Byrsonima myricifolia Grisebach (1875: 125)

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 : 11-16

11. Byrsonima myricifolia Grisebach (1875: 125) . Figure 4A

Trees, 5–14 m tall. Stipules 1.4–2.3 × 0.5–0.9 mm, triangular to lanceolate. Leaf blades 2.26–5.34 × 0.8–2.58 cm, obovate, oblong to elliptic, base attenuate, margins revolute, apex acute, acuminate to rounded, abaxial side glabrous to glabrescent, adaxial side glabrous, midrib prominulous on both sides, lateral veins up to 5, impress on both sides; petioles 5–11 × 11–16 mm, glabrescent to slightly tomentose. Thyrse, bearing ca. 20 flowers, of 1-flowered per cincinnus, main axis 2.9–3.2 cm long, glabrous; bracts ca. 1.8 × 0.7 mm, lanceolate, glabrescent on both sides, non-reflexed; peduncle 0–0.6 mm long; bracteoles ca. 1 × 0.3 mm, lanceolate, non-reflexed; pedicels 7.8–11 × 0.8–1.3 mm, sericeous; sepals 2–3.5 × 1.5–2.5 mm, abaxially glabrous or slightly sericeous to slightly tomentose, adaxially glabrescent, margin flat at anthesis, triangular to widely ovate, not accrescent on the drupes; glands 1.1–2.3 × 0.5–1 mm; all petals white to pink becoming red with age, lateral petals with blades 3.5–5 × 4.7–6 mm, claw 2.6–2.9 × 0.6–0.8 mm, margin slightly entire to erose; posterior petal blade 2.8–4.6 × 2.8–5 mm, claw ca. 3 × 0.5–0.8 mm, margin slightly entire to erose. Stamens free at base, homomorphic, anthers glabrous to glabrescent, connectives ca. 2 × 0.8 mm, surpassing the anther sacs (0–1 mm length); filaments ca. 2.3 × 0.4 mm, sericeous at base. Ovary glabrous to sericeous, styles ca. 2.3 × 0.4 mm. Drupes not seen [according to Grisebach (1875) they are globose to ovoid, up to 4 mm].

Specimens examined:— BRAZIL. Espírito Santo. Santa Teresa: Cabeceira do Rio Bonito , terreno da Aeronáutica, radar, 1030 m, 12 June 2001, bud, L. Kollmann 3901 ( MBML!) ; Estação Biológica Santa Lúcia , 16 October 2004, fl., F.A.G. Guilherme 391 ( MBML!) ; Santo Antônio, terreno do Boza , 750 m, 9 February 1999, fl., L. Kollmann 1867 ( MBML!) .

Notes: — Byrsonima myricifolia is endemic to Brazil. It is distributed from the states of Bahia to Paraná within the Atlantic Forest domain ( Francener 2020). Its main features are the short main axis (ca. 3 cm), with few flowers and small glabrous leaves. It resembles B. ligustrifolia , but it has inflorescences with the main axis longer (up to 5 cm) and tomentose. Rolim (2004) considered B. myricifolia as a variety of B. ligustrifolia . In her opinion, although these plants have differences, the proximity between them is very great and possibly, they may be in a process of speciation or come from hybridizations ( Rolim 2004). It is necessary to highlight the inconsistencies in the description of the material examined Kollmann 1867 ( MBML) with the original ( Grisebach 1875), such as anthers and glabrous ovaries and connective exceeding the anther sacs.

