes up to 8 mm diam......................................................................................................................... B. nitidifolia

-. Leaves elliptical to obovate; drupes ca. 12 mm diam ..................................................................................................... B. Japurensis

13. Drupes less than 7 mm diam; abaxial side of leaves densely sericeous to sericeous (hairs appressed)............................... B. sericea

-. Drupes larger than 7 mm; abaxial side of leaves tomentose or velutine (hairs stalked) or few trichomes appressed (slightly sericeous)..........................................................................................................................................................................................14

14. All petals white becoming pink in age; ovary glabrous ................................................................................................... B. vernicosa

-. All petals yellow becoming orange in age; ovary sericeous ............................................................................................................15

15. Stipules 5–25 mm long, amplexicaul, deciduous; foliar lamina bearing stellate-stalked hairs....................................... B. stipulacea

-. Stipules up to 6,5 mm long, not amplexicaul, persistent; foliar lamina bearing shortly-stalked T-shaped hairs .............................16

16. Stipules narrowly triangular to ovate; style 2.6–4.1 mm long ....................................................................................... B. crassifolia

-. Stipules broadly ovate; style 4–4.8 mm long .................................................................................................................. B. fanshawei

. Byrsonima alvimii W.R. Anderson (1982: 11) . Figure 1A

Trees, 5–22 m tall. Stipules 2–9.8 × 0.8–5.6 mm, ovate, lanceolate to triangular. Leaf blades 3.75–23.7 × 1–9.8 cm, obovate to elliptic, base attenuate to cuneate, margins flat to slightly revolute, apex acute, acuminate, rounded, apiculate, retuse to mucronulated, both side glabrous to glabrescent, with few tricomas on midrib, midrib prominulous on abaxial side, impress to prominulous on adaxial side, principal laterals veins impress on both side and number of ca. 8; petiole 9–18 × 1.3–2.5 mm, glabrous, glabrescent to slightly sericeous. Thyrse of 1-flowered cincinnus, 11–38 flowers, main axis 2.3–13.8 cm long, sericeous; bract 1.4–7.2 × 1–2.5 mm, lanceolate, ovate to triangular, both side glabrous, glabrescent to sericeous, reflexed; peduncle 0–5.6 × 0.6–1 mm; bracteoles 0.8–4.3 × 0.6–2.5 mm, lanceolate, ovate to triangular, non-reflexed, pedicels 3.5–21.4 × 0.7–2 mm, sericeous to tomentose; sepals 2.1–7.6 × 1.5–4.4 mm, abaxial side tomentose to sericeous, adaxial side glabrous, glabrescent to sericeous, margin slightly revolute at anthesis, triangular, ovate to widely ovate, not accrescent on the drupes; glands 1.3–5.3 × 0.8–2.7 mm; all petals white becoming pink in age, lateral petals blade ca. 4.6 × 6.7 mm, claw ca. 3.3 × 0.7 mm, margin slightly erose to entire; posterior petal blade ca. 2.7 × 4.2 mm, claw ca. 2.7 × 0.7 mm, margin erose. Stamens free at base, homomorphic, anthers sericeous, connectives 1.6–2.3 × 0.3–0.8 mm, slightly surpassing the anther sacs (up to 0.4 mm length); filaments ca. 2.3 × 0.6 mm, sericeous at base. Ovary sericeous, style ca. 2.4 × 0.2 mm. Drupes 6.6–13.4 × 3.8–13.2 mm, glabrous, verrucous, globose.

FIGURE: A. Byrsonima alvimii (Kollmann 2279, MBML) B. Byrsonima bahiana (Oliveira 792, VIES), C. Byrsonima fanshawei (Folli 7555, HUFSJ), D. Byrsonima Japurensis (Demuner 59, MBML)

