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Reproductive successThe production of urinary pheromones correlated with male but not female reproductive success (RS; defined in “Materials and methods” section). The most important predictors of male RS were total urinary protein concentration (75%) and social status (69%; Table 1; based on conditional model average sum of weights). The relative importance of age, creatinine, and mass ranged from 23 to 39%; PC ratio (protein:creatinine concentration) was excluded from the model due to collinearity (VIF = 6.97). Total urinary protein concentration during the enclosure phase was positively correlated with RS for males (Spearman R = 0.52, p = 0.01; Fig. 1a), but not females (Fig. 1b). This correlation is explained by the low protein concentration in the urine of non-reproductive males, as it is no longer significant after removing these males from the analysis (R = 0.12, p = 0.62; Supplementary Fig. S2). The median total urinary protein concentration was 5512 µg mL−1 and 5028 µg mL−1 for reproductive and non-reproductive males, respectively (Wilcoxon rank sum test W = 5, p  More

Generic nameRavenala Adans.1 (1763: 67). (equiv) Urania Schreb.22 (1789: 212). –Ravenala Scop.23, nom. illeg. (1777: 96) as “Ravenalla Adans”.Type species Ravenala madagascariensis Sonn.24.Note: Dorr & Parkinson25 proposed to conserve the spelling Ravenala Scop. (and correct Scopoli’s original orthography “Ravenalla”) against Ravenala Adans. on the basis that Adanson’s generic names (using a uninominal nomenclature for species) were invalid. Brummitt26 rejected this proposal and considered that Adanson’s generic names were valid27 and thus that there was no need to use Scopoli’s Ravenala (Ravenalla). Moreover, the exact wording in Scopoli23 (1777: 96) is “Ravenalla Adans.”, citing Adanson explicitly, but with an incorrect spelling for the generic name (the double “l”).Typification and emended descriptionRavenala madagascariensis Sonn. (1782: 2[ed. qto.]: 223, tt. 124–126).(equiv) Ravenala madagascariensis J.F.Gmel.28 (1791: 567). (equiv) Urania madagascariensis (Sonn.) Schreb. ex Forsyth f.29 (1794: 212). (equiv) Heliconia ravenala Willemet30 (1796: 22). (equiv) Urania speciosa Willdenow31 (1799: 7). (equiv) Urania ravenalia (Willemet) A.Rich.32 (1831: 19). –Ravenala madagascariensis Adans.1 (1763: 597), nomen invalid., appearing on page 597, abbreviated in the final index of Adanson’s book as “Ravenala madag. 67”, which can also be construed as referring to Madagascar as a locality.Type Lectotype, here designated: The plate numbered 126, representing the typical lax mature infructescence, in Sonnerat24 (1782: plate 126). Epitype, here designated: MADAGASCAR (bullet) Fort-Dauphin, Forêt de Manantantely, [24°58′ 59.988″S, 46°55′0.012″E, calc. from label], 60–300 m elev., 15 September 1928, H. Humbert 5730 (Epitype: MNHN-P-P02234599!, Isoepitypes: MNHN-P-P02234602!, MNHN-P-P02234604!, MNHN-P-P02234605!).Additional specimen examined: MADAGASCAR (bullet) Toamasina: Foulpointe, Analalava Forest, plant growing close to the main forest station, 17°42.3′S, 49°27.38′E, 50 m elev., 20 March 2016, T.Haevermans, M. Vorontsova, S. Dransfield & J. Razanatsoa 821 (TAN!, P!, K !) (bullet) X. Aubriot et al. 45 (P00696168!, P00696167!, P00685124!, TAN!) (bullet) Along Route #5 from Fenerive to Maroantsetra, disturbed areas along road, 100 m elev., 28 February 1975, T. B. Croat 32540 (L-WAG.1111446!, L-WAG.1111447!, MO-358490!, MO-358491!, MO-358523!) (bullet) Toalagnaro, Ebakika, District de Fort-Dauphin, 12 July 1932, R. Decary 10107 (P02234596!) (bullet) Vondrozo (commune de Farafangana), 16 September 1926, R. Decary 5428 (P02234588!, P02234591!, P02234592!) (bullet) 2 km E of Ranomafana towards Brickaville, 18.965° S, 48.8564° E, 4 March 1992, J. Kress et al. 92-3412 (US00424302!, US00424299!, US00424300!, US00424301!, US00424303!) (bullet) 18 km E of Ranomafana, 25 km W of Brickaville, 18.9453° S, 48.9664° E, 4 March 1992, J. Kress et al. 92-3414 (US00424312!, US00424309!, US00424310!, US00424311!, US00424313!). MAURITIUS (bullet) Isle de France, s.dat., Commerson s.n. (P02234587!, P-JU!, P-LAM!).Identity of Ravenala madagascariensis Sonn. —Figs. 2d, 3d, 4d, 5d— In the absence of a specimen undoubtedly collected or seen by Sonnerat (Commerson’s specimens, collected in Mauritius and preserved in both Jussieu’s and Lamarck’s herbaria at the Paris herbarium (P-JU and P-LAM), might actually be part of original material), we decided to lectotypify from plates 124, 125 and 126 of the protologue in Sonnerat’s valid publication24 of the species. On page 225, Sonnerat24 