Ehretia P.Browne.

[differences from Luebert et al. (2016) in bold] Trees, shrubs, lianas, perennial herbs, rarely with thorns (Rochefortia Sw.); indumentum variable, hirsute to glabrescent. Leaves alternate, entire, petiolate; lamina variable in shape, strongly dissected in the halophytic Cortesia Cav. Inflorescences terminal or axillary thyrses, sometimes congested, or few-flowered corymbs (Keraunea). Flowers pentamerous, cosexual or unisexual and dioecious in Lepidocordia Ducke and Rochefortia, sometimes inserted on the centre of an accrescent bracteole (Keraunea); calyx lobes united in a tube or distinct nearly to the base, tubular to campanulate; aestivation imbricate (mostly quincuncial); corolla sympetalous, generally tubular with spreading lobes, rotate, or campanulate to urceolate, white, red or blue (Halgania, some species of Bourreria P.Browne); stamens 5, the filaments generally adnate to the corolla tube at least at the base, sometimes puberulent at the point of insertion, the anthers usually exerted; gynoecium bicarpellate, the ovary uni- to tetralocular from secondary subdivision, style terminal, the stigma clavate to capitate with 1(2) branches; nectar disc usually present at base of the ovary. Fruits drupaceous, often drying and separating into two two-seeded pyrenes, or 4 1-seeded pyrenes or schizocarps, or 4 nutlets.

Ehretiaceae is a broadly distributed family found throughout tropical and subtropical Asia, Australia, sub-Saharan Africa. In the Americas, its distribution encompasses the eastern United States, Florida, Central America, the Caribbean, the Guyana shield and the Andes. In Brazil, the Ehretiaceae was previously only known from the single species Lepidocordia punctata Ducke (Stapf 2023), found in lowland Amazonian forests in Pará and Roraima states. Our treatment therefore represents new records of the family from the Caatinga and Mata Atlântica phytogeographic regions in Brazil, and from the states of Bahia, Minas Gerais, Espírito Santo and Rio de Janeiro (Fig. 1).

The family includes the following eight genera: Bourreria, Cortesia, Ehretia, Halgania, Keraunea, Lepidocordia Ducke, Rochefortia and Tiquilia Pers. (Luebert et al. 2016).

Our morphological concept of the Ehretiaceae is little changed from that of Luebert et al. (2016). The characters that differ are included in bold in the description above. The first of these is that the four species of Keraunea are the first lianescent species included within the family (versus perennial herbs, shrubs, or trees). Lianas are elsewhere found in the Boraginales in the Cordiaceae (Cordia L.) and Heliotropiaceae (Tournefortia L.). Secondly, we have expanded the concept of the Ehretiaceae to include species with a few-flowered corymb inflorescence structure and where the flower and later fruit are inserted at the centre of an accrescent bracteole. To our knowledge, these characters are unique among not just the Ehretiaceae but the Boraginales, whose members are known for their characteristic scorpioid cymose inflorescences. We suggest the few-flowered inflorescence of Keraunea is the result of secondary reduction rather than a retained ancestral form. Within the Boraginales, reductions to few-flowered inflorescences are present elsewhere in the Boraginaceae, Codonaceae and Wellstediaceae (Luebert et al. 2016).

Keraunea brasiliensis Cheek & Simão-Bianchini.

Scandent shrubs or lianas. Stems woody, cylindrical, hollow, lacking lenticels. Stipules lacking. Leaves on the main stems alternate, simple, petiolate; margins entire; venation pinnate, camptodromous or brochododromous. Side shoots with 2–4 aborted leaves, then 2–6 progressively larger leaves along the shoot. Inflorescences terminal on side shoots, determinate, corymbose, with 3–6 flowers; bracteoles one per flower or rarely lacking, leaf-like, inserted halfway along the pedicel. Flowers 5-merous, cosexual. Calyx campanulate, fused at the base, alternating with the petals; aestivation imbricate. Corolla with the tube campanulate, fused at the base; aestivation imbricate. Stamens epipetalous, inserted at the base of the corolla tube, alternating with the petals; filaments free; anthers basifixed, introrse, dehiscing via lateral slits; connectives extended. Nectary disk present, at base of the ovary. Ovary superior, subglobose, 2-locular, the locules biovulate, each with one functional and one aborted ovule. Style single, with two apical stigmas. Fruit a dry, indehiscent drupe, usually inserted in the centre of the persistent and accrescent bracteole; calyx persistent; corolla caducous; style persistent.