Specimens examined:— BRAZIL. Espírito Santo. Guarapari: Parque Estadual de Setiba , 6 June 1991, fr., L.V. Rosa 111 ( VIES!) ; Parque Estadual Paulo César Vinha , 2 July 1999, fr., A.M. Assis 710 ( VIES!) ; L.C. Fabris 739 (VIES!); ibidem, 25 November 1999, bud, fl., A.M. Assis 755 ( VIES!) ; Parque Estadual Paulo César Vinha , 4 May 2009, fr., G.E. Valente 2597 ( VIC) ; Setiba , 18 March 1991, fr., J.M.L. Gomes 1463 ( VIES!) ; Setiba , 28 November 1990, fl., O.J. Pereira 2285 ( VIES!) . Santa Teresa: Aparecidinha, Terreno de Luis Bringhenti , 750 m, 6 October 1998, fl., L. Kollmann 666 ( MBM, MBML!) ; ibidem, 9 November 1998, fl., L. Kollmann 905 ( MBML!) ; ibidem, 29 March 1999, fr., L. Kollmann 2279 ( MBML!) ; Estação Biológica Santa Lúcia , 650–800 m, 27 July 1993, st., L.D. Thomaz 982 ( MBML!, VIES!) ; Nova Lombardia, Reserva Biológica Augusto Ruschi , 870 m, 21 January 2003, fr., R.R. Vervloet 1682 ( MBML!) ; Santo Antônio, Terreno do Boza , 750 m, 7 October 1998, fl., L. Kollmann 708 ( MBM, MBML!) ; ibidem, 29 October 1998, fl., L. Kollmann 876 ( MBML!) ; ibidem, 800 m, 8 December 1998, fr., L. Kollmann 1256 ( MBML!) ; Santo Henrique , beira de estrada, 26 January 2005, fr., L. Kollmann 7327 ( MBML!) ; Vila Velha: Reserva Ecológica de Jacarenema , 28 June 2000, fr., O.J. Pereira 6229 ( VIES!) .

Notes:— Byrsonima alvimii is endemic to Brazil, occurring only in the states of Bahia and Espírito Santo in the domain of the Atlantic Forest ( Francener 2020). According to the National Flora Conservation Center (CNCFlora 2021), the species is categorized as vulnerable. It is characterized by only bracts reflexed (bracteoles are non-reflexed), deciduous in the maturity, drupes verrucous, leaves glabrous to glabrescent on both sides, with few trichomes on midrib, and margin entire. It resembles B. cacaophila , but this last one has a thyrse of 1–2 flowered cincinnus, bracts, and bracteoles smaller (up to 2.4 mm long) and its drupes are smooth .

. Byrsonima bahiana W.R. Anderson (1982: 110) . Figure 1B

Trees, ca. 10 m tall. Stipules ca. 2.2 × 1.2 mm, triangular. Leaf blades ca. 7 × 4.5 cm, obovate to widely elliptic, base cuneate, margins flat, apex rounded to retuse, both sides glabrous, midrib prominulous on abaxial side, impress on adaxial side, principal laterals veins ca. 10, impress on both sides; petioles ca. 13.6 × 2.3 mm, glabrous. Thyrse of 1-flowered cincinnus with ca. 25 flowers, main axis ca. 9.2 cm long, glabrescent; bracts ca. 1.7 × 0.7 mm, triangular, abaxial side glabrous, adaxial side glabrescent, non-reflexed; peduncle absent; bracteoles ca. 1.2 × 1 mm, ovate, nonreflexed; pedicels ca. 8.5 × 1.2 mm, glabrous. Flowers not seen [according to Anderson (1981) the sepals ca. 2.5 × 3 mm, both sides glabrous except for tiny hairs on the margin, margin revolute at anthesis, rounded, accrescent on the drupes; glands ca. 2.5–3 mm long; all petals white, often tinged with pink, with reddish claw. Anthers glabrous; connectives slightly surpassing the anther sacs (up to 0.1 mm in length); filaments 2–3 mm long, sericeous at base; Ovary glabrous]. Drupes ca. 10 × 8.8 mm, glabrous, verrucous, globose.

Specimen examined:— BRAZIL. Espírito Santo. Conceição da Barra: Itaúnas, Parque Estadual de Itaúnas , Fazenda Jequitaia , 1 May 2010, fr., A.G. Oliveira 792 ( VIES!) .

