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Al-Shehbaz, Ihsan A., German, Dmitry A. (2022): Cardamine pakistanica (Cardamineae; Brassicaceae), a new species from North Pakistan. Phytotaxa 574 (1): 118-120, DOI: 10.11646/phytotaxa.574.1.9, URL: http://dx.doi.org/10.11646/phytotaxa.574.1.9

Fouraeeae Al-Shehbaz, M. Koch, R. Karl, & D. A. German, trib. nov. Type genus: Fourraea Greuter & Burdet Herbs normally perennial, glaucous.Trichomes absent or simple, confined to margins of basal leaves.Leaves exclusively or nearly so entire, middle and upper cauline ones sessile, amplexicaul to strongly auriculate. Racemes ebracteate, elongated in fruit. Petals white, lavender, intense purple-violet. Ovules numerous. Fruits dehiscent, unsegmented, linear siliques, glabrous, latiseptate or nearly so, with distinct or obscure midvein. Seeds uniseriate; cotyledons accumbent. Species of the Fourraeeae can generally (but not exclusively) be distinguished from those of the Arabideae by being either glabrous or at most with simple trichomes confined to leaf margins and that their seeds are always uniseriate. By contrast, species of the Arabideae have most often one or more type of branched trichomes that are sometimes mixed with simple ones, and seeds are often biseriate or very rarely aseriate, but uniseriate seeds are known and common in Arabideae, too. Three species in two genera distributed in southern and central Europe and Morocco. Key to genera. 1a. Fruits with a prominent midvein; trichomes simple, confined to margin of basal leaves; petals white; Europe................. Fourraea 1b. Fruits with obscure midvein; trichomes absent; petals white, pink, or dark purple-violet; Morocco................................... Hurkaea, Published as part of Koch, Marcus A., Karl, Robert, Al-Shehbaz, Ihsan A. & German, Dmitry A., 2022, The new tribe Fourraeeae (Brassicaceae) and two Moroccan Arabis species transferred to the new genus Hurkaea, pp. 41-50 in Phytotaxa 543 (1) on pages 44-45, DOI: 10.11646/phytotaxa.543.1.4, http://zenodo.org/record/6424331

Fourraea Greuter & Burdet (Greuter & Raus 1984: 283) Type: Fourraea alpina (L.) Greuter & Burdet. = “ Caulopsis ” Fourreau (1868: 332), nom. inval. Herbs perennial, with elongated caudex, glaucous. Trichomes simple, confined only to petioles. Multicellular glands absent. Stems erect, simple, leafy, unarmed. Basal leaves long petiolate, ciliate along proximal half of petiole, rosulate; blade simple, entire, sometimes ciliate near base; cauline leaves sessile, amplexicaul to strongly auriculate at base, simple, entire. Racemes few to many flowered, ebracteate throughout, lax, elongated considerably in fruit; rachis straight; fruiting pedicels ascending-divaricate, slightly curved or straight, persistent. Sepals elliptic-oblong, free, deciduous, erect, equal, base of lateral pair not saccate. Petals white, erect to ascending, longer than sepals; blade narrowly oblanceolate, apex rounded; claw shorter than sepals, glabrous, unappendaged, entire. Stamens 6, slightly exserted, erect, tetradynamous; filaments wingless, unappendaged, glabrous, free; anthers oblong, apex obtuse. Nectar glands 4, distinct or confluent; median nectaries ovoid, smaller than laterals; lateral nectaries ringlike. Ovules (30–)36–52 per ovary; placentation parietal. Fruit dehiscent, capsular siliques, linear, erect to ascending, subtetragonal-latiseptate, not inflated, unsegmented; valves thin or thick leathery, with a prominent midvein, glabrous, not keeled, slightly torulose, wingless, unappendaged; gynophore obsolete or to 0.5 mm long; replum rounded, visible; septum