Calea × crassa (Neurolaeneae: Asteraceae), a new nothospecies from Paraná state, Brazil Author Bueno, Vinicius R. Universidade Federal do Rio Grande do Sul, Programa de Pós-graduação em Botânica, Laboratório de Sistemática Vascular, Prédio 43432, sala 107, Porto Alegre, RS 91501 - 970, Brazil & Jardim Botânico da Fundação Zoobotânica de Belo Horizonte - Belo Horizonte, MG 31364 - 450, Brazil Author Cassol, Ana P. V. Universidade Federal do Rio Grande do Sul, Programa de Pós-graduação em Botânica, Laboratório de Ficologia, Museu de Ciências Naturais - Sema / RS, Porto Alegre, RS 90690 - 000, Brazil Author Leroy, Carolina J. Centro Universitário Una, Belo Horizonte, MG, Brazil Author Bueno, Marcelo L. Universidade Estadual de Mato Grosso do Sul, Laboratório de Macroecologia & Evolução (LAMEV), BR- 163, Mundo Novo, MS. Brazil, 79980 - 000, Brazil Author Heiden, Gustavo 0000-0002-0046-6500 Embrapa Clima Temperado, Rodovia BR 392, km 78, Caixa Postal 403, Pelotas, RS 96010 - 971, Brazil gustavo. heiden @ embrapa. br; https: // orcid. org / 0000 - 0002 - 0046 - 6500 gustavo.heiden@embrapa.br text Phytotaxa 2023 2023-10-03 618 2 133 148 http://dx.doi.org/10.11646/phytotaxa.618.2.3 journal article 270730 10.11646/phytotaxa.618.2.3 bd6ae36e-4c62-4c58-b96e-5b616b1d6693 1179-3163 8406652 Calea × crassa V.R.Bueno & G.Heiden , nothosp. nov. ( Fig. 2 ) Type: — BRAZIL . Paraná , Ponta Grossa , Parque Estadual de Vila Velha , campo recém queimado, 19 March 2012 , J . M . Silva , C . B . Poliquesi , L . T . Maranho & R . Ristow 8265 ( holotype : MBM !, isotypes: FUEL !, FURB !, HUCS !, IRAI , digital image!, RB !) . Calea × crassa is differentiated from the parental species C. ilienii and C. monocephala by the third series of phyllaries or just the apex foliaceous (vs. scarious), margins of the fourth series incised (vs. entire), apex of the fifth series obtuse (vs. rounded). Shrubs 0.35–0.7 m tall. Stems cylindrical, striate, sparsely hispid or hispid to densely hispid, eglandular, castaneous, internodes 10–60.8 (–89.7) mm long. Leaves decussate, petiole 1–3.7 (–6.9) mm long; blades 1.5–5.17 × 0.7–3.11 cm , narrow ovate, sometimes ovate-elliptic, base rounded or obtuse, apex acute, rarely cuneate or obtuse, venation acrodromous basal; margins revolute, thickened, white, serrate, mucronate, (6–) 7–17 (–20–23) teeth, 0.3–3.9 mm long, along the entire margin, sometimes from the middle to the apex; basal to median leaves abaxial surface densely hispid or hispid, rarely sparsely hispid, veins hispid, often densely hispid, glandular-punctate, adaxial surface hispid, sometimes sparsely hispid, eglandular; apical leaves glabrous or sparsely hispid, veins sparsely hispid, sparsely glandular-punctate, sometimes glabrous, adaxial surface glabrous; olivaceous, concolorous, coriaceous. Capitulum solitary or capitulescence cymose, slightly branched, axis 0–2.17 cm long, branching to 2 nd ramifications, 1–3 (–4) secondary stems; peduncle 0.58–6.3 cm long, densely hispid, rarely sparsely hispid to hispid, eglandular. Capitulum heterogamous, radiate, involucrum 7.4–15.3 × 10–19.5 mm , wider than longer, 6-seriate. Phyllaries 6-seriate; outermost series of phyllaries margins white, apex revolute, foliaceous, green, conspicuously or inconspicuously striated; first series of phyllaries blade 10.2–12.8 (–18.2) × 7.2–8.75 (–13.6) mm, longer than involucre, sometimes equal, rarely smaller, longer than second series, sometimes smaller, broadly elliptic or orbiculate to suborbiculate, apex obtuse, sometimes attenuate, rarely mucronate, margins serrate mucronate or entire, thickened, glabrous or densely hispid to hispidulous or hispid to sparsely hispid, glandular-punctate, rarely absent, 4–6 striate; second series blade 9.4–14.5 × 6.1–10.5 mm , obovate, sometimes broadly obovate or suborbiculate, apex acuminate, sometimes obtuse or mucronate, margins entire, often serrate, thin, often thickened, sparsely hispid to hispidulous, glandular-punctate, sometimes glabrous, 6–8 striate; third series blade 8.1–13.4 × 4.7–7.9 mm , panduriform, sometimes broadly oblong, apex acuminate, margins entire, apex white, revolute, sparsely hispidulous, eglandular, sometimes glandular-punctate, foliaceous, sometimes foliaceous