Keraunea is endemic to Brazil and thus far known from the states of Bahia (3 spp.), Espírito Santo (1 sp.), Minas Gerais (1 sp.) and Rio de Janeiro (1 sp.). Each of the species is restricted to a single state except for K. capixaba, which is found in both Bahia and Espírito Santo (Fig. 2).

Figure 2. 

Distribution of species of the genus Keraunea. Shading denotes elevation; two letter codes represent the standardised two letter codes for the states of Brazil. Species are coloured as follows: blue, K. brasiliensis Cheek & Simão-Bianchini; black, K. bullata Moonlight & D.B.O.S.Cardoso; yellow, K. capixaba Lombardi; red, K. confusa Moonlight & D.B.O.S.Cardoso; green, K. velutina Moonlight & D.B.O.S.Cardoso.

Two species are known from seasonally dry forest within the Caatinga domain (K. brasiliensis and K. confusa); two from humid forests within the Mata Atlântica domain (K. capixaba and K. velutina); and a fifth from transitional, semi-deciduous forests between the Caatinga and Mata Atlântica domains (K. bullata). The genus has a marked preference for rock outcrops or forest on rocky soils. Both K. brasiliensis and K. confusa are found mostly on karstic outcrops, while K. capixaba and K. velutina have both been collected at the base of granitic inselbergs.

The epithet derives from the Greek, keraunos, or lightning bolt. This was intended to signify the unexpected but now disproven appearance of a neuropeltoid genus of the Convolvulaceae in the Americas.

Cheek and Simão-Bianchini (2013) treated the flower and fruit of Keraunea as being inserted at the centre of a large, leaf-like bract. This ‘bract’ is inserted halfway up and adnate to the pedicel, so we instead follow Lombardi (2014) and treat it as a bracteole rather than a bract.

Keraunea species are distinctive in being semi-scandent shrubs or lianas with simple, alternate, exstipulate leaves and inflorescences terminal on foliose side shoots. The flowers and fruits are highly unusual in being inserted at and appearing to arise from the centre of an accrescent bracteole.

Several specimens of Keraunea were erroneously identified as Bougainvillea Comm. ex Juss. (Nyctaginaceae), which is understandable, as both genera have exstipulate leaves, a semi-lianescent habit, and flowers and fruits associated with showy bracts or bracteoles. Keraunea spp. are however never spinescent, and the flowers and fruits are inserted directly onto the bract, whereas in Bougainvillea several flowers or fruits are clustered on a pedunculate inflorescence surrounded by two or more bracts.

Brazil. Bahia: Mun. Caetité, caminho da Fazenda Boa Vista para Urânio, 13°59'35"S, 42°12'27"W, 560 m, 8 Feb 1997, L. Passos, M.L. Guedes, B. Stannard & E. Saar 5263 (holotype: SPF; isotypes ALCB, CEPEC [CEPEC00077827], HRCB [acc. #38156], HUEFS [HUEFS000058973], K [K000979156]).

Scandent shrub or liana, to 7 m tall. Stems cylindrical, hollow, 3–9 mm in diameter, frequently branching, sericeous-pubescent with silky hairs; internodes 0.7–11 cm long; side shoots 4–9 cm long, hirtellous-pubescent with minute, rigid hairs. Leaves of main stem with the blades 5.0–6.5 × 3.1–3.6 cm, ovate, hirtellous-pubescent with minute, rigid hairs, the venation pinnate, camptodromous, with 4–6 secondary veins; the base acute, the margins entire, ciliate, the apex obtuse; petioles of leaves on main stem ca. 8 mm long, distinctly curved to 90° ⅓ of the way along their length. Side shoots with 3–5 leaves, these progressively larger along the shoot; blades 5.1–7.2 × 2.3–3.9 cm, ovate, hirtellous-pubescent, the venation as on leaves of the main stem, the base acute, the margins entire, ciliate, the apex obtuse; petioles 5.7–6.1 mm long, straight, hirtellous-pubescent like the leaves. Inflorescence corymbose, with 2–4 flowers; free portion of the peduncle ca. 3.5 mm long, 2.5 cm long, the remainder adnate to the bracteole, hirtellous-pubescent; bracteole 5.1–8.1 × 2.4–3.8 cm, inserted ca. ¼ of the way along the pedicel, ovate, hirtellous-pubescent, the venation as the leaves, the base acute, the margins entire, ciliate, the apex obtuse. Flowers incompletely known, 5-merous. Calyx with the tube campanulate, ca. 3.5 mm long, the lobes ca. 12 × 1.5 mm, lanceolate, velutinous-pubescent. Corolla unknown; androecium and gynoecium unknown. Fruit inserted on the accrescent bracteole; calyx persistent, expanding to 14 mm long.

