Calea × crassa (Neurolaeneae: Asteraceae), a new nothospecies from Paraná state, Brazil Bueno, Vinicius R. Cassol, Ana P. V. Leroy, Carolina J. Bueno, Marcelo L. Heiden, Gustavo Phytotaxa 2023 2023-10-03 618 2 133 148 V.R.Bueno & G.Heiden V. R. Bueno & G. Heiden 2023 [136,582,1556,1583] Magnoliopsida Asteraceae Calea true Plantae Asterales 3 136 Tracheophyta species crassa sp. nov.   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.   Ristow8265( holotype: MBM!, isotypes: FUEL!, FURB!, HUCS!, IRAI, digital image!, RB!).   Calea× crassais differentiated from the parental species  C. ilieniiand  C. monocephalaby 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 mtall. Stems cylindrical, striate, sparsely hispid or hispid to densely hispid, eglandular, castaneous, internodes 10–60.8 (–89.7) mm long. Leavesdecussate, 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 mmlong, 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. Capitulumsolitary or capitulescence cymose, slightly branched, axis 0–2.17 cmlong, branching to 2 ndramifications, 1–3 (–4) secondary stems; peduncle 0.58–6.3 cmlong, 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. Phyllaries6-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. Receptacleslightly convex, holopaleaceous; paleae 8.1–10.4 mmlong, longer than disc floret corolla, narrow-oblanceolate or narrow-elliptic, apex long acuminate, often acuminate, conduplicate, light yellow. Ray florets12–20, pistillate, corolla 14.3–22.9 mmlong, liguliform, tube 3–4.3 mmlong, 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 mmlong, yellow. Disc florets50–80, bisexual, corolla (5.2–) 6.2–9 mmlong, 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 mmlong, light yellow, apical anther appendages ovate; style arms 1–1.5 mmlong, linear, yellow. Cypselae2.5–3 (–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; pappusmonotypic, 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 mmlong (when present), oblong, sometimes oblanceolate, apex obtuse, sometimes rounded, margins entire, erose in apex.   Distribution, habitat and predicted area of occurrence:—  Calea × crassais 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. ilieniihas not been collected in Ponta Grossa ( Fig. 3). However, the municipality has the same typeof 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 × crassaoccurs 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. × crassapoints 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 × crassacould be considered Critically Endangered: B1, B2; C2i; D ( IUCN 2022). These different criteria are applicable due the AOO shorter than 10 km² 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. × crassawas 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. × crassais considered Critically Endangered (CE).   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 ilieniiand  C. monocephala( Neurolaeneae, Asteraceae).  Taxonomic comments:—The specimens of  Calea× crassastudied 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 × crassashares 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 × crassadiffers from  C. ilieniiby 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 mmlong (vs. 6.1–8.9 mm), tube 1.9–3.6 mmlong (vs. 1.1–1.9 mm), lobes 1.1–1.7 mmlong (vs. 0.6–1.2 mm), cypselae 2.5–3.8 mmlong ( 2–2.5 mm), and bigger scales 1.8–1.9 mmlong, when present (vs. 1.3 mm, when present).   