. Byrsonima niedenzuiana Skottsberg (1901: 36) . Figure 4B

Trees, ca. 10 m tall. Stipules ca. 2.9 × 1.2 mm, triangular. Leaf blades ca. 6.7 × 3.7 cm, obovate, base attenuate, margins flat to slightly revolute, apex rounded to retuse, both sides glabrous to glabrescent, midrib prominulous on abaxial side and impress on adaxial side, principal laterals veins ca. 8, prominulous to impress on abaxial side and impress on adaxial side; petioles ca. 7 × 1.6 mm, tomentose to glabrescent. Thyrse of 1-flowered cincinnus, with ca. 18 flowers, main axis ca. 6.6 cm long, densely tomentose ferruginous; bracts ca. 1.5 × 0.7 mm, lanceolate, abaxial side densely tomentose, adaxial side glabrescent, non-reflexed; peduncle absent to 1.5 × 0.7 mm; bracteoles ca. 1.2 × 0.5 mm, triangular, non-reflexed; pedicels ca. 7.8 × 0.8 mm, densely tomentose; sepals ca. 3.7 × 2.8 mm, abaxial side tomentose to slightly sericeous, adaxial side tomentose to glabrescent, margin flat at anthesis, triangular, not accrescent on the drupes; glands ca. 2.4 × 1.4 mm; all petals pink, lateral petals blade ca. 5.4. × 4.6 mm, claw ca. 2.3 × 0.3 mm, margin slightly erose; posterior petal blade ca. 3.3 × 3.2 mm, claw ca. 2.5 × 0.8 mm, margin slightly erose. Stamens free, homomorphic, anthers glabrous, connectives ca. 1.9 × 0.6 mm, surpassing the anther sacs (up to 0.7 mm length); filaments ca. 2 × 0.4 mm, sericeous at base. Ovary glabrous, style ca. 2.3 × 0.4 mm. Drupes ca. 9 × 9.8 mm, sericeous on below side, smooth, globose.

FIGURE: A. Byrsonima myricifolia (Kollmann 3901, MBML), B. Byrsonima niedenzuiana (Martinelli 10947, RB), C. Byrsonima nitidifolia (Demuner 1047, MBML) D. Byrsonima perseifolia (Silva 37, CVRD)

Specimens examined:— BRAZIL. Espírito Santo. Santa Teresa: Nova Lombardia, Reserva Biológica Augusto Ruschi , estrada Goipabo-Açu , parte final, 8 April 2003, fr., R.R. Vervloet 2155 ( MBML!) ; Reserva Biológica de Nova Lombardia, Picada da Cachoeira , 800–1000 m, 10 May 1985, fl., G. Martinelli 10947 ( MBML!, RB!) .

Notes: — Byrsonima niedenzuiana is endemic of Brazil, occurs in the states of Espírito Santo, São Paulo, Paraná, Rio Grande do Sul and Santa Catarina, in the domain of the Atlantic Forest ( Francener 2020). Its main features are obovate leaves with apex rounded to retuse, all petals pink, smooth fruits, ca. 9 mm, connectives surpassing the anther sacs (up to 0.7 mm length), and sepals of the same size on drupes. It resembles B. bahiana , but it has the verrucous fruits, the connectives slightly surpassing the anther sacs (up to 0.1 mm length), and all petals white, often tinged with pink, with reddish claw and enlarger fleshy sepals on drupes .

. Byrsonima nitidifolia A. Jussieu (1843: 296) . Figure 4C

Small trees, 2–9 m tall. Stipules 2.5–5 × 0.9–2.6 mm, triangular to lanceolate. Leaf blades 3.8–7.3 × 2–4.6 cm, oblong, base attenuate, cuneate to sessile, margins slightly revolute and undulate, apex acute, acuminate, rounded, apiculate to retuse, both side glabrous, midrib prominulous on both side, principal laterals veins prominulous on abaxial side and impress on adaxial side and number of ca. 8; petiole 4–7 × 1–1.9 mm, sericeous, slightly sericeous to tomentose. Thyrse of 1-flowered cincinnus, ca. 21 flowers, main axis 3.5–8.8 cm long, tomentose; bract ca. 2.5 × 1.7 mm, ovate, both side tomentose, non-reflexed; peduncle 0–1.2 × 1 mm; bracteoles ca. 1.8 × 1 mm, ovate, non-reflexed; pedicels ca. 11 × 1.2 mm, densely tomentose; sepals 2.4–7.6 × 1.6–3 mm, abaxial side densely tomentose, adaxial side sericeous at base, glabrescent to slightly sericeous, margin flat to slightly revolute at anthesis, triangular, not accrescent on the drupes; glands 1.3–3.5 × 0.4–2 mm. Flowers not seen [according to Jussieu (1843), all petals white becoming pink in age. Anthers sericeous, connectives surpassing the anther sacs. Ovary glabrous]. Drupes 6.6–10 × 6–8 mm, glabrous, smooth, globose.