with scarious base, green, 12–14 striate; innermost series scarious, yellowish green, glabrous; fourth series blades 7.2–12.8 × 3.5–6.9 mm , obovate, sometimes broadly obovate apex rounded, margins incised, 14–16 striate; fifth series blades 7.2–15.7 × 3.2–8 mm , obovate, sometimes broadly obovate, apex obtuse, margins entire, 12–14 striate; sixth series, blades 10.6–14.5 × 3.2–3.8 mm , narrow oblong, apex rounded, margins entire, 12–14 striate. Receptacle slightly convex, holopaleaceous; paleae 8.1–10.4 mm long, longer than disc floret corolla, narrow-oblanceolate or narrow-elliptic, apex long acuminate, often acuminate, conduplicate, light yellow. Ray florets 12–20, pistillate, corolla 14.3–22.9 mm long, liguliform, tube 3–4.3 mm long, limb 10–18.9 × 2.5–4 mm , oblanceolate or narrow oblong, apex 2-lobulate, often rounded, sometimes 4-lobulate, 5-veined, abaxial surface glandular-punctate, often sparsely glandular-punctate, rarely glabrous, adaxial surface glabrous, yellow; style arms 1.1–1.4 mm long, yellow. Disc florets 50–80, bisexual, corolla (5.2–) 6.2–9 mm long, tubular, tube 1.9–2.9 (–3.4) mm long, lobes 1.1–1.4 (–1.7) mm long, glabrous, yellow; anthers 3–3.4 mm long, light yellow, apical anther appendages ovate; style arms 1–1.5 mm long, linear, yellow. Cypselae 2.5–3 (–3.8) mm long, prismatic or obconic, ray cypselae 3-angled, disc cypselae 4-angled, glabrous, rarely glabrous to pilose, ribs densely hirsute or densely pilose, sometimes pilose or glabrous, black; pappus monotypic, monolength, rarely bilength, free, scales 9–12 (2 distinctly longer, when present), mostly 1.2–1.7 (–2) mm long, longer scales 1.8–1.9 mm long (when present), oblong, sometimes oblanceolate, apex obtuse, sometimes rounded, margins entire, erose in apex. Distribution, habitat and predicted area of occurrence :— Calea × crassa is endemic to the Paraná state ( Fig. 3 ) and has been recorded in two municipalities: Balsa Nova and Ponta Grossa. The two parental species are sympatric in Balsa Nova, but C. ilienii has not been collected in Ponta Grossa ( Fig. 3 ). However, the municipality has the same type of habitat where the parental species occur in Balsa Nova and is geographically very close. In fact, the available collections of the new hybrid are distributed in a range of 40 km . Calea × crassa occurs in moist sites from the high elevation grasslands of the Atlantic Forest biome in an area nearby the transition with the Cerrado, biome locally known as campos gerais (general grasslands). Considering the recorded C. × crassa points of occurrence ( Fig. 3 ) and the modelled predicted area of the parental taxa ( Fig. 4 ), it is possible that the parental species are sympatric in the red areas of the prediction ( Fig. 4 , 5 ) from Paraná state , providing the conditions where the new nothospecies can be found. Informal conservation status :— Calea × crassa could be considered Critically Endangered: B1, B2; C2i; D ( IUCN 2022 ). These different criteria are applicable due the AOO shorter than 10 km ² and EOO also smaller (only two collect locations were found) and few (less than 50) mature individuals known (pers. comm. of collectors). The nothospecies occurs in two protection areas: Área de Proteç„o Ambiental Escarpa do Devoniano (Ambiental Protection Area of the Devonian Cliff—APAED), the largest protected area in the Paraná state ; and one voucher of C. × crassa was collected in Parque Estadual de Vila Velha (Vila Velha State Park). However, according to Harfouche et al. (2019) , from 1986 to 2015, the anthropized areas increased by 23% and the natural grasslands decreased 12% within the APAED, which shows that despite this being a conservation area, the nothospecies is not fully protected. Although the IUCN (2022) states that hybrids cannot be included on the red list of endangered species, we believe that it is important to score the risk of extinction due to the threats to the area reported above and because the parental species have not yet been evaluated for the level of threat. Moreover, conservation involves the management of genetic variability, species diversity and the ecological and evolutionary processes. Considering the role that hybridization plays in ecology and