complete, membranous, veinless; style absent, obsolete, or 0.7–1 mm long, persistent; stigma capitate, entire, unappendaged. Seeds uniseriate, wingless, oblong-ovoid, plump; seed coat mucilaginous when wetted; cotyledons obliquely accumbent. A monospecific European genus. Fourraea alpina (L.) Greuter & Burdet (Greuter & Raus 1984: 283). ≡ Brassica alpina Linnaeus (1767: 95). For a complete synonymy, see BrassiBase (https://brassibase.cos.uni-heidelberg.de), POWO (https://powo. science.kew.org/), WCVP (https://wcvp.science.kew.org/) [the latter two under Turritis brassica Leers (1775: 147)], and for species descriptions and distributions see Akeroyd (1993) and Jalas & Suominen (1994), respectively, both under Arabis brassica (Leers) Rauschert (1973: 648), as well as Talavera (1993) under A. pauciflora and Euro+Med PlantBase (https://europlusmed.org/). Distribution: Andorra, Austria, Belgium, Croatia, Czech Republic, France, Germany, Italy, Liechtenstein, Luxembourg, Serbia, Slovakia, Slovenia, Spain, Switzerland., Published as part of Koch, Marcus A., Karl, Robert, Al-Shehbaz, Ihsan A. & German, Dmitry A., 2022, The new tribe Fourraeeae (Brassicaceae) and two Moroccan Arabis species transferred to the new genus Hurkaea, pp. 41-50 in Phytotaxa 543 (1) on page 45, DOI: 10.11646/phytotaxa.543.1.4, http://zenodo.org/record/6424331, {"references":["Greuter, W. & Raus, T. (Eds.) (1984) Med-Checklist notulae, 8. Willdenowia 13 (2): 283 - 288.","Fourreau, J. (1868) Catalogue des plantes qui croissent spontanement le long du Rhone. Annales de la Societe Linneenne de Lyon ser. 2, 16: 301 - 404. https: // doi. org / 10.5962 / bhl. title. 12050","Linnaeus, C. (1767) Mantissa plantarum. Generum editionis VI et Specierum editionis II. Salvius, Holmiae [Stockholm], 144 pp.","Leers, J. D. (1775) Flora herbornensis. Hebornae Nassoviorum (sumptibus auctoris), [Herborn], [XXXII] + LIX + 292 pp. + 16 pl.","Akeroyd, J. R. (1993) Arabis. In: Tutin, T. G., Burges, N. A., Chater, A. O., Edmondson, J. R., Heywood, V. H., Moore, D. M., Valentine, D. H., Walters, S. M. & Webb, D. A. (Eds.) Flora Europaea 1. Psilotaceae to Platanaceae. Ed. 2. Cambridge University Press, Cambridge, pp. 352 - 356.","Jalas, J. & Suominen J. (Eds.) (1994) Atlas Florae Europaeae. 10. Cruciferae (Sisymbrium to Aubrieta). Committee for Mapping the Flora of Europe and Societas Biologica Fennica Vanamo, Helsinki, 224 pp.","Rauschert, S. (1973) Zur Nomenklatur der Farn- und Blutenpflanzen Deutschlands (III). Feddes Repertorium 83 (9 - 10): 645 - 662. https: // doi. org / 10.1002 / fedr. 19730830903","Talavera, S. (1993) Arabis. In: Castroviejo, S., Aedo, C., Gomez Campo, C., Lainz, M., Montserrat, P., Morales, R., Munoz Garmendia, F., Nieto Feliner, G., Rico, E., Talavera, S. & Villar, L. (Eds.) Flora Iberica. Plantas vasculares de la Peninsula Iberica e Islas Baleares 4. Real Jardin Botanico, Madrid, pp. 135 - 163."]}

Hurkaea conringioides (Ball) Al-Shehbaz, M. Koch, R. Karl, & D.A. German, comb. nov. ≡ Arabis conringioides Ball (1873: 297). Type indication: “ Hab. in regione superiori Atlantis Majoris---in jugo Tagherot! A 3000 m ad 3500 m; in monte Djebel Tezah! A 2100 m ad 2800 m ”. Lectotype (designated here):— MOROCCO: ‘Ex regione superiori Atlantis Majoris in cacumine Djebel Tezah, alt. 1700–3500 m, 22 May 1871, J. Ball s.n. (K-000239698!; isolectotype: G-00023347!). Note. Ball (1873) cited two syntypes, and the one from Tagherot (K-000230696, K-000230698, G-00023348) was immature and consists of rosette plants. By contrast, the one designated above as lectotype consists of complete mature plants of which some are in flower and others are in fruit. Two subspecies endemic to the High (Grand) Atlas of Morocco. Key to subspecies 1a. Petals pink, 6–9 × 3–5 mm; style in fruit distinct, 0.5–1.5 mm long; central and western Atlas Mts................. subsp. conringioides 1b. Petals white to light pink, 4–5.5(–7) × 1.3–2 mm; style in fruit obsolete; eastern Atlas Mts................................... subsp. humbertii