Specimens examined:— BRAZIL. Espírito Santo. Águia Branca: Pedra da Bandeira, Santa Luzia , prop: Ciro Ferreira , 18°58’40” S, 40°39’56” W, 170–600 m, 26 July 2006, fr., L.F.S. Magnago 1103 ( MBML!) ; ibidem, 18°58’28” S, 40°39’43” W, 250–450 m, 18 May 2007, fr., V. Demuner 3976 ( CEPEC, MBML, SP) ; São Pedro , pedra do CEIER, 19°01’22.2” S, 40°38’52.8” W, 200–550 m, 26 April 2006 fr., V. Demuner 2242 ( MBML!, CEPEC) . Nova Venécia: Área de Proteção Ambiental da Pedra do Elefante, Serra de Baixo , beira da rodovia Nova Venécia-São Gabriel da Palha , prop: Sr. Valdemar , 18°48’16” S, 40°28’11” W, 314 m, 9 May 2008, fr., A.M.A. Amorim 7411 ( MBML!) ; Área de Proteção Ambiental da Pedra do Elefante, Serra de Baixo, morro lado direito na estrada para a Pedra do Elefante , inselberg, 18°46’12” S, 40°26’51” W, 300–600 m, 14 January 2009, bud, A.P. Fontana 5769 ( MBM, MBML!) . Santa Teresa: Estação Biológica de Santa Lúcia , abaixo da cachoeira, 11 May 2000, fr., V. Demuner 1047 ( MBML!) ; Estação Biológica de Nova Lombardia , 850 m, 19 February 2002, fl., L. Kollmann 5562 ( CEPEC, MBML, SP) ;

Additional specimen examined:— BRAZIL. Bahia, Maracás, 13 a 22 Km sul de Maracás, pela Estrada velha de Jequié. 27 April 1978. bud, fl., S.A. Mori s.n. (NY 475421) .

Notes: — Byrsonima nitidifolia is endemic of Brazil, occurs in the state of Pernambuco, Bahia, Ceará and Espírito Santo in the domain of the Atlantic Forest, Cerrado and Caatinga ( Francener 2020). Byrsonima nitidifolia is characterized by leaf blades oblong, elliptic to lanceolate, shiny abaxially, bracts ca. 2.5 mm long, non-reflexed, bracteoles ca. 1.8 mm long, drupes ca. 8 mm, smooth. It resembles B. perseifolia but this one has smaller bracts and bracteoles (ca. 1.5 and 1 mm respectively) and bigger petioles (ca. 12 mm).

. Byrsonima perseifolia Grisebach (1839: 257) . Figura 4D

Trees, 12–20 m tall. Stipules 2.2–4.2 × 0.8–1.7 mm, triangular to lanceolate. Leaf blades 6.27–11.2 × 2.8–4.5 cm, elliptic, base attenuate to cuneate, margins flat to slightly revolute and undulate, apex acute, acuminate falcate to rounded, both side glabrous, midrib prominulous on abaxial side and impress on adaxial side, principal lateral veins ca. 12, prominulous abaxially and impress adaxially; petioles 6–18.5 × 0.9–1.5mm, glabrescent to slightly sericeous. Thyrse of 1-flowered cincinnus, with ca. 25 flowers, main axis 5.7–9.37 cm long, slightly sericeous; bracts 1.2–2 × 0.4–0.8 mm, ovate to triangular, on both sides tomentose, sericeous to glabrescent, non-reflexed; peduncles 0–1.8 × 0–0.6 mm; bracteoles 0.8–2 × 0.5–0.8 mm, ovate to triangular, non-reflexed; pedicels 2.6–10.5 × 0.5–1 mm, tomentose to sericeous; sepals 0.8–3 × 0.5–2.5 mm, abaxial side tomentose to sericeous, adaxial side glabrous, glabrescent to slightly sericeous, margin revolute on apex at anthesis, triangular to widely ovate, not accrescent on the drupes; glands 0.9–2.5 × 0.5–1 mm; all petals white to pink in age, lateral petals with blades 3.1–5.1 × 2.3–5.4 mm, claws 1.5–2.3 × 0.4–0.6 mm, margin slightly erose; posterior petal blades 2.5–3.1 × 2–3.5 mm, claws 2.1–3 × 0.5–0.8 mm, margin erose. Stamens free, homomorphic, anthers glabrous to glabrescent, connectives 1.1–1.7 × 0.5–0.6 mm, surpassing the anther sacs (0–0.3 mm length); filaments 1.1–2.1 × 0.3–0.4 mm, sericeous at base. Ovary glabrous, style 2.4–3.4 × 0.2–0.4 mm. Drupes 8–9 mm diam., glabrous, verrucous, globose.