Figure 3. 

Habitat and morphology of Keraunea brasiliensis Cheek & Simão-Bianchini A–C karstic Caatinga in southwestern Bahia D stem and leafy side shoots E habit F main shoot and side shoots with fruits G–H close up of developing fruits and accrescent bracteole. All photos by Domingos Cardoso.

Keraunea brasiliensis is endemic to Brazil and is known only from the state of Bahia (Coribe, São Félix do Coribe and Caetité municipalities) (Fig. 2).

Keraunea brasiliensis has primarily been collected scrambling over exposed rocks and vegetation in karst formations supporting Caatinga seasonally dry tropical forest vegetation. The two most easterly collections of the species (Pereira-Silva et al. 9111, Passos et al. 5263) were collected in Caatinga vegetation but far from any known karst outcrops.

Keraunea brasiliensis was provisionally assessed by Cheek and Simão-Bianchini (2013) as Endangered (EN B2 a b iii) under IUCN Criteria (IUCN 2019). We alter this assessment as it covered specimens now known to be of two different species. Our narrower concept includes plants from four localities and has an EOO of 4,550 km². None of the known localities are in protected areas. Accordingly, we still assess K. brasiliensis as EN (EN B2 a b iii) under IUCN criteria (IUCN 2019).

Named for the country of Brazil.

The protologue of K. brasiliensis cited individuals we now recognise as two different species and both paratypes collected in Minas Gerais fall under our circumscription of K. confusa. The authors of K. brasiliensis did, however, note the unusual intraspecific morphological variation within the species and speculated that the material collected in Bahia and Minas Gerais may have represented two different species. The original illustration in the protologue (Cheek and Simão-Bianchini 2013) included individuals of both of the species we recognise here so only the subfigures (B, H–I) represent K. brasiliensis. Muñoz-Rodríguez et al. (2022) also published sequence data for K. brasiliensis and K. confusa and a member of the family Malpighiaceae under the name K. brasiliensis.

Our emended description of K. brasiliensis reflects our much narrower circumscription of this species. The specimens we cite represent a morphologically homogenous group of specimens all collected at a similar phenological state, i.e. with fruits but lacking flowers. As such, our description lacks some floral traits. The floral description of K. brasiliensis provided by Cheek and Simão-Bianchini (2013) was made using a dissected bud from Lombardi & Salino 1819 (K), which we now consider as an isotype of K. confusa (see below).

Within its range, K. brasiliensis is most likely to be confused with K. confusa, but the two species can be distinguished with ease based entirely upon vegetative characters. The leaves of K. confusa are larger (to 18 × 12 cm versus to 7.5 × 4 cm) with brochidodromous venation and a rugose texture (versus camptodromous venation and smooth texture in K. brasiliensis). Keraunea brasiliensis is most similar to K. capixaba (though this species is also similar to K. velutina), from which it can be distinguished by its rugose stems (versus smooth) and membranous, dull green leaves (versus chartaceous, glossy green) with indistinct secondary veins (versus raised above the lower leaf surface).