Notes: — Byrsonima bahiana is endemic of Brazil. Occurs in the states of Bahia, Espírito Santo, Sergipe and Alagoas, in the domain of the Atlantic Forest ( Francener 2020). According to the National Flora Conservation Center (CNCFlora 2021), is categorized as near threatened. Byrsonima bahiana has accrescent fleshy sepals on the drupes, the reddish color on the leaves margins and the midrib, and bracts and bracteoles deciduous after anthesis. In addition, the sepals on both sides are glabrous except for tiny hairs on the margin. It resembles B. alvimii , but the last has reflexed bracts, abaxial side of the sepals tomentose to sericeous and sepals not accrescent on the drupes.

. Byrsonima cacaophila W.R. Anderson (1982: 98) . Figure 2A

Trees, 7–26 m tall. Stipules 1.5–3.2 × 1–2.3 mm, triangular to narrowly triangular. Leaf blades 7.8–15.6 × 3.7–7.6 cm, obovate to elliptic, base attenuate to cuneate, margins flat to slightly revolute, apex acute, acuminate to rounded, abaxial side glabrous, glabrescent, with few tricomas on midrib, adaxial side glabrous, midrib prominulous on both side, principal laterals veins impress on both side and number of ca. 9; petiole 8.2–18.5 × 0.9–3 mm, glabrous, glabrescent to slightly sericeous. Thyrse of 1–2 flowered cincinnus, 35–60 flowers, main axis 5.27–9.1 cm long, tomentose to densely tomentose; bracts 0.9–2.4 × 0.6–1.4 mm, lanceolate to triangular, abaxial side sericeous to tomentose, adaxial side glabrous to glabrescent, non-reflexed; peduncle 2.3–4.4 × 0.5–1.2 mm; bracteoles 1–1.4 × 0.6–1.2 mm, triangular, non-reflexed; pedicels 3.7–7.7 × 0.5–1.3 mm, densely tomentose to tomentose; sepals 1.9–3 × 1.6–2.4 mm, abaxial side tomentose to sericeous, adaxial side glabrous to glabrescent, margin flat at anthesis, triangular to ovate, not accrescent on the drupes; glands 1.4–2.5 × 0.4–1.5 mm; all petals white becoming pink in age, lateral petals blade 3.6–7.5 × 4–5.5 mm, claw 1.7–2.7 × 0.4–0.7 mm, margin slightly erose to erose; posterior petal blade 2.6–3.5 × 1.6–3.2 mm, claw 2.2–3.2 × 0.7–0.9 mm, margin erose. Stamens free at base, homomorphic, anthers sericeous, connectives 1.5–2.8 × 0.4–0.9 mm, surpassing the anther sacs (0.1–0.5 mm length); filaments 0.8–2 × 0.3–0.5 mm, sericeous at base. Ovary sericeous, style 2.3–3 × 0.3–0.4 mm. Drupes 10–18 × 12–14 mm, glabrous, glabrescent to slightly sericeous, smooth, globose.

Specimens examined:— BRAZIL. Espírito Santo. Linhares: ES 440 Linhares a Regência , 20 January 2014, bud, fl., D.A. Folli 7175 ( CVRD!, RB!); Reserva Natural da Vale , 02 February 1978. bud, fl., J. Spada 38/78 ( CVRD!, HUFSJ!); ibidem, 18 May 1995, fr., D.A. Folli 2615 ( CVRD!, HUFSJ!); ibidem, 13 January 2010, bud, fl., G.S .

Siqueira 517 (CVRD!, HUFSJ!); ibidem, 24 January 2005, bud, fl., D.A. Folli 5019 ( CVRD!, HUFSJ!); ibidem, 18 April 2013, fr., G.S. Siqueira 882 ( CVRD!, HUFSJ!); Reserva Natural da Vale, estrada Jacarandá-Caveuna , 41 m, 16 April 2011, fr., J.C. Lopes 174 ( RB!) . Marilândia: Liberdade, prop: Reinaldo Bautz , 19°19’57” S, 40°32’5” W, 400–500 m, 27 September 2006, fr., L.F.S. Magnago 1432 ( MBML!) .

FIGURE: A. Byrsonima cacaophila (photo by F. Flores), B. Byrsonima coccolobifolia (photos by L.L. Alves), C. Byrsonima crassifolia (photo by C.F. Hall), D. Byrsonima crispa , detail of bracts and bracteoles strongly reflexed (photo by C.F. Hall), E. Byrsonima intermedia (photo by L.L.Alves).

