Lippia carrascoana P.H.Cardoso & Salimena, 2025

Lippia carrascoana P.H.Cardoso & Salimena sp. nov. ( Fig. 1 View FIGURE 1 , 2 View FIGURE 2 ).

Type:— Meruoca, Sítio Bom Jesus, Serra da Meruoca , 17 February 1981, A. Fernandes & Matos s.n. ( holotype EAC-9699, isotype CESJ-83506)

Diagnosis:— Lippia carrascoana is a distinct species within the Lantana / Lippia complex due to the unique combination of the following traits: leaf blades with margins entire near the base or up the middle, conspicuously dentate towards the apex, abaxial surfaces covered with abundant sessile glandular trichomes and strigose along the veins, wide-ovate, ovate, or lanceolate bracts, with attenuate or aristate apices, nearly covering the corolla tubes, lilac or white corollas without eglandular trichomes externally, densely sericeous ovaries, and drupaceous fruits with two pyrenes, pilose in the external surface.

Shrubs 0.5‒1.8 m tall, aromatic, erect, branches tetragonal, unarmed, strigose-glandulous, white eglandular trichomes and golden sessile glandular trichomes, becoming glabrescent with age. Leaves opposite, patent, petioles 0.3‒2.2 cm long, obscured by the decurrent portion of the leaf blades, blades 1.5‒7.7 × 0.6‒3.8 cm, chartaceous, concolor, ovate or lanceolate, apex acute or obtuse, base attenuate or truncate-attenuate, decurrent into petiole, margins entire near the base or up the middle, conspicuously dentate towards the apex, teeth ca. 0.3 cm long, plan or slightly revolute, ciliate, with white eglandular trichomes, adaxial surface bullate, strigose or puberulent, with white, short and long eglandular trichomes oriented in multiple directions, abaxial surface covered with abundant sessile golden glandular trichomes, densely strigose along the veins, with long white eglandular trichomes, primary and secondary veins strongly prominent. Inflorescences 1 per leaf axil, many-flowered, spikes capituliform, 1‒1.4 × 1‒1.2 cm, elongated up to 2.5 cm during the fructification, rachis strigose, peduncles 1‒4.5 cm long, tetragonal, strigose-glandulous, with white eglandular trichomes and golden sessile glandular trichomes; bracts congested, unequal, external ones larger, wide-ovate or ovate, 0.9‒1.3 × 0.4‒0.7 cm, internal ones narrower, lanceolate, 0.7‒0.9 × 0.2‒0.5 cm, apex attenuate or aristate, nearly covering the corolla tubes, membranaceous, green, abaxial surface strigose-glandulous, with long white eglandular trichomes and abundant sessile golden glandular trichomes, adaxial surface minutely strigose, with short white eglandular trichomes, margins conspicuously ciliate, with long white eglandular trichomes. Flowers sessile, calyces inconspicuous, ca. 0.1 cm long, green, membranaceous, tubular, apex truncate, externally glabrous or inconspicuously minutely strigose, internally glabrous, not accrescent in fruit; corollas lilac or white, throat yellow or lilac, tube 0.6‒1 cm long, hypocrateriform, externally covered with golden sessile glandular trichomes, limb 2-labiate, covered with golden sessile glandular trichomes, apical lobe more conspicuous, stamens 4, didynamous, inserted halfway of the corolla tube; ovary 0.1‒0.15 cm long, globose, densely sericeous, with long white eglandular trichomes, style filiform, stigma oblique. Fruits drupaceous, 2-pyrenate, 0.3‒0.45 × 0.3‒0.45 cm, globose or subglobose, naked, thin fleshy mesocarp, brown, external surface smooth or slightly striate, sparsely to densely pilose.