Specimens examined:— BRAZIL. Espírito Santo. Barra de São Francisco: Cabeceira Córrego do Engenho , terreno de Alfredo Bassi, 235 m, 12 December 2000, bud, L. Kollmann 3476 ( MBML!) . Domingos Martins: BR 262, próximo a Vitor Hugo, Rio Jucu , braço sul, 17 January 1995, fl., G. Hatschbach 61600 ( MBM, MBML!) . Linhares: Reserva Natural Vale , 20 December 1978, bud, fl., I.A. Silva 37 ( CVRD!, HUFSJ!) . Santa Teresa: Reserva Biológica de Nova Linhares , December 1989, bud, L.P.S. Pinto s.n. ( BHCB! 18657, MBML! 5835) . Limoeiro, Castelo , 7 July 1996, fr., G. Hatschbach 65262 ( MBM) .

Notes: — Byrsonima perseifolia is endemic of Brazil, occurs in the states of Minas Gerais, Rio de Janeiro, Espírito Santo and São Paulo, in the domain of the Atlantic Forest and Cerrado ( Francener 2020). It is main features are small bracts and bracteoles (ca. 1.5 × 0.5 mm), erect, just like the flower buds, very small or absent peduncle, large petiole, white petals, elliptical leaves, often with undulate margins. It resembles B. alvimii and B. cacaophila , but B. alvimii has larger and reflexed bracts and B. cacaophila has a peduncle ca. 3 mm long.

. Byrsonima sericea De Candolle (1824: 580) . Figure 3D

Shrubs to small trees, 1.7–4 m tall. Stipules 1.1–4 × 1–2 mm, triangular, lanceolate to ovate. Leaf blades 2.37–9.23 × 1–4.17 cm, obovate, elliptic to lanceolate, base attenuate to cuneate, margins flat to slightly revolute, apex acute, acuminate, mucronulate, apiculate to cuspidate, abaxial side densely sericeous to sericeous, adaxial side glabrous, glabrescent to slightly sericeous on midrib, 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 9–21; petiole 3–8.7 × 1–1.8 mm, glabrescent, sericeous to densely tomentose. Thyrse of 1–2-flowered cincinnus, ca. 20 flowers, main axis 3–8.53 cm long, sericeous; bract 1.2–2 × 0.4–1.8 mm, lanceolate to triangular, abaxial side glabrescent, sericeous to tomentose, adaxial side glabrous, glabrescent, sericeous to tomentose, non-reflexed; peduncle 0–1 × 0–0.8 mm; bracteoles 0.7–1.4 × 0.4–1 mm, triangular, non-reflexed; pedicels 5.2–10 × 0.4–1 mm, sericeous; sepals 2–2.9 × 1.3–2.2 mm, abaxial side sericeous to slightly sericeous, adaxial side glabrous, glabrescent to sericeous, margin flat to slightly revolute at anthesis, triangular, ovate to widely ovate, not accrescent on the drupes; glands 1.5–2.3 × 0.6–1 mm; all petals yellow becoming red to orange in age, lateral petals blade 2.7–5 × 5–5.2 mm, claw 2–2.6 × 0.3–0.6 mm, margin erose; posterior petal blade 2.9–3.4 × 3.1–4.3 mm, claw 2.7–3.4 × 0.8–1 mm, margin erose. Stamens free to connate at base, homomorphic, anthers glabrous to slightly sericeous, connectives 1.7–2.4 × 0.4–0.7 mm, surpassing the anther sacs (0–0.2 mm length); filaments 1.5–2.1 × 0.3–0.6 mm, sericeous at base. Ovary glabrous to sericeous, style 2.7–3.3 × 0.2–0.3 mm. Drupes 4–6.8 × 3–6.6 mm, glabrous, glabrescent to slightly sericeous, verrucous, globose.