Brazil. Bahia: Mun. Coribe, BA-172 sentido São Félix do Coribe, entrada a direita em Colônia do Formoso por estrada de terra até afloramento de calcário a ca. 2.5 km de Ponta d’Água, 13°41'28"S, 44°15'33"W, 4 Dec. 2022, 4 Dec 2022, D. Cardoso et al. 4902 (ALCB, RB, TCD), D. Cardoso et al. 4904 (ALCB, HUEFS, RB, TCD), D. Cardoso et al. 4906 (ALCB, HUEFS, RB, TCD; Mun. São Félix do Coribe, BA-172 sentido Coribe, afloramento de calcário do lado esquerdo, 13°26'6"S, 44°13'5"W, 4 Dec. 2022, D. Cardoso et al. 4909 (ALCB, HUEFS, RB, TCD), D. Cardoso et al. 4910 (ALCB, HUEFS, RB, TCD), D. Cardoso et al. 4912 (ALCB, HUEFS, RB, TCD); Mun. Caetité, Distrito de Maniaçu, Estrada para São Timóteo, km 6, 13°51'18"S, 42°20'45"W, 870 m alt., 22 May 2004, G. Pereira-Silva et al. 9111 (HUEFS [HUFES000117294]); Mun. Santa Maria da Vitória, c. 7.7 km S de Santa Maria da Vitória na Estrada para Lagoinha, extremidade septentrional da Serra do Ramalho, 13°27'0"S, 44°10'16"W, 13 Feb 2000, L.P. de Queiroz et al. 5972 (CEPEC [CEPEC00113512]; HUEFS [HUEFS000110252]); Mun. Coribe, c. 5 km S em estrada de terra que cruza pequeno ramal que sai a 5.1 km E de Ponta d’Água, a 24.4 km S de São Félix do Coribe na estrada para Coribe, 13°35'10"S, 44°19'12"W, 14 Apr. 2007, L.P. de Queiroz et al. 12707 (CEN [CEN00113310], HUEFS [HUEFS000117258]).

Most similar to K. confusa with relatively broad leaves with brochidodromous venation but differing from it in smaller leaves (reaching 10 × 7.4 cm versus 18 × 12 cm) with a bullate texture (versus rugose) and by its tertiary veins, which are raised from the lower leaf surface (versus not raised).

Brazil. Bahia: Mun. Jequié, Bairro Suíça, 13°51'S, 40°5'W, M.L. Guedes, D.M. Loureiro & D.L. Santana 9176 (holotype: ALCB [acc. #55929, ALCB075866, 3 sheets]).

Scandent shrub or liana, to 2 m. Stems cylindrical, hollow, 2–5 mm in diameter, frequently branching, glabrescent to sparsely pubescent; internodes 2.9–8.2 cm long; side shoots 3.0–4.5 cm long, sericeous. Leaves of main stem unknown. Side shoots with 3–4 leaves, these progressively larger along the shoot; blades 4.2–10.5 × 2.4–7.4 cm, ovate to broadly ovate, velutinous-pubescent, the venation pinnate, brochidodromous, with 4–6 secondary veins; the base rounded to truncate, the margins entire, ciliate, the apex obtuse; petioles 8–19 mm long, straight, velutinous-pubescent. Inflorescence corymbose, with 2–6 flowers; free portion of the peduncle 4–6 mm long, 1.8–2.1 cm long, the remainder adnate to the bracteole, sericeous-pubescent; bracteoles 5.0–5.7 × 2.7–3.1 cm, inserted ca. ¼ of the way along the pedicel or rarely lacking, ovate, velutinous-pubescent, the venation as the leaves, the base short-rounded, the margins entire, ciliate, the apex attenuate. Flowers incompletely known, 5-merous. Calyx with the tube campanulate, ca. 1.5 mm long, the lobes ca. 1.5 × 10–12 mm, ensiform, sericeous-pubescent. Corolla unknown; androecium and gynoecium unknown. Fruit inserted on the accrescent bracteole; calyx persistent, expanding to 12 mm long.

Figure 4. 

Holotype of Keraunea bullata Moonlight & D.B.O.S.Cardoso, M.L. Guedes, D.M. Loureiro & D.L. Santana 9176 (ALCB [ALCB075855, acc. #55929, sheet 1]).

Endemic to the state of Bahia (Fig. 2).