Hurkaea Al-Shehbaz, M. Koch, R. Karl, & D.A. German, gen. nov. [Named in honor of Herbert Hurka (5 May 1940 –, Greiz / Thuringia), since 1982 Professor for Botany, Systematics and Evolution at Osnabrück University and retired in 2005, in appreciation of his important contributions in Brassicaceae systematics and phylogenetics paving the way for thorough taxonomic studies]. Type species: Hurkaea conringioides (Ball) Al-Shehbaz, M. Koch, R. Karl, & D.A. German. Herbs normally perennial (sometimes flowering in first year), cespitose or not, with slender or compact caudex, glabrous throughout, glaucous. Trichomes absent. Multicellular glands absent. Stems erect, simple or few to several branched at base. Basal leaves petiolate, rosulate or not, simple, entire (rarely in H. conringioides with 1 tooth or small lobe on each side); cauline leaves sessile, amplexicaul to strongly auriculate at base, entire. Racemes many flowered, ebracteate throughout, initially corymbose in flower, slightly to considerably elongated and lax or somewhat dense in fruit; rachis straight; fruiting pedicels ascending to widely spreading or descending, slender, straight, persistent. Sepals narrowly oblong, free, deciduous, suberect, equal, base of lateral pair not saccate; petals white, pink, or dark purpleviolet, erect at base with flaring blade, distinctly longer than sepals; blade oblanceolate, apex obtuse; claw only slightly differentiated from blade, subequaling sepals, glabrous, unappendaged, entire; stamens 6, slightly exserted, erect, tetradynamous; filaments wingless, unappendaged, glabrous, free; anthers linear to oblong; nectar glands confluent, subtending bases of median stamens, surrounding those of lateral stamens; ovules 30–90 per ovary; placentation parietal. Fruit dehiscent, capsular siliques, narrowly linear, latiseptate, not inflated, unsegmented; valves papery, with obscure midvein, glabrous, smooth, unappendaged; gynophore absent; replum rounded, visible; septum complete, membranous, veinless; style obsolete or distinct and 0.5–1.5 mm long; stigma capitate, entire, unappendaged. Seeds uniseriate, wingless or apically narrow winged, oblong, flattened; cotyledons accumbent. Two species, endemic to Morocco. Key to the species 1a. Plants cespitose; stems 10–30(–40) cm tall; petals white or pink; fruiting pedicels (0.5–) 0.7–1.3 cm long, erect to ascending, subappressed or not; fruits suberect, subappressed, or ascending, 4–6.7 cm long.................................................... H. conringioides 1b. Plants not cespitose; stems 50–130 cm tall; petals dark purple-violet; fruiting pedicels (1.2–) 2.5–4.5 cm long, spreading or divaricate-ascending; fruits widely spreading to descending, (3.5–) 5–13 cm long.............................................................. H. josiae, Published as part of Koch, Marcus A., Karl, Robert, Al-Shehbaz, Ihsan A. & German, Dmitry A., 2022, The new tribe Fourraeeae (Brassicaceae) and two Moroccan Arabis species transferred to the new genus Hurkaea, pp. 41-50 in Phytotaxa 543 (1) on pages 45-46, DOI: 10.11646/phytotaxa.543.1.4, http://zenodo.org/record/6424331