mentions that the plant originated from Madagascar but was transported and established in Mauritius (known at the time as Isle de France) at the “Jardin des Pamplemousses”. We observed plants growing in this garden as well as naturalized plants occurring in the wild in Mauritius; all the plants we saw suckered and possessed the characteristic pointed conical fruits also observed in the Fort-Dauphin population. Sonnerat also specified that the original plant grew in marshy areas, which corresponds exactly to the coastal populations that can be found on the eastern coast of Madagascar (i.e. the “Horonorona” variant of Blanc et al.13). Plate 126 shows the typical mature infructescence of the species, with the space between bracts increasing before releasing the seeds (unlike other species of Ravenala). However, the “tree” pictured on plate 124 is a non-suckering plant, which in our opinion can be explained as artistic license on the part of the illustrator, as all the plants observed in Mauritius consistently sucker, like the plants growing in the south-eastern marshy areas. We also decided to designate an epitype with a documented locality in Madagascar (the material in P-JU and P-LAM does not bear a precise indication of locality) to fix the application of the name R. madagascariensis to the populations occurring in the marshy areas surrounding Fort-Dauphin, where only one morphotype is known.Figure 3Comparison of petiole bases. (a) R. agatheae. (b) R. blancii. (c) R. grandis. (d) R. madagascariensis. (e) R. menahirana. (f) R. hladikorum. Photographs Thomas Haevermans©.Full size imageFigure 4Comparison of inflorescences. (a) R. agatheae. (b) R. blancii. (c) R. grandis. (d) R. madagascariensis. (e) R. menahirana. (f) R. hladikorum. Photographs Thomas Haevermans©.Full size imageFigure 5Species of Ravenala in their natural habitat. (a) R. agatheae. (b) R. blancii. (c) R. grandis. (d) R. madagascariensis. (e) R. menahirana. (f) R. hladikorum. Photographs Thomas Haevermans©.Full size imageEmended description Plants suckering, 6–12 meters tall (adult), trunk circumference (d.b.h.) 20–30 cm, juvenile and adult laminae distributed in a perfect fan, 14–25 leaves simultaneously alive on the adult plant, 1–3 leaves between inflorescences. Leaves adult petiole 380–440 cm long, greenish-yellow, slightly waxy, sheath margin undeveloped to moderately developed (0–9 mm), entire, not drying, slightly splitting when aged (Fig. 3d), petiole/lamina ratio 1.9–(2.2)–2.3, adult lamina (200 times 100) cm, light green, juvenile lamina base non-decurrent. Inflorescences 4–6 live lateral inflorescences at a time, (100 times 100) cm (peduncle excluded), 8–16 bracts per inflorescence, bracts 200–(450 times 50)–100 mm, with some wax to very waxy, margin uniformly green (Fig. 4d), cincinnii of ca. 10 flowers per bract, flowering sequentially, bracteoles without a colored stripe. Flowers 240–280 mm long (ovary included), inferior ovary 40–50 mm long, perianth yellowish, sepals narrowly triangular 240–250 (times 10)–12 mm, sheathing (fused) petals narrowly triangular 220–230(times)ca. 10 mm, free petal acicular 180–190 (times 5) mm, slightly smaller than the remaining perianth with mean free petal/mean fused petal length ratio = 0.8, petal blotches absent, stamens (roughly) the same size as the perianth, 200–210 mm long, style 200–230 mm long, stigma 15–20 mm long, oblong ovoid with a basal constriction. Infructescences lax (bract bases not imbricate at maturity), stiff and coriaceous persisting bracts, old infructescences deciduous, 4–8 fruits per bract. Fruits 70–120 (times 30)–35 mm, trilocular septifragal capsule, apices conical (Fig. 2d), seeds 6–(8.5)–(11 times 5)–(6.4)–8 mm, shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.Ecology Ravenala madagascariensis is a low-altitude species restricted to swampy areas of the eastern coast of Madagascar. Populations outside of Madagascar on nearby islands are reputedly non-indigenous24.Preliminary IUCN assessments We propose a Least Concern status for R. madagascariensis, having an E.O.O ( > 20,000) km2 and an A.O.O. ( > 2,000) km2 (criterion B)33.Note This emended description for R. madagascariensis was drawn up from our own observations and collections, and was made comparable point by point to the descriptions of the five new species presented below, along with a dichotomous identification key to all six species.New species descriptions
Ravenala agatheae Haev. & Razanats. sp. nov.—Figs. 2a, 3a, 4a, 5a, 6