Specimens examined:— BRAZIL. Espírito Santo. Anchieta: Samarco , Área A, 26 July 2000, fr., J. Dalmaschio s.n. ( MBML! 13439) . Colatina: Alto Moacir, Pedra do Cruzeiro , 19°20’53” S, 40°33’3” W, 150–850 m, 17 April 2006, fl., fr., L.F.S. Magnago 796 ( MBML!) . Governador Lindenberg: Pedra de Santa Luzia , 19°16’54” S, 40°27’43” W, 350–650 m, 26 April 2007, fr., V. Demuner 3865 ( MBML!) . Linhares: Pontal de Ipiranga, Degredo , 1 April 2007, fr., L. Kollmann 9594 ( MBML!) ; Reserva Biológica de Comboio —IBAMA, Regência , 08 July 1988, bud, fr., G.L. Farias 197 ( CVRD!) ; Reserva Biológica de Comboio —IBAMA, Regência, próx. ao mar, 50 m do mar, 10 m da estrada, lado esquerdo, Souza , 12 June 1991, fl., V.D. Souza 106 ( CVRD!) ; Reserva Natural Vale , 21 December 1981, bud, fl., I.A. Silva 282 ( CVRD!, HUFSJ!) ; ibidem, 12 June 1991, bud, fl., L.P. Queiroz 2481 ( CVRD!, HUFSJ!) ; Sooretama , sede, perto da lagoa, 15 February 1972, bud, s.col. s.n. ( MBML! 2910) ; Marilândia: Liberdade ( Água-Viva , Pedra do Cruzeiro ), 19°20’53.7” S, 40°33’03.6” W, 150–650 m, 18 January 2006, V. Demuner 1595 ( MBML!) . Praia das Neves , divisa com o estado do Rio de Janeiro, 9 February 1996, fl., fr., C.N. Fraga 292 ( MBML!) ; Praia das Neves , 2 November 1987, bud, fl., J.M.L. Gomes 164 ( VIES!) ; Presidente Kennedy: Propriedade da FERROUS, próximo à Praia das Neves , 21°12’52” S, 40°59’23” W, 14 January 2010, bud, Maielo-Silva 116 ( RB!) . Santa Leopoldina: Luxemburgo, Pedra Preta , 15 March 2005, fl., fr., L. Kollmann 7403 ( MBML!) . São Mateus: Reserva Biológica do Projeto TAMAR, Guriri , 11 April 1991, fr., V.D. Souza 62 ( CVRD!, HUFSJ!) . São Roque do Canaã: Misterioso , fim da estrada, 24 December 2003, fl., A.P. Fontana 664 ( MBML!) . Vila Velha: Barra do Jucu , 14 April 1986, fl., L.C. Fabris 37 ( VIES!) ; Morada do Sol , 20°27’43.7” S, 40°20’35.2” W, 3 m, 14 January 2006, fl., F.A.R. Matos 54 ( MBML!) .

Notes: — Byrsonima sericea occurs in Bolivia, Brazil, French Guiana, Guyana, and Peru. In Brazil is widely distributed, especially on the coast, from São Paulo to Pará, in the domain of the Amazonia, Caatinga, Cerrado, and Atlantic Forest ( Francener 2020). Shrubs to small trees characterize Byrsonima sericea , but we found an unusual specimen with 15 m high. Its main feature is the abaxial side of the leaves densely sericeous with adpressed hairs. It resembles B. chrysophylla , as discussed in item 2.2 in excluded species.