Hurkaea conringioides subsp. conringioides = Arabis werneri Emb. & Maire (Maire 1933: 196). ≡ A. conringioides var. werneri (Emb. & Maire) Galland & Favarger (1985: 233). Type indication: “Hab. in fissuris rupium vulcanicarum Anti-Atlantis: in ditionis Siroua monte Amezdour, ad alt. 2600–2700 m ”. Lectotype (designated here):— MOROCCO: Siroua, Au Tizi-n-Ougdar, Mt. Amezdour (as Amerouz), 2600 m, 30 June 1932, L. Emberger s.n. (MPU-002817; isolectotype: MPU-002818). Hurkaea conringioides subsp. humbertii (Quézel) Al-Shehbaz, M. Koch & R. Karl, comb. nov. ≡ Arabis humbertii Quézel (1953: 198). ≡ Arabis conringioides subsp. humbertii (Quézel) Dobignard & Jordan (1987: 79). Type indication: “Hab. in atlante majore orientale, loco vocato Imi N’Tamd.” Type:— MOROCCO: J. Ayachi, Imi-N-Tamd, June-July 1952, C. Guinet, Ch. Souvage et R. de Villmorin 792 (RAB-000465). Note. Galland & Favarger (1985) recognized three varieties in Arabis conringioides, including var. conringioides, var. werneri, and var. aphanostyla Maire. The last was invalidly published in Maire (1967). By contrast, Dobignard & Jordan (1987) recognized the two subspecies listed here, though they accepted var. aphanostyla, which Dobignard et al. (2011) subsequently reduced to synonymy of subsp. humbertii. Hurkaea josiae (Jahand. & Maire) Al-Shehbaz, M. Koch, R. Karl, & D.A. German, comb. nov. ≡ Arabis josiae Jahandiez & Maire (1923: 66). Type indication: “Hab. in silvis (quercetis et cedretis) Atlantis Medii maroccani supra oppidum Azrou, ubi maio floret – Typus in Herbario Universitatis Algeriensis. … Cette plante a été ensuite récoltée en fleurs, le 16 mai 1921, per Jahandiez, dans le cratère du Djebel Hebbri, entre Azrou et Timhadit, dans les clairières des cédraires, sur basalte vers 1900 m. Elle a été ensuite retrouvée par Maire dans les forêts de Quercus ilex au dessus d’Azrou, sur calcaire jurassique vers 1400 m, en fruits plus que mûrs, le 25 juillet 1921 ”. Type: Morocco, Moyen Atlas, above town Azrou, 1921, É. Jahandiez &/or R. Maire s.n. Note. Although the North African herbarium of the University of Algeria (AL) is listed in Index Herbariorum online (sweetgum.nybg.org/science/ih/), no address or correspondents are given, and inquiry about the type collection of Arabis josiae was not possible. The original protologue (Jahandiez & Maire 1923) lists two gatherings, and relevant syntypes are available in MPU (with printed labels from AL “Université d’Algiere Herbier del’Afrique du Nord”) and AIX. The collections by Jahandiez were made on 16 May 1921 (MPU-000159, AIX-000066) at 1900 m from Cedrus forests, and they are immature plants in early stages of flowering. However, Maire’s collections (MPU-000157, MPU-000158) were made on 25 July 1921 at 1400 m from Quercus forests. They are with fully mature plants. Because the original description includes flowering and fruiting material, and because the fruits are essential for the characterization of the genus Hurkaea, a designation of the specimen with mature fruits at AL as the lectotype would be preferable, and its duplicate at MPU would then be the isolectotype. = Arabis leptopoda Pau & Font Quer (Font Quer 1928: no 236, in sched.). ≡ A. josiae var. leptopoda (Pau & Font Quer) Pau & Font Quer (Font Quer 1929: no 139, in sched.). Type indication: “Hab. in saxosis, lacisque silvaticis montis Iguermalez (Atlante rhiphaeo), 1950 m. alt.; 7 junii”. Lectotype (designated by Nualart et al. 2021: 14):— MOROCCO: In saxosis, lacisque silvaticis montis Iguermalez (Atlante rhiphaeo), 1950 m, 7 June 1928, Font Quer 236-1928 (MA-48513; isolectotypes BC-2978, BCN-17301, BM-000583666, GDA-28946, MPU-006331, MPU 006332)., Published as part of Koch, Marcus A., Karl, Robert, Al-Shehbaz, Ihsan