Type MADAGASCAR (bullet) Antsiranana: Ambanja District, along R.N.6 road to Ankaramibe, 13°45′54.8″S, 48°21′27.7″E, 30 m elev., on degraded lateritic slopes, 28 October 2018, T. Haevermans, A. Haevermans & J. Razanatsoa 830 (Holotype: TAN!, Isotypes: K!, MO!, P!).Figure 6Ravenala agatheae. (a) young infructescence. (b) adult plant habit showing the suckers at the base and the persistent petioles and old infructescences. (c) fruit with a conical apex. (d) infructescence with remains of dried flowers and dried bracts. (e) style apex. (f) inflorescence with open flowers. (g) open flower. Ink drawings on (75 , upmu) polyester tracing paper by Agathe Haevermans© from specimen Haevermans et al. 830, and observations in-situ.Full size imageParatypes MADAGASCAR (bullet) Antsiranana: 57–58 km N of Ambanja, 13°22′59.9″S, 48°48′E, 22 May 1974, A.H. Gentry 11878 (L-WAG.1111448!, L-WAG.1111449!, MO-358489!, TAN) (bullet) Ampasindava, forêts d’Ambilanivy et Rangoty, 13°48′36″S, 48°10′48″E, 29 November 2007, L. Nusbaumer 2658 (G334213/1!, MO!, TAN) (bullet) Mahajanga: Morafenobe, Beravy, 15 km from Beravy, near the road from Orombato to Beravy, 18°3′50″S, 44°31′46″E, 09 June 2016, F. Rakotonasolo et al. 2772 (K, P00782931!, TAN).Diagnosis Similar to Ravenala madagascariensis but differs in its dark green narrower laminae, tricolor petioles with very developed dryish petiole sheath margins, very waxy petioles, the persistence of older infructescences for several years, a purple stripe on the bract margin, longer bracts, a whitish perianth, brown blotches on its mature fused petals, the bracteole apex tinged with pink, an ovoid pointed stigma, dense infructescences, smaller inflorescences, the free petal much shorter than the fused petals, and an end of year flowering period.Distribution Plants restricted to Madagascar, growing in the north-western part of the island. We observed it growing from the southern part of the Diego Suarez area (on the hills along the road leading to Tsingy Rouge and the city of Sadjoavato) in the north to the western part of the Mahajanga province down to the Melaky region, with most observations around Ambanja34. We also observed that the species was cultivated on Nosy Be.Preliminary IUCN assessments We propose a Least Concern status for R. agatheae, having an E.O.O ( > 20,000) km2 and an A.O.O. ( > 2,000) km2 (criterion B)33.Ecology This species is adapted to seasonally dry and warm coastal habitats, growing on slopes at low elevations in north-western coastal areas of Madagascar, from Antsiranana (Diego-Suarez) down to the Melaky region in the Mahajanga province.Etymology This species is named after to the first author’s wife, Agathe Haevermans, a botanical illustrator at the Muséum National d’Histoire Naturelle, who helped discover this species in the field with the collecting team and who contributes greatly to botany by producing illustrations of new taxa from biodiversity hotspots such as Madagascar.Description Plants suckering, 6–10 meters tall (adult), trunk circumference (d.b.h.) 20–30 cm, juvenile and adult laminae distributed like a regular fan, 9–22 leaves simultaneously alive on the adult plant, 1–3 leaves between inflorescences. Leaves adult petiole 300–460 cm long, tricolor (dark green with a waxy white strip and red petiole sheath margin subsequently drying out, Fig. 3a), very waxy, sheath margin very developed (10 mm and more), entire, dryish-papyraceous and protruding at 90 degrees, petiole/lamina ratio 1.7–(1.95)–2.2, adult lamina 174–(210 times 72)–86 cm, dark green, juvenile lamina base non-decurrent. Inflorescences 4–6 live lateral inflorescences at a time, 70–(90 times 90)–100 cm (peduncle excluded), 10–14 bracts per inflorescence, bracts 450–500 (times 80)– 90 mm, with some waxiness (Fig. 4a), margin bearing a purple stripe, cincinnii of 8–10 flowers per bract, flowering sequentially, some pink tinge at the apex of bracteoles. Flowers 260–310 mm long (ovary included), inferior ovary 40–60 mm long, perianth whitish, sepals narrowly triangular 220–250(times)ca. 10 mm, sheathing (fused) petals narrowly triangular 200–(220times)ca. 10 mm, free petal acicular 130–(140 times 5) mm, much smaller than the remaining perianth with a mean free petal / mean fused petal length ratio = 0.6, petal blotches present, stamens (roughly) the same size as the perianth, 210–220 mm long, style 220 mm long, stigma 15 mm long, ovoid-pointed with basal constriction. Infructescences compact (bracts bases imbricate at all stages of maturity), stiff and coriaceous persisting bracts on mature infructescence, persistence of old infructescences, 4–10 fruits per bract. Fruits 90–110 (times) 30–45 mm, trilocular septifragal capsule, apices conical (Fig. 2a), seeds shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.