evolution, we strengthen the importance of evaluating hybrid taxa. Thus, based on these premises C. × crassa is considered Critically Endangered (CE). FIGURE 2. Comparison of informative structures to recognize the new nothospecies Calea × crassa (first column) in relation to its parental species C. ilienii (second column) and C. monocephala (third column) ( Neurolaeneae , Asteraceae ). A–C. Shape of first series of phyllaries and indumentum of the abaxial side. D–I. Ray florets and disc florets. J–L. Indumentum of abaxial surface of ray florets corollas. M–R. Cypselae and pappus scales. A, D, E, J, M and N: Calea monocephala drawn from G. Hatschbach et al. 35885 (MBM). B, F, G, K, O and P: Calea × crassa drawn from J.M. Silva et al. 8265 (MBM). C, H, I, L, Q and R: Calea ilienii drawn from T.F. Santos 347 (ICN). A–R: millimeter scale. Illustration by Débora Dalzotto. FIGURE 3. Occurrence map of Calea × crassa , C. ilienii , and C. monocephala ( Neurolaeneae , Asteraceae ) in Brazil. PR: Paraná State, SC: Santa Catarina State. Etymology :—The epithet “ crassa ” was chosen because the nothospecies have, as well as its parental species, thickened pale olivaceous mucronate serrate margins ( Fig. 1a, c ). Phenology: —The studied specimens were found flowering and fruiting only in March. FIGURE 4. Prediction occurrence map of suitable ecological niche modelling for Calea ilienii and C. monocephala ( Neurolaeneae , Asteraceae ). Taxonomic comments: —The specimens of Calea × crassa studied here were previously identified as their parental species due to their shared morphological similarities, especially the leaves and foliaceous phyllaries with margins serrate mucronate that are thickened and white ( Fig. 1 ). In addition to the margin of leaves and phyllaries already mentioned, Calea × crassa shares with its parents hispid stems and peduncles, slightly branched cymose capitulescences or solitary capitula, cypselae commonly with glabrous intercostal surface and densely hirsute or pilose ribs. Calea × crassa differs from C. ilienii by the concolorous leaves (vs. discolorous leaves), involucre 6-seriate (vs. 5-seriate), foliaceous base of the second series of phyllaries (vs. scarious base), disc florets 8.9–11.3 mm long (vs. 6.1–8.9 mm ), tube 1.9–3.6 mm long (vs. 1.1–1.9 mm ), lobes 1.1–1.7 mm long (vs. 0.6–1.2 mm ), cypselae 2.5–3.8 mm long ( 2–2.5 mm ), and bigger scales 1.8–1.9 mm long, when present (vs. 1.3 mm , when present). Calea × crassa can be distinguished from C. monocephala by the glandular-punctate leaves along the stems, except the apical ones (vs. eglandular in all leaves), third series of phyllaries with acuminate apex (vs. apiculate, sometimes rounded), sixth series of phyllaries narrow oblong (vs. oblanceolate, when present), ray florets 2-lobulate, often rounded, sometimes 4-lobulate apex (vs. 3-lobulate), and bigger scales 1.8–1.9 mm long, when present (vs. 2.1–2.2 mm , when present). The nothospecies diverges from the two parental species by different characters of the phyllaries series, such as the panduriform, sometimes broadly oblong second series (vs. ovate or narrow ovate to oblong or orbiculate), entirely foliaceous, sometimes foliaceous with scarious base third series (vs. scarious), incised margins of the fourth series (vs. entire), and obtuse apex of the fifth series (vs. rounded). These traits highlight the importance of detailing the morphological characterization of phyllaries in Calea , because they have been demonstrated as reliable morphological markers to support the circumscription of other recently described species such as C. funkiana Bueno & Heiden (2021) , C. pruskiana Bueno & Heiden (2022a) and C. subintegerrima Bueno & Heiden (2022a) . The features of the hybrid C. × crassa intermediate and overlapping with those of its parents are described in Table 2 . The PCA ( Fig. 5 ) supports that, despite the characters overlapping with the two parental species, there is a statistical support that corroborates that these are three different taxa: two parental species and one nothospecies. TABLE 2. Intermediate or overlapped morphological features of Calea × crassa and its parents: C. ilienii and C. monocephala ( Neurolaeneae , Asteraceae ).