A. & German, Dmitry A., 2022, The new tribe Fourraeeae (Brassicaceae) and two Moroccan Arabis species transferred to the new genus Hurkaea, pp. 41-50 in Phytotaxa 543 (1) on pages 46-47, DOI: 10.11646/phytotaxa.543.1.4, http://zenodo.org/record/6424331, {"references":["Maire, R. (1933) Contribution a l'etude de la flore de l'Afrique du Nord. Bulletin de la Societe d'histoire naturelle de l'Afrique du Nord 24 (7): 194 - 232.","Galland, N. & Favarger, C. (1985) Notes de taxonomie et de nomenclature sur la flore orophile du Maroc. Candollea 40 (1): 231 - 235. https: // doi. org / 10.5169 / seals- 879783","Quezel, P. (1953) Contribution a la flore de l'Afrique du Nord. Deuxieme contribution a la flore du Haut Atlas Marocain. Bulletin de la Societe d'histoire naturelle de l'Afrique du Nord 44 (5 - 6): 196 - 202.","Dobignard, A. & Jordan, D. (1987) Nouvelles observations sur la flore du Maroc. 1. Contribution a l'etude de la flore du Haut-Atlas. Saussurea 18: 67 - 104.","Maire, R. (1967) Flore de l'Afrique du Nord 13. Paul Lechevalier, Paris, 365 pp.","Dobignard, A., Chatelain, C., Fischer, M., Orso, J. & Jeanmonod, D. (2011) Index synonymique de la flore d'Afrique du Nord 3, Dicotyledoneae: Balsaminaceae-Euphorbiaceae. Conservatoire et Jardin botaniques de la Ville de Geneve, Geneve, 449 pp.","Jahandiez, E. & Maire, R. (1923) Plantae maroccanae novae. Bulletin de la Societe d'histoire naturelle de l'Afrique du Nord 14 (2): 65 - 73.","Font Quer, P. (1928) Iter Maroccanum 1927. Exsiccata schedula. Natural Science Museum, Barcelona, 739 num.","Font Quer, P. (1929) Iter Maroccanum 1928. Exsiccata schedula. Natural Science Museum, Barcelona, 438 num.","Nualart, N., Soriano, I., Perez Prieto, D. & Ibanez, N. (2021) Catalogue and typification of the Moroccan taxa published by Carlos Pau. Phytotaxa 519 (1): 1 - 94. https: // doi. org / 10.11646 / phytotaxa. 519.1.1"]}

The mustard family (Brassicaceae) is a scientifically and economically important family, containing the model plant Arabidopsis thaliana and numerous crop species that feed billions worldwide. Despite its relevance, most published family phylogenies are incompletely sampled, generally contain massive polytomies, and/or show incongruent topologies between datasets. Here, we present the most complete Brassicaceae genus-level family phylogenies to date (Brassicaceae Tree of Life, or BrassiToL) based on nuclear (>1,000 genes, almost all 349 genera and 53 tribes) and plastome (60 genes, 79% of the genera, all tribes) data. We found cytonuclear discordance between nuclear and plastome-derived phylogenies, which is likely a result of rampant hybridisation among closely and more distantly related species, and highlight rogue taxa. To evaluate the impact of this rampant hybridisation on the nuclear phylogeny reconstruction, we performed four different sampling routines that increasingly removed variable data and likely paralogs. Our resulting cleaned subset of 297 nuclear genes revealed high support for the tribes, while support for the main lineages remained relatively low. Calibration based on the 20 most clock-like nuclear genes suggests a late Eocene to late Oligocene ‘icehouse origin’ of the family. Finally, we propose five new or re-established tribes, including the recognition of Arabidopsideae, a monotypic tribe to accommodate Arabidopsis. With a worldwide community of thousands of researchers working on this family, our new, densely sampled family phylogeny will be an indispensable tool to further highlight Brassicaceae as an excellent model family for studies on biodiversity and plant biology., bioRxiv