Ravenala blancii Haev., V. Jeannoda & A. Hladik sp. nov. —Figs. 2b, 3b, 4b, 5b, 7
Type MADAGASCAR (bullet) Andasibe; 18°56′00″S, 48°25′06″E; 940 m elev.; 01 December 2002; A. Hladik & C.-M. Hladik 6760 (Holotype: TAN!, Isotypes: K!, MO!, P!).Paratypes MADAGASCAR (bullet) Andasibe; 18°56′00″S, 48°25′06″E; 940 m elev., 23 Aug. 1998, A. Hladik & al. 6239 (P!, fruits) (bullet) June 2001, A. Hladik & al. 6650 (P!, leaves, fruits, bracts) (bullet) Andasibe-Mantadia area, Vakôna, Kalonora; 18°53′17.3″S, 48°25′51.3″E, 08 November 2018, 934 m elev., T. Haevermans & al. 832 (K!, MO!, P!, TAN!).Diagnosis Similar to Ravenala madagascariensis but differs in its non-suckering habit, decurrent juvenile lamina bases, toroidal distribution of juvenile laminae, smaller number of leaves simultaneously alive on the adult plant, dark green lamina and green non waxy petiole, smaller leaves, smaller number of live inflorescences, smaller number of bracts in an inflorescence, non-waxy bracts, sub-simultaneous flowering, smaller flowers, smaller inflorescences, non-persistence of entire bracts on dry infructescences, October/November flowering period.Distribution Andasibe-Mantadia, Ranomafana21. Restricted to Madagascar.Preliminary IUCN assessments We propose a Data Deficient status for R. blancii; further fieldwork is required to understand its precise distribution and the status of its populations33.Ecology High-elevation species found in eastern rainforests at elevations between 600 and 1,100 m. The species seems to favor cool tropical humid and shady conditions.Etymology This species is named after Dr. Patrick Blanc, world renowned botanist, plant ecologist and street artist, inventor of the planted vertical walls known as “Mur Végétal” and who first recognized the sheer originality of the juvenile phases of this peculiar taxon.Description Plants solitary (never suckering), 10–15 meters tall (adult), trunk circumference (d.b.h.) 20–30 cm, juvenile laminae distributed in a toroidal shape, adult laminae arranged in a regular fan, 9–16 leaves simultaneously alive on the adult plant, 2–4 leaves between inflorescences. Leaves adult petiole 240–310 cm long, green, not waxy, sheath margin undeveloped, entire, not drying, smooth with a worn-out irregular aspect (Fig. 3b), petiole/lamina ratio 1.8–(2.0)–2.2, adult lamina 120–160 (times) 90–104 cm, dark green, juvenile lamina base decurrent. Inflorescences 2–3 live lateral inflorescences at a time, (60 times 70) cm (peduncle excluded), 4–6 bracts per inflorescence, bracts 160–350 (times) 50–90 mm, no waxiness (Fig. 4b), margin color uniformly green, cincinnii of 5–14 flowers per bract, flowering sub-simultaneously, bracteoles sometimes pink colored. Flowers 165–280 mm long (ovary included), inferior ovary 40–50 mm long, perianth whitish-yellowish, sepals narrowly triangular 125–231 (times) 10–12 mm, sheathing (fused) petals narrowly triangular 105–190 (times 10) mm, free petal acicular 105–178 (times 3)–5 mm, free petal and fused petals of sub-equal size with a mean free petal / mean fused petal length ratio = 1.0, petal blotches absent or present, stamens (roughly) the same size as the perianth, 115–186 mm long, style 132–220 mm long, stigma 20-25 mm long, ovoid to ovoid-pointed with a basal constriction. Infructescences compact (bract bases imbricate at all stages of maturity), torn and degraded bracts on mature infructescence, old infructescences deciduous, 5–14 fruits per bract. Fruits 80–120 (times) 32–45 mm, trilocular septifragal capsule, apices conical (Fig. 2b), seeds 6–10 (times) 3.2–6 mm, shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.Note The strong leaf dimorphism between juvenile and adult forms is characteristic of this species13, a phenomenon which is not present in the other taxa. The base of the juvenile plant usually grows buried in the leaf litter due to the action of traction roots13, its decurrent leaves (Fig. 7) giving it the aspect of a bird’s nest fern.Figure 7Ravenala blancii. (a) juvenile plant habit with roots. (b) juvenile plant showing the arrangement of laminae. (c) adult plant habit. (d) mature infructescence segment. (e) juvenile leaf showing the attenuate base of the lamina. (f) inflorescence with sub-simultaneous opening of the flowers. (g) young infructescence with already degraded bracts. (h) seeds with arilla. (i) open flower. (j) details of the stigma. (k) style. Ink drawings on (75 , upmu) polyester tracing paper by Agathe Haevermans© from specimens Hladik 6790, 6239, 6650, Haevermans et al. 832, and observations in-situ.Full size image
Ravenala grandis Haev., Razanats., A. Hladik & P. Blanc sp. nov.—Figs. 2c, 3c, 4c, 5cType. MADAGASCAR (bullet) Ampasimbe Commune, Maromaniry Fokontany, along Route Nationale, 18°57′41.8″S, 48°42′41.4″E, 258 m elev., 08 November 2018, T. Haevermans, A. Haevermans & J. Razanatsoa 831 (Holotype: TAN!, Isotypes: K!, MO!, P!).Paratypes MADAGASCAR (bullet) Varifoana, près d’Ambohimahasoa-sud, 15 May 1964, R. Capuron 26014SF (P02234597!) (bullet) Soanierana-Antasibe[Andasibe], 350 m elev., 10 December 1938, H.J. Lam & A.D.J. Meeuse 5867 (L-WAG.1111450!, L-WAG.1111451!, L-WAG.1111452!, L-WAG.1111453!, L-L.1477714!, L-L.1477715!).Diagnosis Similar to Ravenala madagascariensis but differs in its non-suckering habit, much larger dimensions, very thick leathery laminae, very waxy dark green-yellowish petioles, much larger bracts and overall dimensions, whitish/pure white perianth, strong reddish-pink stripes on its bracteoles, cylindrical stigma without basal constriction, stamens much shorter than perianth, and fruit with a truncated apex.Distribution Eastern rainforests at around 200–500 m elevation in Madagascar13,20.Preliminary IUCN assessments We propose a Data Deficient status for R. grandis; further fieldwork is required to understand its precise distribution and the status of