. Byrsonima stipulacea A. Jussieu (1840: 332) . Figure 3B

Trees, 5–25 m tall. Stipules 5–25 × 2–7.8 mm, triangular, lanceolate to ovate, amplexicaul, deciduous. Leaf blades 5– 19 × 1.7–12 cm, obovate to elliptic, base attenuate to cuneate, margins flat to slightly revolute, apex acute, acuminate to rounded, abaxial side densely tomentose, tomentose to velutine (stellate and stalked hairs), adaxial side glabrous, glabrescent, slightly tomentose, slightly velutine to tomentose, stellate-stalked 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. 10; petiole 8–23 × 1.8–3 mm, velutine to densely tomentose. Thyrse of 1–2-flowered cincinnus, 15–75 flowers, main axis 7–17 cm long, velutine; bract 2.2–10.4 × 1–2.2 mm, lanceolate to triangular, abaxial side velutine, tomentose to densely tomentose, adaxial side glabrous, glabrescent to slightly velutine, non-reflexed; peduncle 0–2.2 × 0–1.5 mm; bracteoles 2.2–5 × 1.4–2.9 mm, triangular, ovate to lanceolate, non-reflexed; pedicels 2–16.9 × 1–1.5 mm, densely tomentose to velutine; sepals 2.9–8.5 × 1.2–5.3 mm, abaxial side densely tomentose, tomentose to densely velutine, adaxial side tomentose to slightly tomentose, margin revolute at anthesis, triangular, ovate to lanceolate, not accrescent on the drupes; glands 1.7–3.2 × 0.7–1.7 mm; petals yellow turning red to orange in age, lateral petals blade 4.2–5.3 × 5.1–7.2 mm, claw 3–3.6 × 0.4–0.7 mm, margin slightly erose to erose; posterior petal blade 2.4–3.4 × 2.3–3.8 mm, claw 3.3–4.2 × 0.5–0.9 mm, margin erose. Stamens connate at base, homomorphic, anthers sericeous, connectives 1.8–3.5 × 0.3–1.2 mm, surpassing the anther sacs (0.4–1.1 mm length); filaments 1.5–4 × 0.5–0.7 mm, sericeous at base. Ovary sericeous, style 1.8–3.7 × 0.2–0.4 mm. Drupes 9–22.7 × 7.2–21 mm, glabrescent, tomentose to densely tomentose, smooth, globose to ovoid.

