A revision of Homalium sect. Nisa (Salicaceae) endemic to Madagascar Wassel, Anna C. Applequist, Wendy L. Candollea 2020 2020-03-24 75 1 1 23 3MCJH Baker Baker [839,1139,417,442] Magnoliopsida Salicaceae Homalium Plantae Malpighiales 5 6 Tracheophyta species confertum     Lectotypus(designated by SLEUMER, 1973: 298):  MADAGASCAR:“ Central Madagascar”, s.d., fr., Baron 3185( K[ K000231489] image seen; isolecto-: BM, L[ L0010973] image seen, P[ P00418088]!).   Syntypus: MADAGASCAR:“Central Madagascar”, s.d., fr., Baron 3256( K[ K000231490] image seen).   Treeto 22 mtall, 60 cmdbh; bark exfoliating in plaques; twigs gray to medium brown, glabrous, leafing internodes (1.2–)1.7–4(–5)mm diam.; stipules fused, sometimes persistent. Leaveselliptic to oblong-elliptic, narrowly or broadly elliptic, or obovate, (5–)6– 16 × (1.8–)3– 6.2(–7) cm; base convex to cuneate; apex cuspidate, obtuse, acute, or rounded (shortacuminate); margin serrate to dentate or slightly crenate, rarely nearly entire, glands in tooth apices elongated and relatively conspicuous; secondary veins 8–14(–16) per side, sometimes with similar intersecondary veins; both surfaces glabrous (aberrantly bearing patches of short trichomes, possibly pathogenic), or abaxial surface moderately glaucous, drying medium to dark brown on both surfaces (rarely light green); petiole 3–7(–10) mm, glabrous. Inflorescencesracemose, axillary (often paired), (2–)4–8.5(– 11) cm; rachis glabrous (sparsely shortpubescent or more densely so in irregular patches); bracts and bracteoles (1.5–) 2–4 mm, glabrous except margins of bracts usually ciliate; flowers borne singly. Flowers5–6-merous, white to green or yellow-green; sepals 3–7 mmin flower, 8–12.5 mmin fruit, oblanceolate to spatulate or narrowly oblanceolateoblong, adaxial surface usually glabrous (midrib sometimes sparsely short-pubescent), abaxial surface usually glabrous (sparsely short-pubescent at base); sepal glands yellow, glabrous; petals ovate, 2–4 mm, both surfaces densely (to moderately) pubescent especially on apical half; filaments glabrous (sparsely pubescent on the basal portion), 1.7–3.1 mm; anthers (0.45–) 0.5–0.6 mm; styles 1.5–2.5 mm.   Vernacular names and use.– “Hazombato” ( Razafindramora 1, Ser vice Forestier 1776, 25277); “Hazombatofotsy ” ( Razanatsima et al. 1136, Service Forestier 7362, 16801, 25277); “Lalondalitra” ( Service Forestier 15406); “Longotra-fotsy” ( Service Forestier 1249); “Marankoditra” ( Réserves Naturelles 6043, Service Forestier 25277); “Tsianihiposa” ( Service Forestier 10148).  Wood is used to make unspecified products ( Service Forestier 25277).   Distribution, ecology and conservation status. – Homalium confertumis widespread in eastern and central regions of Madagascar. It usually occurs in humid forests at mid- to high elevations (range 445–1860 m), but also extends into relatively dry western forests (Ankarafantsika). Reported substrates include gneiss. It is sometimes common and locally dominant ( Randrianaivo et al. 526). It has been collected in several protected areas, including Ambohitantely, Anjanaharibe-Sud, Ankarafantsika, Montagne d’Ambre, and Zahamena. Its conservation status is estimated as “Least Concern” [LC].   