its populations33.Ecology This species seems to favor growing in low discontinuous forests on inselbergs12 and thrives in secondary degraded vegetation on the slopes of eastern rain forests.Etymology The name of this species is in reference to its stature and habit, the most robust species of Ravenala known.Description Plants solitary (never suckering), 20–30 meters tall (adult), trunk circumference (d.b.h.) 30 cm, juvenile and adult laminae distributed in a perfect fan, 15–30 leaves simultaneously alive on the adult plant, usually 3 leaves between inflorescences. Leaves adult petiole 390–440 cm long, dark green/light green-yellowish, very waxy (Fig. 3c), sheath margin moderately developed to undeveloped (0–9 mm), entire on young leaves, splitting and dryish when old, petiole/lamina ratio 1.8–(2.2)–2.6, adult lamina 170–230 (times) 94–120 cm, light green, juvenile lamina base non-decurrent. Inflorescences 4–6 live lateral inflorescences at a time, 100–120 (times) 80–100 cm (peduncle excluded), 10–20 bracts per inflorescence, bracts 440–540 (times) 140–170 mm, some waxiness (Fig. 4c), margin color uniformly green, cincinnii of ca. 20 flowers per bract, flowering sequentially, bracteoles with a strong reddish-pink stripe. Flowers 300 mm long (ovary included), inferior ovary 50–70 mm long, perianth whitish/pure white, sepals narrowly triangular 220–240 (times) 10–15 mm, sheathing (fused) petals narrowly triangular 210–220 (times) 10–12 mm, free petal acicular 150–170 (times 3) mm, slightly smaller than the rest of the perianth with a mean free petal / mean fused petal length ratio = 0.8, petal blotches absent, stamens much shorter than the perianth, 180–200 mm long, style 180–210 mm long, stigma 14–16 mm long, oblong without basal constriction (almost indistinguishable from style). Infructescences lax (bract bases not imbricate at some stages of maturity), stiff and coriaceous persisting bracts on mature infructescence, old infructescences deciduous, 5–18 fruits per bract. Fruits 100–120 (times) 35–40 mm, trilocular septifragal capsule, apices truncate (Fig. 2c), seeds shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.Note The leaves of this species are the most robust and tough of all Ravenala species, with a thick leathery texture, making it the material of choice for building roofs35.
Ravenala hladikorum Haev., Razanats., V. Jeannoda & P. Blanc sp. nov. — Figs. 2f, 3f, 4f, 5fType MADAGASCAR (bullet) Andasibe; 18°56′00″S, 48°25′06″E; 940 m elev.; 05 February 2004; A. Hladik & C.-M. Hladik 6842 (Holotype: TAN!, Isotype: P!). Paratypes. MADAGASCAR (bullet) Andasibe; 18°56′00″S, 48°25′06″E; 940 m elev.; 23 August 1998; A. Hladik & al. 6240 (fruit with seeds: P!). (bullet) Andasibe-Mantadia area, Vakôna, Kalonora; 18°53′17.3″S, 48°25′51.3″E; 934 m elev., 08 November 2018; T. Haevermans & al. 833 (TAN!, P!, K!, MO!).Diagnosis Similar to Ravenala madagascariensis but differs in its non-suckering habit, the alternate positioning of its adult laminae, its dark green leaves, non-waxy petioles with their very papyraceous petiole sheath margins, more than 1 cm long, smaller lamina dimensions, smaller number of simultaneously live inflorescences, purple stripe on bracts and on bracteoles, non-waxy inflorescences, smaller inflorescences, dense infructescences, truncated fruit apices, and short flowering period from November to December.Distribution Andasibe, Mantady, Ranomafana21. Restricted to Madagascar.Preliminary IUCN assessments We propose a Data Deficient status for R. hladikorum; further fieldwork is required to understand its precise distribution and the status of its populations33.Ecology High-elevation species found in eastern rainforests at elevations between 600 and 1100 m. The species seems to favor cool tropical humid and shady conditions.Etymology This species is named in honor of Annette and Claude-Marcel Hladik from the Muséum National d’Histoire Naturelle in Paris, who dedicated their lives to the study of Madagascan biodiversity and contributed greatly to the discovery of this species.Description Plants solitary (never suckering), 10–15 meters tall (adult), trunk circumference (d.b.h.) 20–30 cm, juvenile laminae distributed like a fan, adult laminae arranged in an irregular fan, 9–18 leaves simultaneously alive on the adult plant, 1–3 leaves between inflorescences. Leaves adult petiole 280–440 cm long, greenish-yellow, not waxy (Fig. 3f), sheath margin very developed (10 mm and more), split, very papyraceous with min. 1 cm brown dry expansions, petiole/lamina ratio 2.1–(2.42)–2.8, adult lamina 120–160 (times) 102–116 cm, dark green, juvenile lamina base non-decurrent. Inflorescences 2–3 live lateral inflorescences at a time, (60 times 90) cm (peduncle excluded), 4–7 bracts per inflorescence, bracts 150–510 (times) 64–100 mm, no waxiness (Fig. 4f), margin green with a purple stripe, cincinnii of 5–14 flowers per bract, sequentially flowering, bracteoles with a dark purple colored stripe. Flowers 240–320 mm long (ovary included), inferior ovary 40–60 mm long, perianth whitish, sepals narrowly triangular 210–265(times)ca. 10 mm, sheathing (fused) petals narrowly triangular 190–240(times)ca. 10 mm , free petal acicular 135–220 (times) 5 mm, almost the same size as the fused petals with a mean free petal / mean fused petal length ratio = 0.9, petal blotches unknown, stamens (roughly) the same size as the perianth, 170–230 mm long, style 187–250 mm long, stigma 20–25 mm long, ovoid with a basal constriction. Infructescences compact (bract bases imbricate at all stages of maturity), stiff and coriaceous persistent bracts on mature infructescences, old infructescences deciduous, 5–14 fruits per bract. Fruits 82–108 (times) 34–48 mm, trilocular septifragal capsule, apices truncate (Fig. 2f), seeds 4–9 (times) 3–6 mm, shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.