Keraunea bullata is a species of forests in the transition from humid forests in the coastal Mata Atlântica phytogeographic domain and seasonally dry forests in the inland Caatinga domain. This forest is described on labels as “Caatinga arborea” (Guedes et al. 9176) and (Mori et al. 11534) as growing on rocky soils. This area has a metamorphic bedrock (often called crystalline in the literature, de Queiroz et al. 2017) and is quite edaphically distinct from the karstic outcrops that are home to K. brasiliensis and K. confusa.

Keraunea bullata is known from only three localities with a combined EOO of ca. 6,300 km². We know of no specific conservation threats to this species so provisionally assess it as Vulnerable (VU D2) under IUCN Red List criteria (IUCN 2019).

Named for the bullate texture of the leaves and bracteoles.

Keraunea bullata is highly distinct within its range. Despite also occurring in the same Mata Atlântica phytogeographic domain as K. capixaba, their habitat does not overlap. Keraunea bullata can be morphologically distinguished by its leaf venation (brochidodromous versus camptodromous) and texture (bullate and dull green versus smooth and glossy green).

Brazil. Bahia: Mun. Itambé, rod. BA-265, trecho Itapatinga/Caatiba, Faz. Serra Verde a 17 km da Rod. BR-415, [15°6'S, 40°22'W], 14 Mar. 1979, S.A. Mori et al. 11534 (CEPEC [CEPEC00016207], NY [03147645], RB [00725192]). Mun. Juraci, Vicinity of Machado Portela [13°10'S, 40°46'W], 19–23 Jun 1915, J.N. Rose & P.G. Russell 19979 (US [01341072, acc. # 762304).

Brazil. Espírito Santo: Mun. Jaguaré, perto da Comunidade São Jorge de Paduá, sentido para Fátima, 18°54'29.0"S, 40°8'44.9"W, 26 Sep. 2013, G.S. Siqueira 891 (holotype: CVRD [acc. # 14565]; isotypes HRCB [acc. # 76196], SP [SP003725]).

Scandent shrub or liana, to ca. 4 m. Stems cylindrical, hollow, 2–4 mm in diameter, glabrescent, rarely branching, internodes 2.4–11.5 cm long; side shoots 1.3–5.5 cm long, glabrescent. Leaves of main stem with the blades 7.0–13.5 × 2.1–4.5 cm, elliptic, glabrescent, the venation pinnate, camptodromous, with 4–6 secondary veins; the base cuneate, the margins entire, ciliate, the apex attenuate; petioles 5–6.5 mm long, straight. Side shoots with 4–6 leaves, these progressively larger along the shoot; blades 1–9.5 × 0.5–6.5 cm, ovate, rarely obovate or broadly ovate, glabrescent, the venation as on leaves of the main stem, the base cuneate to rounded, the margins entire, rarely crenulate toward the apex (Siqueira 891), ciliate, the apex acute to rounded; petioles 5.5–12 mm long, straight, glabrescent. Inflorescence corymbose, with 2–4 flowers; free portion of the pedicel, 8–14 mm long, 1.8–2.6 cm long, the remainder adnate to the bracteole, sericeous-pubescent with silky hairs; bracteole 3.3–3.5 × 1.9–2.4 cm, inserted ca. ½ of the way along the pedicel or rarely lacking, ovate, glabrescent, the venation as the leaves, the base short acute, margin entire, not ciliate, the apex acute. Flowers 5-merous. Calyx with the tube campanulate, ca. 3.5 mm long, the lobes 6.5–18 × 1.5–2.5 mm, ensiform, glabrescent to minutely pubescent. Corolla with the tube campanulate, ca. 6.5 mm long, the lobes 19–26 × 7.5–11.5 mm, elliptic to oblong, glabrous. Stamens epipetalous, inserted at the base of the corolla tube, the filaments 2–3 mm long, the anthers ca. 9 mm long, connective extending to 3.2 mm. Ovary subglobose, 2-locular, the locules biovulate; style single, conduplicate, unbranched; stigmas 2, truncate. Fruit inserted on the accrescent bracteole; calyx persistent, expanding to 18 mm long.

Figure 5. 

Morphology of Keraunea capixaba Lombardi A habit B inflorescence, front view C inflorescence, side view. For photograph of fruit with accrescent bracteole see Lombardi (2014). All photographs by Geovane Siqueira.