Notes. – Homalium confertumwas lumped into  H. nudiflorumby SLEUMER (1973). It differs from  H. nudiflorum s.s.in its larger, often strongly toothed leaves with many marginal glands and its more densely pubescent petals; the final perianth size may also be larger. The two overlap in range, but  H. confertumis more tolerant of dry climate and high altitude, with much of its habitat on the central plateau. It is similar to  H. pachycladumand  H. retusum, which also overlap in range, in maximum leaf size, but  H. pachycladumand  retusumare distinguished by entire leaf margins, fewer leaf glands, and petals glabrous on the abaxial surface. Several possible hybrids between  H. confertumand  H. nudiflorumare seen in two regions where both are known, and gene flow is suspected to be responsible for some of the range of variation in  H. nudiflorum; these are discussed under the latter species. Hybridization is seen in most sections of  Homalium, even rarely among species of different sections ( APPLEQUIST, 2018b), so the ability of these species to hybridize does not suggest that they should not be recognized as distinct, given their significant morphological and ecological differences.   Additional material examined.–   MADAGASCAR. Reg. Alaotra-Mangoro[Prov. Toamasina]: PN Zahamena, 17°38'25"S 48°38'34"E,  989 m,  30.III.2000, buds,  Ratovosonet al. 184( MO, P);  Manaka, Ambatondrazaka,  20.II[?].1953, fl.,  Reserves Naturelles6043( MO);  P. K. 39 route d’Anosibe, canton and souspréfecture Anosibe,  4.V.1965, fr.,  Service Forestier25277( P).   Reg. Amoron’i Mania[Prov. Fianarantsoa]: commune d’Ambalamanakana, 20°44'01"S 47°11'42"E,  1860 m,  27.III.1996, fl.,  Rakotomalazaet al. 681A( MO, P).   Reg. Analamanga[Prov. Antananarivo]: Tampoketsa-Ankazobe, RS Ambohitantely, 18°11'52.5"S 47°17'3"E,  1620 m,  7.III.2004, fl.,  Almeda et al.8636( MO);  Anjozorobe, commune d’Ambongamarina, Andreba, 18°16'55"S 47°54'24"E,  1400 m,  11.III.2000, fl.,  Randrianaivoet al. 526( MO);  Ambohitantely RS, 18°11'46"S 47°17'24"E,  1620 m,  12.III.1997,  Ravololonanaharyet al. 69( MO).   Reg. Analanjirofo[Prov. Toamasina]: Beanana,  700 m,  22.II.1954, fl.,  Service Forestier9058( MO, P[2 sheets]).   Reg. Anosy[Prov. Toliara]: Antsotso Avaratra, forêt Tsitongambarika, 24°34'S 47°12'E,  2.IV.2008, fr.,  Razakamalalaet al. 4148( MO, P).   Reg. Atsinanana[Prov. Toamasina]: Ambodilendemy, Ankerana, 18°25'49"S 48°47'14"E,  890 m,  24.III.2011, fl.,  Antilahimena7778( MO);  Ambodilendemy, vers le sommet d’Ankerana, 18°25'49"S 48°47'14"E,  900m,  14.III.2011, fl.,  Ravelonarivo& Felix3663( MO);  Vatomandry, Ambalabe, 19°09'27"S 48°34'57"E,  596 m,  11.III.2013, fl.,  Razanatsimaet al. 1136( BR, G, K, MO, USMS).   Reg. Boeny[Prov. Mahajanga]: Réserve Naturelle Intégrale d’Ankarafantsika, 16°18'00"S 46°49'00"E,  22.IV.1994, fl.,  Rajemisa24( MO).   