Specimens examined:— BRAZIL. Espírito Santo. Aracruz: Estrada para Santa Rosa , 19°53.32’ S, 40°16.58’ W, 4 April 2007, bud, H.C. Lima 6599 ( RB!) ; Reserva do Putiri , 19°82’03” S, 40°27’33” W, 10 May 1994, fr., J.N. Neves 1 ( VIES!) . Fundão: Goiapaba-Açu , 700 m, 15 July 1998, fr., L. Kollmann 255 ( MBM, MBML!) . Linhares : OCEMADE, 1 February 1972, fl. D. Sucre 8353 ( RB!) ; Fazenda Trajano , 100 m da sede da fazenda em linha reta, 6 February 1991, bud, fl., V.D. Souza 15 ( CVRD!, HUFSJ!) ; Flona de Goytacazes , 9 April 2011, fr., J.M.L. Gomes 3864 ( VIES!) ; Flona de Goytacazes , perto da sede, 23 April 2011, bud, J.M.L. Gomes 3714 ( VIES!) . Reserva Biológica de Sooretama , 12 May 1985, fr., G. Martinelli 10962 ( MBML!, RB!) ; Reserva da Companhia Vale do Rio Doce , 8 April 2006, bud, G.O. Romão 1282 ( CVRD!, RB!) ; Reserva da Companhia Vale do Rio Doce, Estrada MME, 24 July 2006, fr., J. Paula-Souza 6358 ( RB!) ; Reserva Florestal de Linhares, próximo Estrada 154, 7 March 1972, fr., A.M. Lino 17 ( CVRD!, RB!) ; Reserva Natural da Vale, Sooretama , 19°08’32.1” S, 40°03’58.1” W, 38 m, 02 February 2013, bud, M.G. Caxambu 4553 ( RB!) ; Reserva Natural Vale , 9 January 1982, bud, fl., I.A. Silva 309 ( CVRD!, HUFSJ!) ; ibidem, 16 August 1991, fr., V.D. Souza 129 ( CVRD!) ibidem, 19 September 1991, fr., D.A. Folli 1416 ( CVRD!, HUFSJ!) ; ibidem, 17 April 2011, fr., D.F. Lima 207 ( RB!) ; Reserva Natural Vale do Rio Doce (particular), próximo a estrada X-I, Talhão 606, 8 March 1972, bud, fr., J. Spada 20 ( RB!) ; Reserva Natural Vale, Estrada Gávea , 19°11’S 39°54’W, 9 February 1999, bud, fl., R. Mello-Silva 1546 ( BHCB!, CVRD!, RB!, HUFSJ!) . Santa Leopoldina: Fazenda Caioaba , 20°3’33” S, 40°28’14” W, 150–650 m, 17 July 2007, fr., R.R. Vervloet 2909 ( MBML!) ; Fazenda Caioaba, prop: Virloni, trilha do Córrego Caioaba , 20°3’33” S, 40°28’14” W, 150–650 m, 8 August 2006, fr., L.F.S. Magnago 1241 ( MBML!) ; Pedra Branca, mata na Serra Santa Lúcia, prop: Cristiano Bremencampi , 20°12’2” S, 40°29’26” W, 250– 380 m, 18 April 2007, fl., V. Demuner 3622 ( MBML!) ; Serra do Ramalhete, prop: Cláudio Virloni, trilha do Córrego Caioaba , 20°03’30.8” S, 40°28’14.3” W, 200–500 m, 15 February 2006, bud, fl., V. Demuner 1833 ( MBML!) ; ibidem, 4 April 2008, fl., fr., L. Kollmann 10794 ( MBML!) ; Suíça , 20°5’417” S, 40°35’351” W, 460 m, 12 April 2008, fl., M. Simonelli 1481 ( MBML!) . Santa Teresa: Bairro do Eco, mata do Banestes , 650 m, 22 October 1998, fr., L. Kollmann 769 ( MBML!) ; Penha, mata do Tabajara , 22 March 2005, fl., L. Kollmann 7528 ( MBML!) ; Rio Saltinho, estrada para Goiapaba-Açu , 350 m, 29 August 2001, fr., L. Kollmann 4404 ( MBML!) ; Santo Antônio, terreno do Boza , 850 m, 27 April 1999, fl., L. Kollmann 2536 ( MBML!) ; Valsugana Velha, Estação Biológica Santa Lúcia , 19°58’ S, 40°32’ W, 600–900 m, 15 May 1991, fl., W. Pizziolo 352 ( MBML!) . Sooterama: Reserva Florestal da Companhia Vale do Rio Doce , 23 February 2000, fl., A.M Amorim 3355 ( CVRD!, MBML!) .

Notes: — Byrsonima stipulacea occurs in Bolivia, Brazil, French Guiana, Guyana, Peru, Suriname, and Venezuela. Its distribution in Brazilian territory goes from the northern, southeastern, and some northeastern states, in the domain of the Atlantic Forest and Amazonia ( Francener 2020). B. stipulacea has stellate-stalked hairs admixture with simple hairs, also large stipules (ca. 4 mm long), sometimes deciduous. It resembles B. duckeana as discussed in item 2.3 in excluded species.