Endemic to Espírito Santo state and known from the municipalities of Jaguaré, Sooretama and Nova Venécia (Fig. 2).

Collections of Keraunea capixaba have primarily been made in “mata tabuleiro” or flat, semi-deciduous forests found within the coast of the Mata Atlântica domain of Brazil. A more recent collection (Gurtler & Dutra 371) was made growing over rocks in forest understory at the base of a granitic outcrop. Specimen labels describe the species as scandent or lianescent.

Keraunea capixaba was provisionally assessed by Lombardi (2014) as Endangered because it was at that time known from four collections made within disturbed forest patches adjacent to the Sooterama Biological Reserve. We add five collections to the known distribution, including one collected within the Sooterama Biological Reserve (Covre s.n.), two collected at the base of inselbergs > 30 km from the reserve (Gurtler & Dutra 371, Demuner et al. 3799), and a gathering made a remarkable 500 km to the north near Ipauí, Bahia. The species is now known from six localities with a collective Extent of Occurrence (EOO) of ca. 11,000 km². The species is known from < 10 localities and has a EOO of < 20,000 km², so we assess K. capixaba as Vulnerable (VU B1 a b iii) under IUCN criteria (IUCN 2019).

The epithet is an indigenous term referring to people or objects from Espírito Santo state.

Keraunea capixaba is most morphologically and ecologically similar to the newly described K. velutina, the two species of the genus that are associated with more humid settings in the Mata Atlântica phytogeographic region, yet they were never found growing sympatrically. Keraunea capixaba differs in lacking an indumentum on the stems, side shoots and lower leaf surface (versus a velutinous-pubescent in K. velutina), and its ovate leaves on the side shoots (versus elliptic to narrowly-lanceolate). Its range is close to that of K. confusa, from which it is readily distinguished by its camptodromous venation (versus brochidodromous).

Brazil. Bahia: Rod Ipiau-Ibirataia, [14°6'S, 39°41'W], 13 Nov. 1971, T.S. Santos 2139 (HUEFS). Espírito Santo: Mun. Sooretama, Reserva Biológica de Sooretama, [19°1'S, 40°7'W], 19 May 2015, C. Covre s.n. (SAMES [SAMES03696]); Mun. Rio Bananal, Alto Bananal, 19°14'56"S, 40°24'59"W, 300–600 m alt., 25 Apr. 2007, V. Demuner et al. 3799 (BHCB [BHCB017211]); Mun. Jaguaré, Perto da Comunidade São Jorge de Paduá, 25 Sep. 2013, D.A. Folli 7117 (CVRD [acc. #14563]; HRCB [acc. #16194], NY [02687787], RB [00895205]); Mun. Sooretama, Barro Roxo a Córrego Rodrigues, 19°4'35"S, 40°13'25"W, 156 m alt., 8 Oct. 2014, D.A. Folli 7273 (CVRD [acc. #15122]; RB [01103439]); Mun. Nova Venécia, fazenda Santa Rita, ao pé da pedra da torre (P3), 18°47'7"S, 40°26'29"W, 2 Feb. 2018, J. Gurtler & S.C. Dutra 371 (VIES [VIES036853]); Mun. Sooretama, Rodovia ES 358, distrito de Bom Jardim, 19°2'20.31"S, 40°14'48.23"W, 1 Sep. 2012, A. Moreira de Assis & J. Freitas 3340 (HRCB [acc. #76193], MBML [2 sheets: MBML00016754, MBML00016755]); Mun. Jaguaré, perto da Comunidade São Jorge de Paduá, 18°54'29.0"S, 40°8'44.9"W, 6 Oct. 2013, G.S. Siqueira 893 (CVRD [acc. #14570], HRCB [acc. #76199], K [K001275507], MBML [MBML00016491], NY [02687786], RB [00895202])

Most similar to K. brasiliensis but differing in its larger leaves (to 18 × 12 cm versus to 7.5 × 4 cm) with brochidodromous (versus camptodromous) leaf venation and a rugose (versus flat) leaf texture and by its leaves and stems with a sericeous (versus hirtellous) indumentum.