Paratypes:— Ceará: Aiuaba, Serra Nova, 13 January 1998, M.A. Figueiredo & L.W. Lima-Verde 942 (EAC-041273, HUEFS-137185); Aiuaba, Serra Nova, Serra do Ermo, 15 January 1998, M.M.A. Souza & J.F.J. Araújo 289 (EAC-41271); Aiuaba, Serra do Ermo, 5 February 1997, L.W. Lima-Verde et al. 373 (EAC-41268); Aiuaba, Estrada da Confiança, Serra do Ermo, 5 February 1997, L.W. Lima-Verde et al. 385 (CESJ-83511, EAC-41249); Aiuaba, Estrada da Confiança, Serra do Ermo, 5 February 1997, L.W. Lima-Verde et al. 399 (EAC-41251); Aiuaba, Gameleira, Estação Ecológica de Aiuaba, 24 May 2013, M.I.B. Loiola & F.R.S. Tabosa 2019 (EAC-55734); Cratéus, Serra das Almas , 26 February 2002, F.S. Araújo & L.C. Girão 1312 (EAC-33955, HUEFS-67599); Cratéus, Reserva Serra das Almas , 17 May 2001, M.S. Sobrinho & M.M.A. Bruno s.n. (EAC-32990); Cratéus, RPPN Serra das Almas , 25 February 2002, F.S. Araújo & L.C. Girão 1292 (CESJ-83510, EAC-33957, HUEFS-67597); Cratéus, RPPN Serra das Almas , 26 February 2002, F.S. Araújo & L.C. Girão 1313 (EAC-22956, HUEFS-67599); Cratéus, Serra das Almas , around plot 1., 29 March 2017, P.W. Moonlight 610 (HUEFS-233564); Cratéus, RPPN Serra das Almas , 3 May 2005, F. Prado & M. Goretti 3 (EAC-35552); Cratéus, RPPN Serra das Almas , 17 May 2001, M.S. Sobrinho 64 (EAC-33381); Cratéus, Serra das Almas , 22 February 2000, L.W. Lima-Verde s.n. (EAC-39337); Cratéus, Serra das Almas , 23 February 2007, E. Silveira s.n. (EAC-39775, UFRN-5841); Cratéus, Tucuns, Sertão de Cratéus, 20 May 1997, M.A. Figueiredo & J. Augusto s.n. (EAC-25607); Guaraciaba do Norte, Andrade, 26 May 1981, A. Fernandes & P. Martins s.n. (EAC-10344); Guaraciaba do Norte, Andrade, 27 February 1981, A.P. Fernandes & P. Martins s.n. (CESJ-83514, EAC-9814); Ipueiras, Sítio Olho d’Água dos Galvão, Serra da Ibiapaba , 31 January 2012, A.S.F. Castro 2616 (EAC-51841); Novo Oriente, Planalto da Ibiapaba , 15 February 1991, F.S. Araújo 279 (EAC-19405); Novo Oriente, Planalto da Ibiapaba , 8 November 1990, F.S. Araújo 187 (EAC-19313); Poranga, congl. 164, sub. 2, 4 February 2014, W. Batista 224 (EAC-56711, RB-787799); São Benedito, Chapada de Ibiapaba , 7 December 1990, A. Fernandes et Matos s.n. (EAC-17082); São Benedito, Faveira, 27 June 1981, A. Fernandes & Matos s.n. (CESJ-83508, EAC-10459); Tianguá, Chapada da Ibiapaba , 29 July 1989, A. Fernandes et al. s.n. (CESJ-83513, EAC-16660); Tianguá, Chapada da Ibiapaba , 13 January 1982, A. Fernandes & Matos s.n. (CESJ-83507, EAC-11070); Tianguá, Serra da Ibiapaba , 11 August 1978, A. Fernandes & Matos s.n. (EAC-4096); Ubajara, Jaburuna/Sul, Planalto da Ibiapaba , 27 January 1996, F.S. Araújo s.n. (EAC-23597); Ubajara, Planalto da Ibiapaba, Faz. Japtaraco , 29 March 1994, F.S. Araújo 614 (EAC-22791); Ubajara, Planalto da Ibiapaba , 3 July 1994, F.S. Araújo 853 (EAC-22790). Viçosa do Ceará, Pindaré, 11 April 1992, F.S. Cavalcanti 46 (EAC-18517). Pernambuco: Moreilândia, 2 April 1995, E.R. Silveira s.n. (EAC-22107). Piauí: Canto do Buriti, Fazenda Itaueira, 9 January 1995, I. Pinto & E. Alves s.n. (EAC-22096); Caracol, PARNA Serra das Confusões , 23 March 2006, R. Barros 2716 (HUEFS-135904, TEPB-23224); São Raimundo Nonato, Serra da Capivara , 19 December 1978, A. Fernandes et al. s.n. (EAC-5176).

Distribution and ecology:— Lippia carrascoana is distributed in the Northeast Region of Brazil, in the states of Ceará, Pernambuco, and Piauí, within the Caatinga domain ( Fig. 3 View FIGURE 3 ). It grows in the carrasco vegetation, at elevations between 500 and 830 m, in dystrophic red-yellow latosol, which are sandy or sandy-loam acidic soils with low nutrient and water availability. Most of the specimens were collected on the Ibiapaba Plateau or Ibiapaba-Piauí District, a sedimentary formation located along the border between the states of Ceará and Piauí with high floristic richness ( Moro et al. 2024). Its steep face descends toward Ceará, while the opposite side slopes gently toward Piauí, forming a characteristic cuesta landform ( Claudino-Sales et al. 2020, Lima et al. 2022a).