Ravenala menahirana Haev. & Razanats. sp. nov.—Figs. 2e, 3e, 4e, 5eType MADAGASCAR (bullet) Foulpointe, Analalava Forest; 17°42.3′S, 49°27.38′E; 50 m elev.; 20 March 2016; T.Haevermans, M. Vorontsova, S. Dransfield & J. Razanatsoa 826 (Holotype: TAN!, Isotypes: P!, K !, MO!).Diagnosis Similar to Ravenala madagascariensis but differs in its non-suckering habit, the alternate dark green laminae tending not to form a perfect fan (Fig. 5e), dark red petioles with a zigzagging well developed dryish sheath margin, more strongly obovoid laminae, smaller number of simultaneously live inflorescences, smaller inflorescences tinged with red, pure white/whitish perianth, smaller flowers, dense infructescences, the fruit apices truncate with a mucro, and subequal free and fused petals.Distribution Appears to be restricted to the east coast in the area around Analalava-Foulpointe up to the Mananara-Avaratra area. Two human observations from Marojejy (North-East) and Tampolo (Masoala) seem also to be this species. Restricted to Madagascar.Preliminary IUCN assessments We propose a Data Deficient status for R. menahirana; further fieldwork is required to understand its precise distribution and the status of its populations33.Ecology This coastal forest-dwelling species favors low-elevation tropical humid conditions in the Analalava-Foulpointe area, extending north to Mananara-Avaratra area, and maybe up to Marojejy.Etymology The name of this species is in reference to one of its local names “menahirana”, given to the species in the Analalava-Foulpointe area and meaning “red ravenala”.Description Plants solitary (never suckering), 6–10 meters tall (adult), trunk circumference (d.b.h.) 20–30 cm, juvenile laminae distributed like a fan, adult laminae arranged in an irregular to regular fan, 12–18 leaves simultaneously alive on the adult plant, 3 leaves between inflorescences. Leaves adult petiole 200–230 cm long, dark red, slightly to very waxy, sheath margin very developed (10 mm and more), red, entire, forming a three dimensional zigzag pattern (Fig. 3e), then splitting and drying on old leaves, petiole/lamina ratio 1.4–(1.7)–1.9, adult lamina (350 times 120) cm, lamina color dark green, juvenile lamina base non-decurrent. Inflorescences 1–2 live lateral inflorescences at a time, (60 times 70) cm (peduncle excluded), 10–12 bracts per inflorescence, bracts 260–360 (times) 50–80 mm, very waxy (Fig. 4e), margin color uniformly reddish-green, cincinnii of 8–12 flowers per bract, flowering sequentially, no colored stripe on bracteoles (apices sometimes suffused with pink). Flowers 220–250 mm long (ovary included), inferior ovary 40–60 mm long, perianth pure white to whitish, sepals narrowly triangular 180–230 (times) 12–16 mm, sheathing (fused) petals narrowly triangular 160–180 (times) 5 mm, free petal acicular 160–170 (times) 5 mm, free petal the same size as the remaining perianth with a mean free petal / mean fused petal length ratio = 1.0, petal blotches absent, stamens the same size (roughly) as the perianth, stamen 150–160 mm long, style 150–200 mm long, stigma 10 mm long, oblong with a basal constriction. Infructescences compact (bract bases imbricate at all stages of maturity), stiff and coriaceous persisting bracts on mature infructescences, old infructescences deciduous, 8–12 fruits per bract. Fruits 80–100 (times) 30–35 mm, trilocular septifragal capsule, apices truncate with a mucro (Fig. 2e), seeds shiny, dark brown, mostly globose, varying in shape according to their distribution in the capsule, ultramarine blue aril.Note This species is similar to R. hladikorum but is easily distinguished by, in addition to its petioles and its ecology, its truncate mucronate fruit apices, the shape of the synflorescence bracts and the absence of a red stripe on the cyme bracteoles.Identification key to the species of genus Ravenala More

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Eukaryote composition (V9_PR2)Using V9_PR2 we were able to assign a total of 15 668 (shotgun) and 90 689 reads for the shotgun and amplicon data, respectively. These reads represented 14%, 54%, 0 and 32% (shotgun), and 0%, 29%, 0 and 71% (amplicon) unassigned cellular organisms, Bacteria, Archaea and Eukaryota, respectively. Within the eukaryotes, we determined 51 and 64 taxa for shotgun and amplicon data, respectively. Abundant taxa (average abundance >0.1% across all samples; 31 and 27 taxa in shotgun and amplicon, respectively) are shown in Fig. 2. The latter includes 23 taxa (including assignments made on “Eukaryota” level) that were shared between shotgun and amplicon, and four taxa only detected in the amplicon data (Fig. 2C).Fig. 2: Eukaryote composition in five Santa Barbara Basin sediment samples post-alignment with V9_PR2 database.Composition is shown in relative abundances for (A) shotgun, and (B) amplicon data (phylum-level). The surface sample should be considered with caution in both (A) and (B) due to the possibility of contamination (see “Methods”). C Venn diagram showing eukaryote taxa richness (phylum