. Byrsonima vernicosa Niedenzu (1901: 33) . Figure 3C

Trees, 8–15 m tall. Stipules 2.5–5.5 × 1.4–3.4 mm, triangular, lanceolate to ovate. Leaf blades 2–9.1 × 1–4.9 cm, obovate to elliptic, base attenuate to cuneate, margins flat, slightly revolute to revolute, apex acute, acuminate, retuse, apiculate to rounded, abaxial side densely tomentose to tomentose, adaxial side glabrescent to tomentose on midrib, midrib prominulous on side abaxial and prominulous to impress on side adaxial, principal laterals veins prominulous to impress on side abaxial and impress on side adaxial and number of ca. 10; petiole 0.95–7.8 × 1–2.6 mm, glabrescent, tomentose to densely tomentose. Thyrse of 1-flowered cincinnus, ca. 25 flowers, main axis 3.87–11.14 cm long, densely tomentose; bract 1.4–4 × 0.6–1.4 mm, lanceolate, triangular to ovate, abaxial side tomentose, adaxial side glabrescent, non-reflexed; peduncle 1.5–5.3 × 1–1.7 mm; bracteoles 1.3–2.3 × 0.8–2 mm, triangular, ovate to lanceolate, nonreflexed; pedicels 1.6–11 × 1–1.7 mm, densely tomentose; sepals 3–5.1 × 2–4.2 mm, abaxial side densely tomentose to tomentose, adaxial side glabrescent to tomentose, margin flat after anthesis, widely triangular, ovate to widely ovate, not accrescent on the drupes; glands 1.1–3.4 × 0.8–2.3 mm; all petals white becoming pink in age, blade 5.2–6.5 × 5–6.3 mm, claw 2.4 × 0.6 mm, margin erose; posterior petal blade 3.4–3.9 × 2.7–4.3 mm, claw 2.7–3.5 × 0.5–0.6 mm, margin erose. Stamens connate at base, homomorphic, anthers glabrous, connectives 1.5–2.4 × 0.4–0.8 mm, surpassing or not the anther sacs (ca. 0.1 mm length); filaments 2.5–3.1 × 0.3–0.5 mm, glabrescent to sericeous at base. Ovary glabrous, style 3.2–3.5 × 0.3–0.5 mm. Drupes 9.6–12 × 10–13.8 mm, glabrous to slightly sericeous at apex, verrucous, discoid.

Specimens examined: BRAZIL. Espírito Santo. Castelo: Parque Estadual do Forno Grande , trilha da balança, 20°31’37” S, 41°06’6” W, 1250–1600 m, 18 July 2008, fr., R. Goldenberg 1177 ( MBM, MBML!) . Santa Maria de Jetibá: Rio das Pedras , terreno de Paulo Kuzanki (área 2), 700 m, 20 January 2003, bud, fr., L. Kollmann 5921 ( MBML!, MICH) ; Rio Nove , ter. L. Kollmann, 24 February 2000, bud, fl., fr., V. Demuner 778 ( ESA, MBML!) . Santa Teresa: Estação Biológica Santa Lúcia , 650–800 m, 8 December 1992, st., L.D. Thomaz 985 ( MBML!, VIES!) ; ibidem, 7 March 1995, fl., L.D. Thomaz 987 ( MBML!, VIES!) ; Estação Biológica Santa Lúcia , trilha do palmiteiro, 27 January 2000, bud, V. Demuner 637 ( MBML!) ; Nova Lombardina, Reserva Biológica Augusto Ruschi , 800 m, 4 October 2001, fr., L. Kollmann 4801 ( MBML!) ; Nova Lombardina, Reserva Biológica Augusto Ruschi , cabeceira do rio Saltinho , 800 m, 13 March 2002, fl., L. Kollmann 5646 ( MBML!) ; Nova Lombardina, Reserva Biológica Augusto Ruschi , divisa, 800 m, 21 January 2003, bud, fl., fr., R.R. Vervloet 1688 ( MBML!) ; Nova Lombardina, Reserva Biológica Augusto Ruschi , linha de divisa, lado direito, saída para João Neiva , 6 March 2003, fl., R.R. Vervloet 1940 ( MBML!) ; Reserva Biológica Augusto Ruschi , primeira divisa à esquerda, 750 m, 8 January 2002, bud, L. Kollmann 5212 ( MBML!) ; Santo Antônio, terreno do Boza , 850 m, 16 March 1999, fl., L. Kollmann 2164 ( MBML!) . São Roque do Canaã: Alto Misterioso , 973 m, 25 November 2007, bud, M. Simonelli 1363 ( MBML!) .

Notes: — Byrsonima vernicosa is endemic to Brazil, occurring in the state of Espírito Santo and Rio de Janeiro, in the domain of the Atlantic Forest ( Francener 2020). It is characterized by the main axis, pedicels and abaxial side of leaves densely tomentose to tomentose and drupes ca. 12 mm diam, verrucous. It resembles B. variabilis , as discussed later under excluded species .