Xylopia amplexicaulis (Lam.) Baill. An [ n ] ona amplexicaulis Lam., Encyclopédie méthodique, Botanique 2 (1): 127 (1786) . Pseudannona amplexicaulis (Lam.) Saff., Journal of the Washington Academy of Sciences 3: 18 (1913) A revision of Xylopia L. (Annonaceae): the species of Madagascar and the Mascarene islands Johnson, David M. Murray, Nancy A. Adansonia 2020 2020-02-05 42 1 1 88 5CLZ7 (Lam.) Baill. Baill. Lam. [277,659,1651,1678] Magnoliopsida Annonaceae Xylopia Plantae Magnoliales 17 16 Tracheophyta species amplexicaulis   ( Figs 5; 6A)    Adansonia; Recueil d’Observations botaniques4: 142 (1864). —   An[ n] ona  amplexicaulis Lam., Encyclopédie méthodique, Botanique2 (1): 127 (1786).—   Pseudannona  amplexicaulis(Lam.) Saff., Journal of the Washington Academy of Sciences3: 18 (1913). — Type:   Mauritius.“ Madagascar”, s. d. (fl.), Commerson s. n.(lecto-, here designated, P[ P030364]!).  MATERIAL EXAMINED. —   Mauritius.Without definite locality, s.d. (fl.), Ayres 11( MAU[ MAU0000966]); without definite locality, s. d. (fl.) [presented in 1863], herb. Blackburn s. n.(K);  Pér[r] ier (reserve), s. d. (fl., in spirit collection only, see Fig. 5),  Bosser21824(P[P01954053]); upland forests, without definite locality, s. d. (st.),  Bouton10( MAU[ MAU0000986]);  Île de France, without specific locality,  Commersons. n.(B fragment, FI-W, G[G197204/2, G197204/1], MPU[2 sheets], P[P030363, P01954064, P01954065, Drake Herb.ex Herb. Richard, P01954066]);  Crown Land Perrierin the National Reserves,  21.III.1950(fl.),  Duljeet F. D.77( MAU[ MAU0000961, MAU0000962]); without definite locality, Herbier du Petit-Thouars s. n.(P[P01954067]);  Gaulettesserrées, 20°12’40”S, 57°38’23”E,  8.XII.1998(st.),  Florenset al. 22975( MAU[ MAU0000982]); without definite locality, 1858 (fl.), Grey s. n.(K);  Perrier Nature Reserve,  5.III.1974(fl.),  Guého16133( MAU[ MAU0000984, MAU00000985]);  Perrier Nature Reserve, near Mareaux Vacoas,  1800 ft,  3.II.1973(fl.),  Lorence M57( MO);  Perrier Nature Reserve,  14.III.1975(fr.),  Lorence1180( MAU[ MAU0000988, spirit collection only]);  Bel Ombre forestreserve,  200-400 m,  16.II.1979(buds),  Lorence& Lecordier2405(K n.v., MAU[ MAU0000983], MO);  Perrier Nature Reserve, near Mareaux Vacoas,  600 m,  14.III.1979(fl.),  Lorence2535(K n.v., MO, P);  Grand Port, s. d. (fl.),  Wallichs. n.( BM).  DISTRIBUTION, ECOLOGY AND CONSERVATION STATUS. —  Xylopia amplexicaulisis restricted to the island of Mauritius( Fig. 7), where it occurs in forests and thickets, sometimes along streams and sometimes in habitats dominated by Sideroxylonspecies, at elevations of 200-600 m. Baker (1877)reported it as occurring in mountain woods at Nouvelle Découverte and La Savanne, but no specimens from these localities have been seen; C. Baider (personal communication) lists the localities where the plant has most recently been observed and photographed as Perrier, Bel Ombre, Gaulettes Serrées, Mt. Lion, Brise Fer, Mare Longue/Machabe, and Florin. Specimens with flowers have been collected in February and March and with fruits in March. The species was initially assessed as Endangered by Page (1998a), and is currently listed as Critically Endangered on the IUCN Red List. In 1998 it was only known from 20 mature trees distributed across three localities on the island. Our EOO and AOO estimates of 334 km 2and 16 km 2, respectively ( Table 2), support the assessments.  DESCRIPTION  Treeup 5 mtall; d.b.h. 6 cm.  