level) in the shotgun and amplicon data after alignment with the V9_PR2 database (diagram areas are proportional to the total number of taxa included, for a list of shared/non-shared taxa see Supplementary Material Fig. 1). Only taxa abundant on average >0.1% are included, as they make up >99% of the eukaryote composition.Full size imageWithin shotgun, the most abundant eukaryotes were Ascomycota (53%), Telonemia (11%), Eukaryota (not further determined, 8%), Polycystinea (4%), Dinophyceae (3.8%), Streptophyta (3.2%), Amoebozoa (3%), Cercozoa (1.6%), Bacillariophyta (1.6%), Arthropoda (1%). In the amplicon data, the most abundant eukaryotes were Ascomycota (33%), Apicomplexa (30%), Dinophyceae (9.5%), Stramenopiles (6.3%), Eukaryota (4.9%), Polycystinea (3.5%), Foraminifera (3.2%), Cercozoa (1.1%) and Chordata (1%). Thus, a total of 10 and 9 taxa were abundant with >1% (average across all samples) in the shotgun and amplicon data, including only five taxa (Ascomycota, Eukaryota, Dinophyceae, Polycystinea, Cercozoa) that were picked up by both methods (i.e., are amongst the shared taxa in Fig. 2C, Supplementary Material Fig. 1). Taxa detected by one method or the other were slightly rarer species (between 0.1 and 1% average relative abundance across all samples; Supplementary Material Table 3).The shotgun EBC detected two taxonomic groups, one prokaryotic (Gammaproteobacteria) and one eukaryotic (Poacea). The amplicon EBC detected 46 taxa, of which 12 were prokaryotes and 34 were eukaryotes, including dinoflagellate taxa (Dinophysis and Alexandrium), Calanoida and Bacillariophyta (copepods and diatoms, respectively; Supplementary Material Table 1). While any reads assigned to EBC taxa were removed from samples, including reads assigned to the Bacillariophyta node, reads assigned to Bacillariophyta at lower taxonomic levels (e.g., Bacillariophycidae, Bacillariaceae, etc.) remain summarised under the phylum-level Bacillariophyta node (Fig. 2).Relationship between Eukaryota composition and V9_PR2 reference sequence lengthV9_PR2 reference sequence-lengths for the relatively abundant taxa ( >0.1% across all samples, including all taxa that were shared and assigned below eukaryote-level, i.e., 22 taxa, see Supplementary Material Table 3) were around the overall average sequence length of the V9_PR2 database (121 bp) (Fig. 3). However, considerable length variation was observed, with most of the abundant taxa being represented by shorter than average reference sequences in the V9_PR2 database, and a few taxa (e.g., Arthropoda, Opisthokonta and Amoebozoa) with a number of reference sequences longer than average (Fig. 3).Fig. 3: Average sequence lengths for individual eukaryote taxa as per in the V9_PR2 database (A) and read counts for these taxa in shotgun (SG) and amplicon (Ampl) data (B).Listed are all taxa that occurred on average >0.1% across all samples in either the shotgun or amplicon dataset, or both. Only taxa that were determined in both shotgun and amplicon data are included.Full size imageWe determined a negative correlation between the average V9_PR2 reference sequence length (V9PR2AL) and the A:SG read counts ratio per taxon for all samples (rV9PR2AL,A:SG_1.2 = −0.27269, rV9PR2AL,A:SG_4.3 = −0.33233, rV9PR2AL,A:SG_7.3 = −0.28064, rV9PR2AL,A:SG_11.8 = −0.32559, rV9PR2AL,A:SG_16.4 = −0.30078). This means that shorter V9_PR2 reference sequences for our abundant taxa were associated with an overamplification of these taxa in the amplicon data (for average V9_PR2 reference sequence length of the abundant taxa and A:SG ratios see Supplementary Material Table 4).Eukaryota and Bacillariophyta sequence length and coverage post-V9_PR2 alignmentSequences assigned to Eukaryota in shotgun were on average 112 bp and in amplicon data 161 bp, i.e., shotgun reads were around ~50 bp shorter than amplicon reads (Table 2). Bases covered in shotgun were ~40 bp shorter than in amplicon data (Table 2). Similarly, sequences assigned to Bacillariophyta were on average 124 and 167 bp in shotgun and amplicon data, respectively, so showed an ~40 bp difference. For Eukaryota, there was a difference of ~23 bp and 29 bp between sequence length and coverage in shotgun and amplicon data, respectively. For Bacillariophyta, we found a ~36 and ~37 bp difference between sequence length and coverage in shotgun and amplicon data, respectively.Table 2 Lengths and coverage of sequences assigned to Eukaryota and Bacillariophyta in shotgun and amplicon data.Full size tableBacillariophyta read lengths and coverage were similar to those of Eukaryota, for both shotgun and amplicon data (Table 2). Variation in sequence lengths and coverage was much higher in shotgun than in amplicon data. We found no trend towards shorter (i.e., more fragmented) sequences with increasing subseafloor depth for either Eukaryota or Bacillariophyta in the shotgun data. Eukaryota shotgun read lengths were on average ~9 bp shorter (112 bp) than the average reference sequences in the V9_PR2 database (121 bp).Diatom composition detected via diat-rbcL