Reg. Bongolava[Prov. Antananarivo]: Tsiroanomandidy, Ambaravaranala, 18°25'S 45°40'E,  1190–1200 m,  25.VI.1997, fl.,  Rakotomalazaet al. 1353( G, MO).   Reg.DIANA [Prov. Antsiranana]: Montagne d’Ambre(partie N.O.),  1000–1800 m,  23.XII.1967, fr.,  Bernardi12003( MO, P);  contreforts occidentaux du Massif de Marojejy( NE) près du col de Doanyanala (limite des bassins de la Lokoho et de l’Andraronga),  800–1200 m,  25.I–25. II.1949, fl.,   Humbert23040( P[3 sheets]);  montagnes au N de Mangindrano(haute Maevarano) jusqu’aux sommets d’Ambohimirahavavy(partage des eaux Mahavavy-Androranga),  1500 m,  19.I–12.II.1951, fr.,   Humbert& Capuron24888( P);  flanc S du M. Tsaratanana,  1700 m,  XII.1912, fl.,  Perrier de la Bâthie6705( P).  Reg. Haute Matsiatra[Prov. Fianarantsoa]: Ampamaherana, Fianarantsoa,  13.XII.1949, fl.,  Service Forestier1249( P).   Reg.SAVA [Prov. Antsiranana]: Daraina, forêt d’Antsahabe, 13°13'03"S 49°32'46"E,  780 m,  2.XII.2004, fr.,  Gautier4806( MO, P);  Ampasindava, forêt de Bongomihirahavavy, 13°46'11"S 48°04'58"E,  445 m,  4.XII.2013, fr.,  Gautieret al. 6092( G, P);  massif de l’Anjanaharibe(pentes et sommet N) a l’W d’Andapa(haute Andramonta, bassin de la Lokoho: NE),  1600–1800 m,  10.XII.1950–  3.I.1951, fl.,   Humbertet al. 24704( P);  Doany, Anjialavahely, 14°17'41"S 49°32'50"E,  949 m,  15.III.2006, fl.,  Rakotovaoet al. 2986( MO, P);  Anjialavabe, Ankiakabe, 14°10'47"S 49°22'01"E,  1567 m,  23.II.2007, fl.,  Ravelonarivoet al. 2238( MO, P);  Antalaha, Bealampona, village de Befingotra, 14°45'11"S 049°28'49"E,  762 m,  21.IV.1997, fl.,  Razafindramora1( MO).   Reg. Sofia[Prov. Mahajanga]: Marsoandakana, 14°55'39"S 49°25'52"E,  1176 m,  23.II.2008, fl.,  Bernardet al. 876( MO);  NW du campement Bemafo, 14°13'17"S 49°03'55"E,  1830 m,  26.X.2005, fl.,  Buerkiet al. 106( MO, P).   Reg. Vakinankaratra[Prov. Antananarivo]: Angodona, Tsinjoarivo,  6.VI.1950, fl.,  Service Forestier1766( P[2 sheets]).   Reg. Vatovavy-Fitovinany[Prov. Fianarantsoa]: Ampasinambo, Nosy-Varika,  21.IV.1954, fl., fr.,  Service Forestier10148( MO, P). 3698440398 K, BM, L, P Madagascar Central Madagascar 5 6 K000231489, L0010973, P00418088 1 lectotype 3698440382 K Madagascar MADAGASCAR 5 6 K000231490 1 syntype 3698440323 2000-03-30 MO, P PN Zahamena & Ratovoson Madagascar 989 -17.640278 Reg. Alaotra-Mangoro 21 48.64278 7 8 1 Toamasina 3698440350 20. II [?]. 1953 MO Reserves Naturelles Madagascar Ambatondrazaka Manaka 7 8 1 3698440411 1965-05-04 P, K Service Forestier Madagascar canton and sousprefecture Anosibe 7 8 1 3698440480 1996-03-27 MO, P Rakotomalaza Madagascar 1860 -20.733612 commune d'Ambalamanakana 21 47.195 Reg. Amoron'i Mania 7 8 1 Fianarantsoa 3698440467 2004-03-07 MO Almeda et al. Madagascar 1620 -18.197916 Tampoketsa-Ankazobe 14 47.284164 Reg. Analamanga 7 8 1 Antananarivo 3698440442 2000-03-11 MO Randrianaivo Madagascar Anjozorobe 1400 -18.281944 Andreba 21 47.90667 d'Ambongamarina 7 8 1 3698440424 1997-03-12 MO Ravololonanahary Madagascar 1620 -18.19611 Ambohitantely RS 21 47.29 7 8 1 3698440469 1954-02-22 MO, P Service Forestier