Brazil. Bahia: Mun. Januária, distrito de Fabião, junto ao Abrigo do Malhador, 15°7.85'S, 44°15.17'W, 25 May 1997, J.A. Lombardi & A. Salino 1819 (holotype: BHCB [BHCB017209]; isotypes: HRCB [acc. #26438]; K [K001395055], UEC [UEC117783]).

Scandent shrub or liana, to 4 m tall. Stems cylindrical, hollow, 2–5 mm in diameter, rarely branching, sericeous-pubescent with silky hairs; internodes 3.8–16.5 cm long; side shoots 2.6–3.5 cm long, sericeous-pubescent. Leaves of the main stem with the blades 9.5–17 × 5.6–13.5 cm, ovate, velutinous-pubescent, the venation pinnate, brochidodromous, with 5–7 secondary veins, the base truncate to subcordate, the margins entire, ciliate, the apex attenuate to acute; petioles of leaves on main stem 10–16 mm long, distinctly curved to 90° ¼ of the way along their length. Side shoots with 2–5 leaves, these progressively larger along the shoot; blades 2.1–18 × 3.0–12 cm, velutinous-pubescent, the venation as on leaves of the main stems, the base rounded to truncate, the margins entire, ciliate, the apex obtuse; petioles 3–25 mm long, sericeous-pubescent. Inflorescence corymbose, with 2–4 flowers; free portion of the peduncle 5–13 mm long, 2.2–2.8 cm long, the remainder adnate to the bracteole, sericeous-pubescent; bracteole 4.9–5.6 × 2.7–5.1 cm, inserted ca. ⅓ of the way along the pedicel or rarely lacking, ovate, sericeous-pubescent, the venation as the leaves, the base short cuneate, the margins entire, ciliate, the apex attenuate. Flowers incompletely known, 5-merous. Calyx with the tube campanulate, ca. 1.5 mm long, the lobes ca. 7–9 × 1.5 mm, ensiform, sericeous-pubescent. Corolla unknown; androecium and gynoecium unknown. Fruit inserted on the accrescent bracteole; calyx persistent, expanding to 9 mm long.

Figure 6. 

Habitat and morphology of Keraunea confusa Moonlight & D.B.O.S.Cardoso A–C karstic Caatinga seasonally dry tropical forest in northern Minas Gerais D–G habit H–I branching pattern J close up of side shoot. All photographs by Domingos Cardoso.

Keraunea confusa is endemic to Brazil and to the state of Minas Gerais (Fig. 2).

All collections of K. confusa are from karstic (limestone) areas, where plants have been collected growing over rocks and vegetation in the understory of Caatinga seasonally dry tropical forest vegetation.

Keraunea confusa is known from two localities and has an AOO of ca. 12 km². Two of the known collections (Lombardi & Salino 1819, Lombardi 2107) were made within Parque Nacional Cavernas do Peruaçu, which includes extensive karst limestone habitat suitable for the species. We know of no specific threats to this species but provide a preliminary assessment as Vulnerable (VU D2) because of the species’ few known localities (IUCN 2019).

The epithet refers to the confusion that has surrounded the taxonomic history of this species, which was both included within the original circumscription of the type species and the placement of the genus in the wrong family by Cheek and Simão-Bianchini (2013).

Keraunea confusa is one of two species of the genus with brochidodromous venation, the other of which is K. bullata. It is distinguished from K. bullata by its generally bigger leaves measuring 9.5–17 cm long (versus 4.2–10.5 cm long), rugose (versus bullate) leaf texture and by the tertiary veins, which are plane to the lower leaf surface (versus raised).

(paratypes). Brazil. Minas Gerais: Mun. Manga, BR-135, entre Montalvânia e Manga, afloramento de calcário do lado esquerdo a ca. 400 m da rodovia sentido Manga-Januária, 14°30'44.8"S, 44°10'31.8"W, 5 Dec. 2022, D. Cardoso et al. 4916 (ALCB, HUEFS, TCD, RB), D. Cardoso et al. 4918 (ALCB, HUEFS, TCD, RB); Mun. Januária, distrito de Fabião, junto ao Abrigo do Malhador, 15°7'16”–15°8'57"S, 44°15'20”–44°14'13"W, 26 Oct. 1997, J.A. Lombardi 2107 (BHCB [BHCB017210], K [K000593363]).