Calea ilienii	Calea × crassa	Calea monocephala
Abaxial surface indumentum in leaves along the stems Abaxial surface indumentum in apical leaves	Hispid or sparsely hispid, rarely glabrous to sparsely hispidulous	Densely hispid or hispid, rarely sparsely hispid Glabrous or sparsely hispid	Glabrous, sometimes sparsely puberulent or sparsely hispid to glabrous
Glands in abaxial surface in leaves along the stems	Densely glandular-punctate, sometimes	Glandular-punctate
glandular-punctate, rarely glabrous with	Absent
Glands in abaxial surface in apical	apex glandular-punctate	Sparsely glandular-punctate or
leaves	absent
Peduncle indumentum	Hispid to densely hispid, sometimes hispidulous to hispid	Densely hispid, rarely sparsely hispid to hispid	Sparsely hispid to hispid, rarely densely hispid
Indumentum of first series of phyllaries	Densely hispid to hispid, often hispid to sparsely hispid	Glabrous or densely hispid to hispidulous or hispid to sparsely hispid	Glabrous, rarely glabrous with base sparsely hispid
Glands in first series of phyllaries	Glandular-punctate to densely glandular- punctate, sometimes just the apex glandular-punctate, rarely absent	Glandular-punctate, rarely absent	Absent
Indumentum of second series of phyllaries	Densely hispid, often hispid, sometimes sparsely hispid	Sparsely hispid to hispidulous, sometimes glabrous	Glabrous
Glands in second series of phyllaries	Densely hispid, often hispid, sometimes sparsely hispid	Sparsely hispid to hispidulous, sometimes glabrous	Absent
Corolla of the ray florets (length)	13.7–18.1 mm	14.3–22.9 mm	14.6–24.2 mm
Abaxial surface indumentum of ray florets	Densely glandular-punctate, often glandular-punctate	Glandular-punctate, often sparsely glandular-punctate, rarely glabrous	Sparsely glandular-punctate, rarely glandular-punctate
Disc florets (length)	7.6–8.9 mm	8.9–11.3 mm	9–11 mm
Cypselae (length)	13.7–18.1 mm	2.5–3.0 (–3.8) mm	3.2–3.6 mm
Pappus scales (length)	0.6–1.6 mm	1.3–1.7 (–2) mm	1.3–2.3 mm
Bigger pappus scale, when present (length)	1–1.5 mm	1.8–1.9 mm	2.1–2.3 mm

FIGURE 5. Prediction occurrence map of suitable ecological niche modelling for Calea × crassa based on overlapping occurrence areas of C. ilienii and C. monocephala ( Neurolaeneae , Asteraceae ). Paratypes : — BRAZIL . Paraná , Balsa Nova, Tamanduá, Ponte dos Arcos , 8 March 2012 , E . Barbosa , & J. Cordeiro 3380 (MBM, UB). Additional specimens examined :— Calea ilienii — BRAZIL . Paraná , Balsa Nova , Serrinha , S „o Luiz do Purun „, 6 April 1914 , G . Jonsson 141a ( G ); Lapa , Hotel Fazenda Colônia Municipal , 2 March 2002 , J . Cordeiro et al. 1967 ( HUCS ) . Santa Catarina , Campo Alegre , Serra do Quiriri , 11 February 2001 , O . S . Ribas & J . M . Silva 3246 ( HAS ); Garuva , campos do Quiriri , 15 February 2020 , T . F . Santos 347 ( ICN ); Lages , Morro Pinheiro Seco , 3 km East of Lajes , 15 January 1957 , L . B . Smith & R . Reitz 10053 ( RB ) . Calea monocephala — BRAZIL . Paraná : Bocaiúva do Sul , Campina dos Tavares , 28 January 1970 , G . Hatschbach et al. 23442 ( MBM ); Campo Largo , S „o Luiz do Purun „, 10 January 1977 , L . T . Dombrowski et al. 6959 ( MBM ); Palmeira , Córrego da Anta , 2 January 1975 , G . Hatschbach et al. 35885 ( MBM ); Piraquara , 20 March 1979 , L . T . Dombrowski et al. 10329 ( MBM ); Ponta Grossa , estrada para Castro , 20 December 1952 , G . Hatschbach 2907 ( MBM ) .