Notes: — Byrsonima cacaophila is endemic of Brazil. It occurs only in the states of Bahia and Espírito Santo in the domain of the Atlantic Forest ( Francener 2020). According to the National Flora Conservation Center (CNCFlora 2021), B. cacaophila is considered near threatened. It is characterized by the presence of a large peduncle (ca. 3 mm long), thyrse 1–2 flowered cincinnus, large petiole (ca. 10 mm long), bracts not-reflexed, drupes ca. 14 mm long. The exsiccate examined with flowers had a sericeous ovary, but they are at variance with the original description ( Anderson 1982). B. cacaophila resembles B. perseifolia and B. alvimii as mentioned above.

. Byrsonima coccolobifolia Kunth (1822: 148). Figure 2B

Small trees, 1–2 m tall. Stipules ca. 2.6 × 1.2 mm, triangular. Leaf blades ca. 6.6 × 4.2 cm, widely elliptic to ovate, base subcordate to cuneate, margins flat, apex rounded, both sides glabrous, midrib prominulous on both sides and principal laterals veins impress on both sides and number of ca. 6; petiole 0–1.5 × 0–2.3 mm, glabrous. Thyrse of 1-flowered cincinnus, main axis ca. 6.5 cm long, densely tomentose; bract ca. 2 × 0.7 mm, lanceolate, abaxial side slightly sericeous, adaxial side glabrous to glabrescent, non-reflexed; peduncle absent; bracteoles ca. 1.5 × 0.8 mm, triangular, non-reflexed; pedicels ca. 2.5 × 0.7 mm, sericeous; sepals ca. 2.4 × 1.5 mm, both sides sericeous, margin slightly revolute at anthesis, ovate, not accrescent on the drupes; glands ca. 1.2 × 0.7 mm. all petals white to pink becoming pink in age; lateral petals blade ca. 4.5 × 4 mm, claw ca. 2.7 × 0.5 mm, margins entire; posterior petal blade ca. 2.5 × 4.6 mm, claw ca. 4.3 × 0.6 mm, margins entire. Stamens free at base, homomorphic; anthers sericeous; connectives ca. 1.8 × 0.8 mm, not surpassing the anther sacs; filaments ca. 2.6 × 0.7 mm, densely sericeous at base. Ovary glabrescent; styles ca. 4.8 × 0.3 mm. Drupes not seen [according to Anderson (1981), 7–8 mm diam, glabrous, globose or ovoid].

Specimens examined:— BRAZIL. Espírito Santo. Conceição da Barra: Área 157 da Aracruz Celulose S.A., 26 March 1992, fl., O.J. Pereira 3200 ( VIES!); Parque Estadual de Itaúnas , 26 April 2000, bud., O.J. Pereira 6157 ( VIES!) .

Additional specimens examined: BRAZIL. Minas Gerais, São João del-Rei: loteamento Pinheiros , distrito do Rio das Mortes, 21º19’89” S, 44º34’42” W, 26 November 2017, bud., fl., L.L. Alves 227 (HUFSJ) .

Notes: — Byrsonima coccolobifolia occurs from Cuba descending from South America to Paraguay, passing through Colombia, Peru, Venezuela, and Brazil. In Brazil is widely distributed in all regions, in the domain of the Atlantic Forest, Amazonia, and Cerrado ( Francener 2020). It is characterized by its leaves widely elliptic to ovate, base subcordate to cuneate, margins flat, apex rounded, usually midrib, and principal laterals veins pink, petiole absent or small (ca. 1.5 mm), bracts not reflexed. This species does not resemble any other of the 16 species in the study area.