Carrasco is a type of xerophilous vegetation within the Caatinga domain that occurs at higher elevations, ranging usually from 500 to 900 meters, on sedimentary terrains with sandy soils. This phytophysiognomy is characterized by a high density of woody individuals, including multi-branched and typically spineless shrubs, lianas, as well as small trees typically ranging from 3 to 4 m in height, with occasional emergent trees reaching 6–8 m in height, which have thin trunks and irregular crowns. Herbaceous plants are scarce, with cacti and bromeliads nearly absent ( Araújo & Martins 1999, Bezerra et al. 2003, Vasconcelos et al. 2010, Moro et al. 2015a). In recent years, some new plant species have been described as occurring in carrasco, highlighting the endemism of this vegetation type (Liola 2013, Santos et al. 2020, 2021, Lima et al. 2022b).

Preliminary conservation assessment:— The Caatinga domain, originally covering 844,000 km ², has experienced severe degradation from extensive anthropogenic activities, including deforestation for wood extraction, livestock grazing, agriculture expansion, and road construction. Compounded by the spread of invasive species, these pressures have resulted in a 40% loss of native vegetation cover, with large areas now undergoing desertification ( Leal 2005, Albuquerque et al. 2012, Antongiovanni et al. 2018).

Among the Caatinga ’s vegetation types, carrasco —where Lippia carrascoana occurs—is particularly threatened. According to Moro et al. (2015b), this vegetation faces increasing pressure from agricultural expansion, driven by the spread of large-scale mechanized monocultures. On the Ibiapaba Plateau , where the new species is most frequently found, habitat loss has intensified since 2011 due to urban expansion, large-scale agriculture, and wind farm development ( Lima et al. 2022a). Additionally, human-induced wildfires have become a major driver of environmental degradation in the region, with 1,707.85 km ² burned between 2005 and 2020 ( Lima et al. 2022a).

Currently, only 7.96% of the Caatinga domain is legally protected, with just 1.3% designated as Full Protection reserves—the most effective category for safeguarding biodiversity (Teixeira et al. 2020). Lippia carrascoana has an EOO of 131,402.651 km 2 and an AOO of 100.000 km 2, occurring in only a few protected areas, including Aiuaba Ecological Station, Serra das Almas Private Natural Heritage Reserve, and Serra das Confusões National Park. However, ongoing habitat degradation continues to reduce both the quality of its habitat and extent of its distribution, suggesting that the new species should be considered as Endangered (EN) under criterion B (B2a,b i, ii, iii).

Etymology:— The specific epithet carrascoana is derived from “carrasco,” referring to the distinctive vegetation type where the species grows.

Vernacular names:— According to specimen labels, Lippia carrascoana is a shrub with a strong, distinctive odor. The following vernacular names appear on vouchers: Alecrim, Camará, Camará-Branco , Camará-da-Serra, Cidreira, Cidreira-Brava, Cidreira-Braba-de-Moreilândia, and Cidreirinha-da-Flor-Roxa.

Taxonomic notes:— Lippia carrascoana displays distinctive reproductive traits that are uncommon within Lantaneae , including corolla tubes lacking eglandular trichomes, densely sericeous ovaries covered with long eglandular trichomes, and fruits with a pilose external surface. In contrast, most species of Lantana and Lippia typically possess corolla tubes covered with eglandular trichomes, as well as glabrous ovaries and fruits.

The presence of ovaries covered with eglandular trichomes has previously been documented only in Lantana restingensis Salimena & Silva (2017: 128) , a species endemic to restinga (a vegetation complex typical of Brazil’s coastal lowlands, established over sea deposits of sandy sediment) in the states of Alagoas and Sergipe, within the Atlantic Forest domain ( Silva et al. 2017, 2025). Our comparative analysis further revealed that the fruits of Lantana restingensis are naked, with a pilose external surface (as depicted in the original illustration), and contain two pyrenes, characteristics shared with Lippia carrascoana . However, Silva et al. (2017) described the fruits of Lantana restingensis as 1-pyrenate and enclosed by the surrounding and pilose calyx. This constitutes a mischaracterization resulting from the fact that the fruits were not dissected.

Although current generic delimitations would require the transfer of Lantana restingensis to Lippia ( Silva & Salimena 2002, Atkins 2004), we refrain from formally proposing a new combination at this time, given the nonmonophyly of both Lantana and Lippia (Lu-Irwing et al. 2021) . A phylogenetic classification of the tribe Lantaneae , following the principles of the PhyloCode, has recently been proposed by O’Leary et al. (2023), in which names can be applied to monophyletic groups anywhere in the hierarchy without regard to rank. On the other hand, we underscore the need for a revised classification within Lantaneae , in accordance with the International Code of Nomenclature for algae, fungi, and plants ( Turland et al. 2018), ideally through the recognition of a broadly circumscribed Lippia , which would minimize the number of required new combinations. Despite their current placement in different genera, these taxonomic and nomenclatural considerations are essential for contextualizing the morphological similarity between Lippia carrascoana and Lantana restingensis . The distinguishing traits between these two species are summarized in Table 1.