Twigsglabrous or sparsely appressed-pubescent, the hairs 0.1- 0.2 mmlong, but soon glabrate; number of axillary branches per node unknown.  Leaveswith larger blades 9.7-13.4 cmlong, 5.5-7.0 cm wide, subcoriaceous, discolorous, dull green to brown adaxially, tan and occasionally somewhat glaucous abaxially, ovate to lanceolate, apex acute, obtuse, rounded, or emarginate, base cordate-auriculate with the c. 1 cmlong lobes clasping the stem, margin flat, not revolute, glabrous on both surfaces; midrib concolorous with blade adaxially, secondary veins weakly brochidodromous, 13-15 per side, diverging at 65-70° from midrib, these and higher-order veins slightly raised and forming a conspicuous reticulum on both surfaces; petiole 0-2 mmlong, broad and flattened, wrinkled, glabrous.  TABLE 2. — Preliminary and published conservation assessments for Madagascar and Mascarene  Xylopiaspecies.Abbreviations: CR, Critically Endangered; D, based on IUCN Criterion D; DD, Data Deficient; EN, Endangered; VU, Vulnerable. Sources: 1, Page 1998a; 2, Lantoarisoa 2015; 3, Callmander et al.2009; 4, Page 1998b; 5, Page 1998c. *, See Discussion under  X. flexuosaDiels.     EOO  AOO   Species  Status  (km 2 )  (km 2 )  Localities    X. ambanjensisCavaco & Keraudren EN (D) 5498 48 12    X. amplexicaulis(Lam.) Baill. CR1 334 16 4    X. anomala, sp. nov. EN (D) 3468 24 6    X. australis, sp. nov. EN (D) 185 24 6    X. beananensisCavaco & Keraudren DD 192 983 80 20    X. bemarivensisDiels DD 171 046 172 43    X. buxifoliaBaill. DD 149 436 96 24    X. capuroniiCavaco & Keraudren CR (D) 3846 16 4    X. carinata, sp. nov. EN (D) 1110 8 2    X. danguyellaGhesq. ex Cavaco & Keraudren EN (D) 2191 28 7    X. dielsiiCavaco & Keraudren EN (D) 461 12 3    X. fananehanensisCavaco & Keraudren EN (D) 1510 24 6    X. flexuosaDiels* EN2 41 558 48 12    X. galokothamna, sp. nov. EN (D) 5 8 2    X. ghesquiereanaCavaco & Keraudren CR (D) 0 4 1    X. humbertiiGhesq. ex Cavaco & Keraudren CR (D) 0 4 1    X. humblotianaBaill. DD 41 472 240 60    X. kalabenonensisD.M.Johnson, Deroin, & Callmander CR3 0 4 1    X. lamarckiiBaill. CR4 270 32 8    X. lamiiCavaco & Keraudren EN (D) 6316 24 6    X. lastellianaBaill. DD 124 820 72 18    X. lemuricaDiels DD 38 756 68 17    X. lokobensis, sp. nov. EN (D) 144 8 2    X. longirostra, sp. nov. EN (D) 5003 24 6    X. madagascariensisCavaco & Keraudren EN (D) 3780 24 6    X. marojejyana, sp. nov. CR (D) 0 4 1    X. perrieriDiels DD 50 587 120 30    X. ravelonarivoi, sp. nov. EN (D) 7118 40 10    X. retusa, sp. nov. EN (D) 1751 32 8    X. richardiiBoivin ex Baill. VU5 558 36 9    X. sahafariensisCavaco & Keraudren EN (D) 853 12 3    X. sclerophylla, sp. nov. EN (D) 859 20 5    X. sericolampraDiels EN (D) 1940 12 3  Inflorescencesaxillary, 1-2-flowered, if more than one flower then peduncle present, sparsely appressed-pubescent except for distal half of peduncle, which is glabrous and sometimes glaucous; peduncle c. 3.4 mmlong; pedicels 1-2 per peduncle, 7-15 mmlong, 2-2.2 mmthick; bracts 3-4, on proximal half of pedicel, caducous or occasionally with distal bract persistent, distal bract 1.7-4 mmlong, semicircular; buds oblong, apex obtuse.  Sepalserect to slightly spreading at anthesis, c.¼-connate, 5-8 mmlong, 6.5-7 mmwide, coriaceous, broadly triangular and concave, apex acute, sparsely pubescent abaxially.  