. Byrsonima crassifolia ( Linnaeus 1753: 426) Kunth (1822: 149). Figure 2C

Trees, 3–25 m tall. Stipules 2.8–6.4 × 2–4.5 mm, triangular, narrowly triangular to ovate, not amplexicaul, persistent. Leaf blades 7.9–16 × 3.7–7.4 cm, obovate, ovate to elliptic, base attenuate to cuneate, margins flat to slightly revolute, apex acute, acuminate, rounded to apiculate, abaxial side of the leaves tomentose when young but deciduous with maturity, adaxial side glabrous, glabrescent to tomentose on midrib, but deciduous with maturity, shortly-stalked Tshaped hairs, midrib prominulous on abaxial side, impress on adaxial side, principal laterals veins prominulous on abaxial side, impress on adaxial side and number of ca. 8; petiole 9–14 × 1.5–2.5 mm, glabrescent to sericeous. Thyrse of 1–2-flowered cincinnus, 35–60 flowers, main axis 6.5–15.2 cm long, glabrescent, tomentose to sericeous; bract 1.7– 3.7 × 1–1.4 mm, triangular, abaxial side sericeous, adaxial side glabrous to glabrescent, non-reflexed; peduncle 0–2.2 × 0–1.6 mm; bracteoles 1.4–2.1 × 0.6–0.9 mm, triangular, non-reflexed, pedicels 6.6–11.8 × 0.4–1.6 mm, glabrescent, tomentose to sericeous; sepals 1.6–5.4 × 1.4–4.6 mm, abaxial side glabrous, sericeous to tomentose, adaxial side glabrous, glabrescent to sericeous, margin flat to slightly revolute at anthesis, triangular to ovate, not accrescent on the drupes; glands 2.1–3.7 × 0.9–1.3 mm; all petals yellow becoming red to orange in age, lateral petals blade 3.9–5.5 × 3.6–5.4 mm, claw 2.1–3.2 × 0.2–0.5 mm, margin flat to slightly erose; posterior petal blade 2.3–3.3 × 2.4–2.9 mm, claw 2.9–3.8 × 0.7–1 mm, margin erose. Stamens free at base, homomorphic, anthers sericeous, connectives 2–2.4 × 0.5–0.9 mm, surpassing the anther sacs (0.4–0.5 mm length); filaments 2–2.3 × 0.4–0.6 mm, sericeous at base. Ovary sericeous, style 2.6–4.1 × 0.2–0.3 mm. Drupes 11.5–20 × 10–17.6 mm, glabrous to glabrescent, smooth, globose to ovoid.

Specimens examined:— BRAZIL. Espírito Santo. Linhares: Área Experimental da Reserva Natural Vale , 20 April 2006, fr., G.S. Siqueira 250 ( CVRD!, HUFSJ!); ibidem, 29 September 2006, bud, fl., D.A. Folli 5361 ( CVRD!, HUFSJ!); Reserva Natural Vale, 08 January 1973, bud, fl., J. Spada 138/73 ( CVRD!) ; ibidem, 05 January 1978, bud, fl., J. Spada 17/78 ( CVRD!) ; ibidem, 24 July 1991, fr., G.L. Farias 461 ( CVRD!, HUFSJ!); ibidem, 17 July 2002, fr., D.A. Folli 4289 ( CVRD!) ; ibidem, 8 April 2005, fr., D.A. Folli 5045 ( CVRD!, HUFSJ!) . Serra: Morro do Mestre Álvaro, 22 March 2010, fr., D.A. Folli 6593 ( CVRD!, HUFSJ!); Reserva Florestal , próximo a estrada X-I talhão 301, 25 January 1973, bud, fl., J. Spada 162 ( CVRD!, RB!) .

Notes: — Byrsonima crassifolia is a widespread species, from Mexico and West Indies to South America, reaching Bolivia and Brazil ( Francener 2020). Occurs in almost all Brazilian states, except the southern states and Rio de Janeiro, Acre, Alagoas, and Sergipe. Byrsonima crassifolia is characterized by yellow petals turning red or orange in age, thyrse 1–2-flowered, the main axis ca. 10 cm long, glabrescent, tomentose to sericeous (hairs stalked), ovary and anther sericeous and drupes ca. 16 mm diam. Byrsonima crassifolia resembles B. alvimii , but it should be noted that the latter has glabrous leaves with few trichomes in the central vein, while B. crassifolia has tomentose leaves when young, deciduous with maturity, and when in flowers the colors the petals are different between these two species.