Madagascar 700 Beanana Reg. Analanjirofo 7 8 1 Toamasina 3698440380 2008-04-02 MO, P Razakamalala Madagascar -24.566668 Antsotso Avaratra 1250 47.2 Reg. Anosy 7 8 1 Toliara 3698440482 2011-03-24 MO Antilahimena Madagascar Reg. Atsinanana 890 -18.430277 Ankerana 21 48.78722 Ambodilendemy 7 8 1 Toamasina 3698440329 2011-03-14 MO Ravelonarivo & Felix Madagascar 900 -18.430277 vers le sommet d'Ankerana 21 48.78722 Ambodilendemy 7 8 1 3698440444 2013-03-11 BR, G, K, MO, USMS Razanatsima Madagascar 596 -19.1575 Ambalabe 21 48.5825 Vatomandry 7 8 1 3698440315 1994-04-22 MO Rajemisa Madagascar -16.3 Reserve Naturelle Integrale d'Ankarafantsika 21 46.816666 Reg. Boeny 7 8 1 Mahajanga 3698440431 1997-06-25 1949-02-25 1949-01-25 G, MO Rakotomalaza Madagascar Reg. 1195 -18.416666 Tsiroanomandidy 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440333 1967-12-23 1949-02-25 1949-01-25 MO, P Montagne d'Ambre & Bernardi Madagascar Reg. 1400 -18.416666 Reg. 1275 45.666668 Reg. Bongolava 7 8 1 Antsiranana 3698440347 1949-01-25 1949-02-25 1949-01-25 NE, P Humbert Madagascar Reg. 1000 -18.416666 contreforts occidentaux du Massif de Marojejy 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440354 1951-01-19 1951-02-12 1951-01-19 P Humbert & Capuron Madagascar Reg. 1500 -18.416666 jusqu'aux sommets d'Ambohimirahavavy 1275 45.666668 Maevarano 7 8 1 Antananarivo 3698440340 1912-12 1949-12-13 1912-12 P Service Forestier Madagascar Reg. 1700 -18.416666 Reg. Haute Matsiatra 1275 45.666668 Reg. Bongolava 7 8 1 Fianarantsoa 3698440474 2004-12-02 1949-02-25 1949-01-25 MO, P Gautier Madagascar Reg. 780 -18.416666 foret d'Antsahabe 1275 45.666668 Daraina 7 8 1 Antsiranana 3698440438 2013-12-04 1949-02-25 1949-01-25 G, P Gautier Madagascar Reg. 445 -18.416666 Ampasindava 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440407 1950-12-10 1951-01-03 1950-12-10 P Humbert Madagascar Reg. 1700 -18.416666 massif de l'Anjanaharibe 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440473 2006-03-15 1949-02-25 1949-01-25 MO, P Rakotovao Madagascar Reg. 949 -18.416666 Anjialavahely 1275 45.666668 Doany 7 8 1 Antananarivo 3698440339 2007-02-23 1949-02-25 1949-01-25 MO, P Ravelonarivo Madagascar Reg. 1567 -18.416666 Ankiakabe 1275 45.666668 Anjialavabe 7 8 1 Antananarivo 3698440400 1997-04-21 1949-02-25 1949-01-25 MO Razafindramora Madagascar Reg. 762 -18.416666 Bealampona 1275 45.666668 Antalaha 7 8 1 Antananarivo 3698440440 2008-02-23 1949-02-25 1949-01-25 MO Bernard Madagascar Reg. 1176 -18.416666 Marsoandakana 1275 45.666668 Reg. Sofia 7 8 1 Mahajanga 3698440370 2005-10-26 1949-02-25 1949-01-25 MO, P Buerki Madagascar Reg. 1830 -18.416666 Tsiroanomandidy 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440413 1950-06-06 1949-02-25 1949-01-25 P Service Forestier Madagascar Reg. 1195 -18.416666 Reg. Vakinankaratra 1275 45.666668 Reg. Bongolava 7 8 1 Antananarivo 3698440365 1954-04-21 1949-02-25 1949-01-25 MO, P Service Forestier Madagascar Reg. 1195 -18.416666 Ampasinambo 1275 45.666668 Reg. Vatovavy-Fitovinany 7 8 1 Fianarantsoa