Calea × crassacan be distinguished from  C. monocephalaby 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 mmlong, 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. × crassaintermediate 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 × crassaand its parents:  C. ilieniiand 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× crassabased on overlapping occurrence areas of  C. ilieniiand  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 .   Jonsson141a( G); Lapa, Hotel Fazenda Colônia Municipal,  2 March 2002,  J .   Cordeiroet al.1967 ( HUCS).  Santa Catarina, Campo Alegre, Serra do Quiriri,  11 February 2001,  O .  S.   Ribas&  J .  M.   Silva3246( HAS); Garuva, campos do Quiriri,  15 February 2020,  T .  F.   Santos347( ICN); Lages, Morro Pinheiro Seco,  3 kmEast of Lajes,  15 January 1957,  L .  B.   Smith&  R .   Reitz10053( RB).   Calea monocephala—  BRAZIL. Paraná: Bocaiúva do Sul, Campinados Tavares,  28 January 1970,  G .   Hatschbachet al. 23442( MBM); Campo Largo, S „o Luiz do Purun„,  10 January 1977,  L. T .   Dombrowskiet al. 6959( MBM); Palmeira, Córrego da Anta,  2 January 1975,  G .   Hatschbachet al. 35885( MBM); Piraquara,  20 March 1979,  L. T .   Dombrowskiet al. 10329( MBM); Ponta Grossa, estrada para Castro,  20 December 1952,  G .   Hatschbach2907( MBM). 4420038303 [216,1332,1632,1654] 2012-03-19 J, M Brazil Parque Estadual de Vila Velha Ponta Grossa 3 136 1 Parana 4420038322 [1345,1448,1632,1653] C, B Brazil Silva 3 136 1 Parana 4420038317 [189,326,1668,1689] L, T Brazil Poliquesi 3 136 1 Parana 4420038311 [335,472,1668,1689] R Brazil Maranho 3 136 1 Parana 4420038314 [482,1386,1668,1690] MBM, FUEL, FURB, HUCS, IRAI, RB Brazil Ristow 3 136 1 Parana holotype 4420038301 [189,1269,1319,1346] 2012-03-08 E Ponte dos Arcos Brazil Parana 9 142 1 Parana paratype 4420038304 [770,1239,1391,1418] S Brazil Serrinha Balsa Nova 9 142 1 Parana 4420038312 1914-04-06 G Brazil Luiz do Purun 9 142 1 Parana 4420038315 [331,1209,1427,1454] 2002-03-02 G, J Lapa & Hotel Fazenda Colonia Municipal Brazil Jonsson 9 142 1 Parana 4420038310 HUCS Brazil Cordeiro 9 142 1 Parana 4420038305 [245,1044,1463,1490] 2001-02-11 O Brazil Serra do Quiriri Campo Alegre 9 142 1 Santa Catarina 4420038316 [1048,1066,1464,1490] 2001-02-11 S Brazil true Serra do Quiriri Campo Alegre 9 142 1 Santa Catarina 4420038320 [1078,1196,1463,1489] J Brazil Ribas 9 142 1 Santa Catarina 4420038325 [1200,1228,1464,1489] M Brazil true Ribas 9 142 1 Santa Catarina 4420038307 2020-02-15 HAS, T Brazil Silva campos do Quiriri Garuva 9 142 1 Santa Catarina 4420038321 [676,694,1500,1525] 2020-02-15 F Brazil Silva true campos do Quiriri Garuva 9 142 1 Santa Catarina 4420038309 1957-01-15 ICN, L Lages & Morro Pinheiro Seco Brazil Santos 9 142 1 Santa Catarina 4420038318 [358,378,1536,1561] 1957-01-15 B Lages & Morro Pinheiro Seco Brazil true Santos 9 142 1 Santa Catarina 4420038324 [389,508,1535,1561] R Brazil Smith 9 142 1 Santa Catarina 4420038302 [519,719,1535,1561] RB Brazil Reitz 9 142 1 Santa Catarina 4420038306 [442,1308,1571,1598] 1970-01-28 G Brazil Bocaiuva do Sul Tavares Campina 9 142 1 Parana 4420038323 MBM, S Campo Largo Brazil Hatschbach 9 142 1 Parana 4420038319 [578,1057,1607,1634] 1977-01-10 L, T Brazil Luiz do Purun 9 142 1 Parana 4420038308 1975-01-02 MBM, G Brazil Dombrowski Corrego da Anta Palmeira 9 142 1 Parana 4420038313 [692,1448,1643,1669] 1979-03-20 MBM, L, T Piraquara Brazil Hatschbach 9 142 1 Parana 4420038326 [136,1233,1679,1706] 1952-12-20 MBM, G Ponta Grossa & Castro Brazil Dombrowski 9 142 1 Parana 4420038327 MBM Brazil Hatschbach 9 142 1 Parana