. Byrsonima crispa A. Jussieu (1840: 335) . Figure 2D

Trees, 5–19 m tall. Stipules 2–4 × 0.8–1.8 mm, triangular. Leaf blades 12.2–17.2 × 5.5–6.4 cm, ovate to elliptic, base attenuate to cuneate, margins flat, apex acute to acuminate, abaxial side glabrous to glabrescent, with few tricomas on midrib, adaxial side glabrous, midrib and principal laterals veins prominulous and number of ca. 11; petiole 24–34 × 1.4–2.4 mm, glabrous, glabrescent to slightly sericeous. Thyrse of 1-flowered cincinnus, ca. 55 flowers, main axis 4.2–10 cm long, tomentose to sericeous; bract 3–4.6 × 0.4–1 mm, lanceolate to triangular, both side tomentose to sericeous, reflexed; peduncle 0–1.4 × 0.4 mm; bracteoles 1.4–1.6 × 0.9–1.4 mm, triangular, reflexed; pedicels 11–33 × 0.5–1.8 mm, tomentose to slightly sericeous; sepals 4–5.6 × 2.8 mm, both side sericeous to glabrescent, margin slightly revolute at anthesis, triangular, accrescent on the drupes; glands 2.2–2.9 × 0.9–1.1 mm; petals yellow becoming red to orange in age, lateral petals blade ca. 5.3 × 4.9 mm, claw ca. 2.9 × 0.5 mm, margin slightly erose; posterior petal blade ca. 3.1 × 3.2 mm, claw ca. 3.1 × 0.9 mm, margin slightly erose. Stamens free, homomorphic, anthers sericeous, connectives ca. 1.9 × 0.9 mm, not surpassing the anther sacs; filaments ca. 2 × 0.3 mm, sericeous at base. Ovary sericeous, style ca. 4 × 0.3 mm. Drupes 6.4–8.4 × 5.3–8 mm, sericeous on the apex, smooth, globose to ovoid.

Specimens examined:— BRAZIL. Espírito Santo. Águia Branca: Santa Luzia, prop. Ciro Ferreira , 18°59’5.4” S, 40°39’44.9” W, 150–270 m, 20 December 2007, bud, fl., V. Demuner 4843 ( MBML!) . Colatina : 30 January 1992, bud, fl., V.D. Souza 306 ( CVRD!, HUFSJ!) . Conceição da Barra: Reserva Biológica de Córrego Grande , km 3 a 4 da estrada interna, 13 July 2011, fr., M. Ribeiro 570 ( VIES!) . Marilândia : Estrada ES 356, 8 February 2012, fr., D.A. Folli 6845 ( CVRD!, HUFSJ!) .

Notes: — Byrsonima crispa is a widespread species, from south of Central America and north of South America on Amazon Forest, with disjunct distribution on Atlantic Forest ( Francener 2020). It is characterized by leaves elliptic, margin undulate, with acuminate apex, midrib and lateral veins prominulous, bracts and bracteoles strongly reflexed and usually persistent to maturity of the fruit, long petiole and sepals accrescent in fruit ( Anderson 1981). Although bracts and bracteoles are usually persistent at maturity, one of the examined exsiccate did not preserve this characteristic (Folli 6845, HUFSJ). B. crispa resembles B. sericea , but the latter has a densely sericeous abaxial side and appressed bracts and bracteoles.

. Byrsonima fanshawei W.R. Anderson (1981: 84) . Figure 1C

Trees, ca. 12 m tall. Stipules ca. 6.2 × 2.5 mm, broadly ovate, not amplexicaul, persistent. Leaf blades ca. 13.7 × 6 cm, elliptic, base cuneate to attenuate, margins flat to slightly revolute, apex acute, cuspidate to acuminate, abaxial side tomentose to slightly sericeous, adaxial side glabrous to glabrescent, shortly-stalked T-shaped hairs, midrib prominulous on side abaxial and impress on side adaxial, principal laterals veins prominulous on side abaxial and impress on side adaxial and number of ca. 11; petiole ca. 16 × 1.5 mm, tomentose, sericeous to glabrescent. Thyrse of 1-flowered cincinnus, ca. 60 flowers, main axis ca. 13.6 cm long, tomentose; bract ca. 2 × 1.4 mm, triangular, abaxial side sericeous, adaxial side glabrescent, non-reflexed; peduncle ca. 2.7 × 1.2 mm; bracteoles ca. 1.4 × 1.1 mm, triangular, non-reflexed; pedicels ca. 11.2 × 1 mm, tomentose; sepals ca. 4.8 × 3.0 mm, sericeous to glabrescent on both sides, margin flat to slightly revolute at anthesis, triangular, not accrescent on the drupes; glands ca. 2.5 × 1 mm. Flowers not seen [according to Francener et al. (2018), all petals yellow becoming red to orange in age, lateral petals blade 5.5–5.2 × 5–5.2 mm, claw 2.3–3 × 0.8–0.9 mm, margin slightly erose; posterior petal blade 3.8–4 × 3.5–4 mm, claw 3–3.5 × 0.9 mm, margin slightly erose. Stamens free, homomorphic, anthers sericeous, connectives 2–2.7 × 0.6–1 mm, surpassing or not the anther sacs (0–0.4 mm length); filaments 2–2.3 × 0.7–1 mm, sericeous at base. Ovary sericeous, style 4–4.8 mm long]. Drupes ca. 9.5 × 8.5 mm, sericeous to glabrescent, smooth, globose.