Lippia carrascoana also resembles species formerly placed in Lantana sect. Sarcolippia Schauer (1847: 595) , a group characterized by drupaceous fruits containing two pyrenes, which was later transferred to Lippia View in CoL by Silva & Salimena (2002). Species within this group likewise exhibit naked fruits, whereas most Lippia species typically have fruits surrounded by persistent calyces ( Mirra et al. 2024). In addition to Lantana restingensis View in CoL , other species known to have drupaceous fruits with two pyrenes are Lippia aonae Zavatin & P.H.Cardoso in Zavatin et al. (2024: 2), Lippia brasiliensis (Link 1822: 126) T.R.S.Silva View in CoL in Silva & Salimena (2002: 58), Lippia lippioides ( Chamisso 1832: 224) Rusby (1896: 106) View in CoL , Lippia macrophylla Chamisso (1832: 214) View in CoL , Lippia magentea Silva (2001: 472) View in CoL , Lippia maximilianii ( Schauer 1847: 595) T.R.S.Silva in Silva & Salimena (2002: 58), Lippia procurrens ( Schauer 1847: 596) T.R.S.Silva View in CoL in Silva & Salimena (2002: 59), Lippia pubescens ( Moldenke 1970: 435) T.R.S.Silva View in CoL in Silva & Salimena (2002: 59), Lippia pusilla T.R.S.Silva View in CoL in Silva & Salimena (2002: 59), Lippia raoniana P.H.Cardoso View in CoL in Cardoso et al. (2021b: 44), and Lippia spiraeastrum View in CoL (Martius & Schauer in Schauer 1847: 596) T.R.S.Silva in Silva & Salimena (2002: 59). However, they did not emerge as a monophyletic group (Lu-Irwing et al. 2021).

Among the 35 specimens of Lippia carrascoana examined, one was previously misidentified as Lippia brasiliensis View in CoL , four as Lippia magentea View in CoL , and one as Lippia maximilianii . These three species exhibit the closest morphological resemblance to Lippia carrascoana , sharing inflorescence peduncles measuring less than 5 cm in length, and conspicuous bracts with long and narrow apices. However, the new species can be distinguished based on the diagnostic traits presented in Table 1. The remaining specimens were identified as follows: 18 as Lantana sp. , two as Lippia sp. , one as Lantana camara Linnaeus (1753: 627) View in CoL , one as Lantana canescens Kunth (1818: 259) View in CoL , and seven as Lantana fucata Lindley (1824: 788) View in CoL . These misidentifications were observed in herbarium collections and some floristic studies, such as Araújo et al. (2011) and Moro et al. (2014). Although Lantana camara View in CoL , Lantana canescens View in CoL , and Lantana fucata View in CoL show few morphological similarities with Lippia carrascoana , they are among the most collected and well-known species of Lantaneae in Brazil ( Silva 1999, Cardoso et al. 2021c), and are frequently involved in misidentifications. A comparison between Lippia carrascoana and these three common Lantana species involved in past misidentifications is also provided in Table 1.

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Of the three species of Lippia morphologically most similar to Lippia carrascoana , both Lippia magentea and Lippia maximilianii are also found in carrasco, although they are not exclusive to this vegetation type ( Salimena & Cardoso 2025). Lippia magentea also occurs in forest formations, cerrado s.l., and caatinga s.s., but it is restricted to the states of Bahia and Minas Gerais ( Salimena & Cardoso 2025), thus occupying a distributional range that does not overlap with that of Lippia carrascoana . In contrast, Lippia maximilianii has a broader distribution in Brazil, occurring across the states of Minas Gerais, Bahia, Ceará, Maranhão, and Piauí ( Salimena & Cardoso 2025). It primarily inhabits forest formations, being rarely found in carrasco. Like Lippia carrascoana , this species occurs on the Ibiapaba Plateau , with some specimens collected in the municipality of Crateús (vouchers: M.A. Figueiredo s.n., EAC-11870 and J.R. Lima 109). Furthermore, both Lippia carrascoana and Lippia maximilianii were recorded in Serra das Confusões National Park; however, their distributions do not overlap within this protected area, with Lippia maximilianii found in a Seasonally Deciduous Forest in the municipality of Guaribas (voucher: J.A. Siqueira Filho 2595). On the other hand, despite its wide distribution across South America, Lippia brasiliensis was not recorded in carrasco vegetation ( Múlgura et al. 2012, Salimena & Cardoso 2025). This species predominantly inhabits humid environments, particularly the interior of forests ( Salimena & Cardoso 2025).