and read length characteristicsA total of 60 (shotgun) and 80 674 (amplicon) reads were assigned to diatoms (Fig. 4). In total, 27 taxa were determined in the shotgun, and 140 in the amplicon dataset. When considering the “abundant” taxa (on average >0.1%), 27 and 49 diatoms were determined in the shotgun and amplicon data, respectively (Fig. 4). A total of 10 taxa were shared between the two datasets Bacillariophyta, Bacillariophycidae, Chaetoceros, C. cf. pseudobrevis 2 SEH-2013, Pseudo-nitzschia, P. fryxelliana, Thalassiosiraceae, Thalassiosirales, Thalassiosira and T. oceanica (Fig. 4C, Supplementary Material Fig. 2). Sequences assigned to diatoms via diat-rbcL were shorter (by ~16 bp) in the shotgun than in the amplicon data, with amplicon read lengths and coverage all 76 + 1 bases (Table 3).Fig. 4: Diatom composition in the Santa Barbara Basin sediment samples post-alignment with diat-rbcL database.Diatom composition is shown as relative abundance for (A) shotgun and (B) amplicon data. The surface sample should be considered with caution in both (A) and (B) due to the possibility of contamination (see “Methods”). C Venn diagram showing diatom taxa richness (species level) in the shotgun and amplicon data after alignment with the diat-rbcL database (diagram areas are proportional to the total number of taxa included, for a list of shared/non-shared taxa see Supplementary Material Fig. 2). Only taxa abundant on average >0.1% are included (in A, B, C).Full size imageTable 3 Bacillariophyta sequence lengths in shotgun and amplicon datasets.Full size tableNo diatoms were detected in the shotgun EBC, however, 45 taxa were determined in the amplicon EBC with most reads assigned to Chaetoceros spp. (especially, Chaetoceros debilis, C. socialis and C. radicans), several Thalassiosira and Pseudo-nitzschia species, as well as others (Supplementary Material Table 2).Comparison of V9_PR2 vs. diat-rbcL derived diatom compositionIn the shotgun data, 79 and 60 sequences were assigned to diatoms using V9_PR2 and diat-rbcL as the reference database, respectively, and composition differed considerably (Fig. 5). Using V9_PR2, diatoms were mostly assigned on relatively high taxonomic levels (e.g., Bacillariophyta) with few taxa being differentiated sporadically in the different samples (Fig. 5A, Supplementary Material Fig. 3). Using diat-rbcL, Chaetoceros, Thalassiosira and Pseudo-nitzschia were more prominent (Fig. 5B).Fig. 5: Comparison of diatom composition in Santa Barbara Basin sediment samples determined in shotgun and amplicon data using the V9_PR2 and diat-rbcL databases.Relative abundance of diatoms (genus level) in the shotgun data after aligning to (A) V9_PR2 and (B) diat-rbcL. Relative abundance of diatoms (genus level) in the amplicon data after aligning to (C) V9_PR2 and (D) diat-rbcL. The surface sample should be considered with caution in (A–D) due to the possibility of contamination (see “Methods”). Venn diagrams of shared and non-shared diatom taxa after alignment to the V9_PR2 (18S-V9) and diat-rbcL databases for the shotgun (E) and amplicon (F) data (species level, diagram areas are proportional to the total number of species included). For a complete species list and their read counts per sample see Supplementary Material Fig. 3, Supplementary Material Table 5.Full size imageIn the amplicon data, 329 sequences were assigned to diatoms using V9_PR2, and 80 674 using diat-rbcL. Using V9_PR2, few taxa were detected in the two top samples (Leptocylindrus and Fragilariaceae at 1.2 mbsf, Bacillariophycidae and Bacillariaceae at 4.3 mbsf) while the lowermost samples were more diverse (Fig. 5C). Using diat-rbcL, most reads were assigned to Thalassiosira, Chaetoceros, and Pseudo-nitzschia, with other taxa sporadically occurring at different depths (Fig. 5D). For a complete species list and their read counts see Supplementary Material Fig. 3, and Supplementary Material Table 5.We found large differences in the number of shared vs. non-shared taxa between shotgun and amplicon data, and V9_PR2 and diat-rbcL alignments (Fig. 5E, F). Database inspections showed that all taxa detected via V9_PR2 were also represented in the diat-rbcL database, except Rhizosoleniaceae. However, out of the 22 taxa exclusively detected via diat-rbcL in shotgun (Fig. 5E, F), 10 are only represented in the diat-rbcL database (Pseudo-nitzschia caciantha, P. dolorosa, Chaetoceros cf. contortus 1 SEH-2013, C. cf. lorenzianus 2 SEH-2013, C. cf. pseudobrevis 2 SEH-2013, Thalassiosirales, Thalassiosiraceae, Coscinodiscus wailesii, Arcocellulus mammifer, Meuniera membranacea, Supplementary Material Fig. 3). Similarly, out of the 134 taxa exclusively detected via diat-rbcl in amplicon, 84 were in this database only, noticeably including several species and strains of Chaetoceros, Pseudo-nitzschia, Thalassiosira and Cylindrotheca (eg., additions SHE-2013, BOF in species names), amongst others (see Supplementary Material Fig. 3, Supplementary Material Table 5). More

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