Specimens examined:— BRAZIL. Espírito Santo. Conceição da Barra: Reserva Biológica do Córrego Grande , 14 May 2017, fr., D.A. Folli 7555 ( CVRD!, HUFSJ!) . Linhares: Reserva Natural Vale , 19 June 2006, fr., D.A. Folli 5303 ( CVRD!, HUFSJ!) .

Notes: — Byrsonima fanshawei occurs in Guyana and Brazil, in the states of Espírito Santo, Bahia, and Amazonas, in the domain of the Atlantic Forest and Amazon Forest ( Francener et al. 2018). It resembles B. stipulacea , but the latter has longer and deciduous stipules (amplexicaul), stellate leaf-hairs, and is larger, while B. fanshawei is shorter, persistent stipules (sometimes deciduous), shortly-stalked T-shaped leaf-hairs, and by small fruits ( Anderson 1981).

. Byrsonima intermedia A. Jussieu (1833: 82) . Figure 2E

Shrubs, ca. 1 m tall. Stipules ca. 3.8 × 2.2 mm, widely triangular. Leaf blades ca. 5.2 × 2.9 cm, elliptic to lanceolate, base attenuate, margins flat, apex rounded to acute, abaxial side glabrous to glabrescent, with a few sericeous hairs in midrib, adaxial side glabrous, midrib and principal laterals veins prominulous at both sides and number of ca. 8; petiole ca. 3.3 × 3.1 mm, glabrescent. Thyrse of 1-flowered cincinnus, ca. 25 flowers, main axis ca. 5.8 cm long, tomentose; bract ca. 1 × 0.7 mm, ovate, abaxial side sericeous, adaxial side slightly sericeous, non-reflexed; peduncle absent; bracteoles ca. 1 × 0.6 mm, ovate, non-reflexed; pedicels ca. 3.5 × 1 mm, sericeous; sepals ca. 2.2 × 1.7 mm, both the sides sericeous, margin slightly revolute at anthesis, widely ovate, not accrescent on the drupes; glands ca. 2.4 × 1.4 mm; all petals yellow becoming red to orange in age, lateral petals blade ca. 4.7 × 5.3 mm, claw ca. 2.2 × 0.6 mm, margin entire to slightly erose; posterior petal blade ca. 2 × 3.7 mm, claw ca. 2.4 × 0.8 mm, margin erose. Stamens free, homomorphic, anthers sericeous, with few trichomes, connectives ca. 1.8 × 0.8 mm, equaling pollen sacs; filaments ca. 1.7 × 0.7 mm, sericeous at base. Ovary glabrous, style ca. 2.6 × 0.4 mm. Drupes ca. 4 × 5 mm, glabrous, smooth, ovoid.

Specimen examined:— BRAZIL. Espírito Santo. Santa Teresa: Pedra da Paulista , 17 February 2000, bud, fl., fr., V. Demuner 741 ( MBML!) .