Petalscreamy white, wine-colored with a white patch at the base adaxially and with a pink to purple tinge on the outer petals abaxially in vivo; outer petals possibly erect at anthesis, fleshy, 31-45 mmlong, 10-14 mmwide at base, 14-15 mmwide at midpoint, fleshy, oblong, flat, apex obtuse, keeled on apical ¼-¹⁄5 but becoming concave toward the base adaxially, densely but finely appressed-pubescent abaxially; inner petals with position at anthesis unknown, c. 26 mmlong, c. 4 mmwide at base and midpoint, fleshy, lanceolate, laterally compressed, apex acute, base concave with undifferentiated margin, indument not determinable.  Stamens c.400; fertile stamens 0.9-1.2 mmlong, narrowly oblong, anther connective apex 0.1-0.2 mmlong, depressedglobose to bluntly conical, slightly overhanging anther thecae, papillate, anthers 13-14-locellate, filament 0.3-0.4 mmlong; outer staminodes 0.8-1.0 mm long, oblong, apex truncate to rounded; inner staminodes c. 0.9 mmlong, oblong to quadrate, apex truncate; staminal cone c. 3.5 mmin diameter, c. 0.9 mmhigh, completely concealing the ovaries, rim even.  Carpels c.25; ovaries 0.9-1.3 mmlong, lanceolate-oblong, glabrous, stigmas loosely appressed, c. 1 mmlong, filiform, glabrous.  Fruitof c.17 monocarps borne on a pedicel 25-30 mmlong, glabrate; torus c. 10.5 mmin diameter, c. 9 mmhigh, depressed-globose; monocarps with light yellow faintly glaucous exterior and bright red endocarp in vivo, 2-4.5 cmlong, 0.6- 1 cmwide and thick, clavate to fusiform, not torulose, apex obtuse, base gradually contracted into a stipe 8.8-10 mmlong, 1.7-3.4 mmthick, smooth, glabrous; pericarp thin, leathery.  FIG. 5. —  Xylopia amplexicaulis(Lam.) Baill.and  X. lamarckiiBaill. A -H,  X. amplexicaulis: A, habit; B, C, flower, longitudinal section showing carpels, staminal cone, inner petals, and outer petals; D, outer petal, adaxial view; E, inner petal, adaxial view; F, stamen, abaxial view; G, fruit; H, seed, side view; I,  X. lamarckii, habit. Flowers and fruits are based on fluid-preserved specimens, and the sizes are larger than indicated in the species descriptions. A, From Commerson s. n.(P); B -F, from Bosser 21824(P, spirit collection and photograph); G, H, from Lorence 1180(MAU, spirit collection); I, from Commerson s. n.(P). Figure reproduced from Keraudren-Aymonin 1980, re-used with permission of T. Mourier, Service Editions, Mission culture scientifique et technique, Institut de recherche pour le développement (IRD), Montpellier. Scale bars: A, B, D, E, G, I, 3 cm; C, H, 2 cm; F, 1 mm.  FIG. 6. — A, Shoot of  X. amplexicaulis(Lam.) Baill., showing flower bud at left and clasping leaf bases visible at several nodes; B, branch of  X. ravelonarivoi, sp. nov., with fruit ( Ravelonarivo 4171); C, flower of  X. humblotianaBaill., side view ( Schatz 4475); D, buds of  X. richardiiBoivin ex Baill.; E, close-up of flower interior of  X. lamarckiiBaill., showing red coloration at bases of petals, and the stamens and stigmas; outer staminodes are visible along the edge of the androecium; F, flower of  X. richardiiat anthesis, one outer petal and one inner petal removed, so that androecium and stigmas are visible; G, leaves and fruit of  X. anomala, sp. nov., ( Rakotonirina 48); H, leaves and fruit of  X. sericolampraDiels. Photos: A, Carlos Magdalena; B, D. Ravelonarivo; C, G. E. Schatz; D -F, C. Kaiser-Bunbury; G, C. Birkinshaw; H, R. N. Rabarijaona. B, C, G, used under the terms of the Creative Commons license for Tropicos, the botanical information system at the Missouri Botanical Garden; H, used under the terms of the the Creative Commons license for iNaturalist.  