Most similar to Keraunea capixaba but differing in its velutinous indumentum on the stems, side shoots and lower leaf surface (versus glabrescent); its strigose indumentum on the upper leaf surface (versus glabrescent); its elliptic to narrowly lanceolate leaves on the side shoots (versus ovate).

Brazil. Rio de Janeiro: Mun. Cardoso Moreira, Santíssimo, Fazenda Borges, [21°29'S, 41°37'W], 10 Sep. 2013, J.G. Costa 257 (holotype: RB [00852871]).

Scandent shrub or liana, size unknown. Stems cylindrical, hollow, 2–3 mm in diameter, rarely branching, velutinous-pubescent; internodes 2–5.9 cm long; side shoots 2.2–4.7 cm long velutinous-pubescent. Leaves of the main stem with the blades 7.9–8.1 × 2.7–3.1 cm, lanceolate, velutinous-pubescent, the venation pinnate, camptodromous, with 6–8 secondary veins; the base obtuse, the margins entire, ciliate, the apex attenuate; petioles of leaves on main stem 10–11 mm long, distinctly curved to 90° ¼ of the way along their length. Side shoots with 6–8 leaves, these progressively larger along the shoot; blades 0.5–3.5 × 0.3–1.7 cm, elliptic, rarely or lanceolate or obovate, strigulose-pubescent above, velutinous below, the base cuneate to rounded, the margins entire, ciliate, the apex rounded to attenuate. Inflorescence corymbose, with 2–4 flowers; free portion of the peduncle 3–5 mm long; pedicel 0.5–1.5 cm long, the remainder adnate to the bracteole, sericeous-pubescent; bracteole 2–3.8 × 0.7–1.6 cm, inserted ca. ⅓ of the way along the pedicel or rarely lacking, ovate, velutinous- pubescent, the venation as the leaves, the base short-cuneate, margin entire, ciliate, the apex attenuate. Flowers 5-merous. Calyx with the tube campanulate, ca. 1.5 mm long, the lobes 0.7–1.4 × 6.7–12 mm, ensiform, velutinous-pubescent. Corolla with the tube campanulate, 5–5.9 mm long, the lobes 14 × 7 mm, elliptic, glabrous. Stamens epipetalous, inserted at the base of the corolla tube, filaments 3–4 mm long, anthers ca. 7 mm long, connective extending to 1.3 mm; ovary subglobose, 2-locular, the locules biovulate; style single, conduplicate, unbranched; stigmas 2, truncate. Fruit inserted on the accrescent bracteole; calyx persistent, expanding to 15 mm long.

Figure 7. 

Holotype of Keraunea velutina Moonlight & D.B.O.S.Cardoso, J.G. Costa 257 (RB [00852871]).

Keraunea velutina is endemic to Brazil and known only from the type from Cardoso Moreira municipality in the state of Rio de Janeiro (Fig. 2).

The single specimen known of K. velutina was collected within a forest fragment around isolated inselbergs. Cardoso Moreira municipality includes several granitic inselbergs and deforestation appears reduced around these compared to the surrounding land. We suspect that, like the similar Keraunea capixaba, this species was collected scrambling over rocks at the base of an inselberg.

Keraunea velutina is known from a single specimen collected within a small forest fragment. Forest cover within Cardoso Moreira municipality declined from 5.2% in 1985 to 4.0% in 2012 before recovering to 4.6% by 2021 (Mapbiomas 2022). This picture is however complex, with Google Earth imagery showing that deforestation of primary forest is continuing in some areas, but forest is recovering in other areas (Google Earth Pro 2020). As such, due to single collection with imprecise locality information from a largely deforested municipality, we provisionally assess K. velutina as Critically Endangered (CR B1 B2 a b ii) under IUCN criteria (IUCN 2019).

Named for the plant’s velvety indumentum.

Keraunea velutina is a distinct species easily distinguished from all others in the genus by the dense, velvety indumentum of the stems and lower leaf surfaces. It is also distinctive in its side shoots with more numerous leaves than other members of the genus (6–8 versus 2–5), which are almost all elliptic (versus variously ovate to broadly ovate).