Notes: — Byrsonima intermedia occurs in Paraguay and Brazil, in the states of Pará, Tocantins, Bahia, Distrito Federal, Goiás, Mato Grosso do Sul, Mato Grosso, Minas Gerais, São Paulo and Paraná, in the domain of the Atlantic Forest, Amazonia, Cerrado and Pantanal ( Francener 2020). It is characterized by size–shrubs, ca. 1 m tall, leaves elliptic to lanceolate, base attenuate, apex rounded to acute, abaxial side slightly sericeous in the midrib, adaxial side glabrous, bracts not reflexed, yellow petals becoming orange or red in age. It resembles B. crassifolia but it has the abaxial side of the leaves tomentose when young, connective surpassing the anther sacs (0.4–0.5 mm length) and ovary sericeous. B. intermedia is recorded for the first time for the Espírito Santo state.

. Byrsonima japurensis A. Jussieu (1840: 335) . Figure 1D

Trees, 3–20 m tall. Stipules ca. 2.1 × 0.6 mm, triangular. Leaf blades ca. 7.9 × 5.15 cm, elliptic to obovate, base attenuate, margins flat to slightly revolute, apex acuminate to rounded, both side glabrous, midrib prominulous on side abaxial and impress on side adaxial, principal laterals veins prominulous on side abaxial and impress on side adaxial, and number of ca. 8; petiole ca. 8.6 × 1.75 mm, densely tomentose. Thyrse 1-flowered cincinnus, ca. 15 flowers, main axis ca. 7.5 cm long, densely tomentose; bracts not seen [according to Rolim (2004), 0.5–1.5 × 0.5–1 mm, lanceolate, ovate, triangular to rarely suborbicular, abaxial side slightly sericeous to slightly tomentose, adaxial side glabrous, non-reflexed]; peduncle ca. 0.8 × 1.5 mm, densely tomentose; bracteoles not seen [according to Rolim (2004), 0.5–1 × 0.5–0.8 mm, lanceolate, triangular, ovate, oblong to rarely suborbicular, non-reflexed]; pedicels ca. 13.8 × 1.2 mm, densely tomentose; sepals ca. 2.8 × 2.9 mm, abaxial side glabrous, sericeous to tomentose, adaxial side glabrous to slightly sericeous, margin flat to slightly revolute at anthesis, triangular, not accrescent on the drupes; glands ca. 1.8 × 1 mm. Flowers not seen [according to Rolim (2004), all petals white becoming pink in age, lateral petals blade 3–4.45 × 4–6 mm, claw ca. 3 × 0.6 mm, margin erose; posterior petal blade ca. 3–4 × 3–3.5 mm, claw ca. 3 × 0.9 mm, margin erose. Stamens free, homomorphic, anthers sericeous, connectives 0.7–3.2 × 0.5 mm, surpassing the anther sacs (0.2– 1.7 mm length); filaments 1.8–2 × 0.3–0.7 mm, sericeous at base. Ovary glabrous, style 3.4–4.6 mm long]. Drupes ca. 11.3 × 12 mm, glabrous, verrucous, globose.

Specimens examined:— BRAZIL. Espírito Santo. Santa Teresa: Estação Biológica Santa Lúcia , trilha do palmiteiro, 30 September 1999, fr., V. Demuner 59 ( MBML!) ; Reserva Biológica de Nova Lombardia , December 1989, st. L.P.S. Pinto s.n. ( BHCB!, MBML!) .

Notes: — Byrsonima Japurensis occurs in Brazil, Colombia, Ecuador, Peru, and Venezuela. The Brazilian territory has a wide distribution of these species: in the north, northeast, and southeast regions, in the domain of the Atlantic Forest and Amazonia ( Francener 2020). Leaves obovate to elliptical, petiole ca. 8.5 mm long, densely tomentose, the main axis ca. 7.5 mm long, densely tomentose, bracts non-reflexed and deciduous at maturity, characterize B. Japurensis . It resembles B. perseifolia but it has a larger petiole (ca. 12 mm long), glabrescent to slightly sericeous, the main axis ca. 6 mm long, slightly sericeous, and smaller drupes (ca. 8.5 mm).

FIGURE: A. Byrsonima ligustrifolia (photo by R.F. de Almeida), B. Byrsonima stipulacea , detail of the amplexicaule stipules and starshaped trichome (photo by M. Pastore), C. Byrsonima vernicosa (Vervloet 1940, MBML), D. Byrsonima sericea (Fontana 664, MBML)