Seedsup to 6 per monocarp, in a single row, oblique to parallel to long axis, 10-12 mmlong, 4-5 mmwide, narrowly ellipsoid, circular in cross-section, black, smooth, shiny, perichalazal ring not evident; sarcotesta absent; aril translucent white in vivo, bilobed, lobes c. 3.8 mmlong and wide, fleshy, semicircular. NOTES The flowers of  X. amplexicaulis, with their broad and thick outer petals, are among the largest in the entire genus, and were the basis for the original placement of the plant in the genus  Annonaby Lamarck, and the recognition of section  Pseudanonaby Baillon and later a genus of the same name by Safford. The auriculate clasping leaf bases are unique in the genus and unusual among Annonaceae.  Xylopia lamarckii, also endemic to Mauritius, has similar flowers but a distinctly petiolate leaf with a rounded base. Labels of the collections made by D. Lorence indicate that the branches are erect, forming a compact crown; such a growth habit is unusual within both the genus and family. Two collections of  Xylopia amplexicaulisin the Lamarck herbarium with handwritten labels state that they were collected by Commerson: one from Isle de France, the other from Madagascar. This agrees with the provenance information given in Lamarck’s protologue: “Cette plante croît à l’Isle de France& à Madagascar, où Commerson l’a observée”. Commerson apparently collected mostly around Taolagnaro (Fort Dauphin) during his brief visit to Madagascar( Morel 2012), and, as noted above, the plant has never been collected or reported from Madagascarsubsequently. This provenance is regarded as a labeling error. It is unclear whether the two locality labelings represent different collections, or the mis-labeling of some duplicates, but in either case, and to avoid future difficulties, a lectotypeis designated above. Lamarck’s diagnosis states “Anona foliis cordato-oblongis acutis distichis amplexicaulibus”. The leaves of the specimen marked as collected from “isle de france” have distinctly acute apices in addition to the cordate bases, while the leaves of the specimen marked as collected from “ Madagascar“ have cordate bases but rounded apices. Lamarck’s description carefully describes the sepals and petals of the flowers, which are only present on the “ Madagascar” specimen. This is also the more complete specimen, and is designated as the lectotype. There is uncertainty as to whether other collections are duplicates and thus isolectotypes, and they are therefore not listed as types. 2632071497 P Mauritius Madagascar 17 16 P030364 1 lectotype 2632071331 17 16 2632071315 17 16 2632071676 Bosser Per 17 16 1 2632071668 MAU Bouton 17 16 MAU0000986 1 2632071679 Drake Herb. & Herb. Richard, P Commerson Ile de France 17 16 1 2632071596 1950-03-21 MAU Duljeet F. D. National Reserves Crown Land Perrier 17 16 MAU0000961, MAU0000962 1 2632071767 17 16 2632071502 1998-12-08 MAU Florens -20.211111 Gaulettes 21 57.639725 17 16 MAU0000982 1 2632071711 17 16 2632071529 1974-03-05 MAU Gueho Perrier Nature Reserve 17 16 MAU0000984, MAU00000985 1 2632071377 1973-02-03 MO Lorence M Perrier Nature Reserve 549 Vacoas Mare 17 16 1 2632071423 1975-03-14 MAU Lorence Perrier Nature Reserve 17 16 MAU0000988 1 2632071516 1979-02-16 MAU, MO Lorence & Lecordier 300 Bel Ombre forest 17 16 MAU0000983 1 2632071401 1979-03-14 MO Lorence Perrier Nature Reserve 600 Vacoas Mare 17 16 1 2632071462 [904,1307,1107,1132] BM Wallich Grand Port 17 16 1 Grand Port