Paracephaelis cinerea (A.Rich.) De Block

Block, Petra De, 2022, Monograph of the western Indian Ocean genus Paracephaelis (Rubiaceae - Pavetteae), with description of thirteen new species, European Journal of Taxonomy 801, pp. 1-99 : 31-38

Paracephaelis cinerea (A.Rich.) De Block

Figs 1A, E, I , 2I , 8

Systematics and Geography of Plants 73: 100 ( De Block 2003).

Basionym

Pavetta cinerea A.Rich. in DC., Prodromus Systematis Naturalis Regni Vegetabilis 4: 492 ( De Candolle 1830); Mémoire sur la Famille des Rubiacées: 101 ( Richard 1830); Mémoires de la Société d’Histoire naturelle de Paris 5: 181 (Richard 1834). Type: MADAGASCAR – Antananarivo Province, Analamanga Region, Antananarivo Renivohitra District • in monte Tananarivo ; s.dat.; fl; A. Richard in Drake s.n.; holotype: P[ P00115278 ].

Homotypic synonyms

Ixora cinerea (A.Rich) Drake, Histoire physique, naturelle, et politique de Madagascar 36: pl. 421 (Drake del Castillo 1897); Bulletin mensuel de la Société linnéenne de Paris 2: 1308 (Drake del Castillo 1898); Annuaire du Conservatoire et du Jardin botaniques de Genève 11–12: 107, fig. 14 ( Hochreutiner 1908). – Chomelia cinerea (A.Rich.) Dubard & Dop , Journal de Botanique (Morot), sér. 2, 3: 11 ( Dubard & Dop 1925). – Tarenna cinerea (A.Rich.) Bremek. , nom. illeg., Repertorium Novarum Specierum Regni Vegetabilis 37: 7, 189 ( Bremekamp 1934; see Note 2); Bulletin de la Société botanique de France 85: 607 ( Homolle 1938); Révision des Rubiacées de Madagascar et des Comores: 170 ( Capuron 1973). – Tarenna spodia Bremek., Repertorium Novarum Specierum Regni Vegetabilis 47: 96 ( Bremekamp 1939).

Heterotypic synonym

Webera hispidula Baker , Journal of Botany (London) 20: 137 ( Baker 1882). Type: MADAGASCAR – Fianarantsoa Province • chiefly in Betsileo land; s.dat.; fl; Baron 150; holotype: K, isotype: P.

Other collections examined

MADAGASCAR – Antsiranana Province, Diana Region , Antsiranana II District • montagne des Français, 4 km from the junction of the main road to Ramena; 78 m a.s.l.; 21 Jul. 2007; fr; Andriamahay & Rakotoarisoa 1794; K • Analamera RS, 80 km S d’Antsiranana par route et 2.5 km E du village d’Ambery (Antranonaomby); 507 m a.s.l.; 17 Jun. 1994; fr; Andrianantoanina & Rabeharinosy 705; BR, MO • montagne Andrao à Lingvatou [Lanivato]; s.dat.; fl; Bernier 319; BR, P • forêt d’Orangea , E de Diego- Suarez; 22 Feb. 1962; fl; Capuron 20942-SF; BR, P, TEF • forêt de Sahafary , bassin de la Saharaina; 200 m a.s.l.; 20 Feb. 1962; fl; Capuron 20969-SF; BR, P, TEF • lisière supérieure de la forêt d’Andranomadiro, rebord S du plateau de Sahafary , entre les bassins de la Saharenana et de l’Irodo; 300 m a.s.l.; 27 Dec. 1963; fl, fr; Capuron 23073-SF; P, TEF • environs d’Antsoha , S du mont Raynaud, piste de la baie de Rigny ; 28 Feb. 1964; fl; Capuron 23293-SF; BR, P, TEF • rebord S du plateau de Sahafary, haut de la forêt d’Andranomadiro ; 26 Feb. 1964; fr; Capuron 23309-SF; BR, P, TEF • Analamera, along Ambtabe river ; 40 m a.s.l.; 7 Jan. 2002; fl; De Block, Rakotonasolo & Randriamboavonjy 1111; BR, MO, P, TAN, WAG • Analamera, bank of Irodo river , close to Irodo camp; 40 m a.s.l.; 8 Jan. 2002; fl; De Block, Rakotonasolo & Randriamboavonjy 1133; BR, G, MO, TAN, UPS • ibid.; 10 Jan. 2002; fl; De Block, Rakotonasolo & Randriamboavonjy 1174; BR, K, MO, P, TAN • baie de Sakalava ; 0–100 m a.s.l.; 19 Jan. 2002; fl; De Block, Rakotonasolo & Randriamboavonjy 1291; BR, K, MO, P, TAN, WAG • forêt d’Orangea, side closest to Ramena ; 0–100 m a.s.l.; 24 Jan. 2002; fl; De Block, Rakotonasolo & Randriamboavonjy 1384; BR, G, K, MO, TAN • commune Mahavanona , fkt. Andranomanitra , Antafiankoroka , massif forestier de la montagne des Français ; 204 m a.s.l.; 4 Feb. 2005; fl; Rabefarihy 52; BR, MO, P, TAN • montagne des Français ; 12 Feb. 2003; fr; Rakotonasolo & Rogers 582; P, K • montagne des Français, road to Ivovona ; 12 Feb. 2003; fr; Rakotonasolo & Rogers 585; BR, K • Andrafiabe , Ambolobozokely , Anosy , 4 km NE du village d’Ambolobozokely; 22 m a.s.l.; 30 Mar. 2007; fr; Rakotondrafara 497; BR, CNARP n.v., MO, P, TAN • commune Sadjoavato, fkt. Saharenana, forêt d’Andranomadiro, SW of Sahafary , S-facing watershed draining plateau; 300 m a.s.l.; 14 Feb. 2005; fr; Schatz, Ratovoson & Riri Guittou 4274; BR, CNARP n.v., MO, P, TAN • commune Sadjoavato, fkt. Saharenana, forêt de Sahafary; 210 m a.s.l.; 15 Feb. 2005; fl; Schatz, Ratovoson & Guittou 4299; CNARP n.v., MO n.v., P online, TAN n.v. – Nossi-Be District • Nossi-Be, bords de la mer entre Amponbilava et Djabul ; Feb. 1851; fl; Boivin s.n.; P • Nossi-Komba; Dec. 1879; fl; Hildebrandt 3227; BM, G, K, P. – Sava Region, Vohemar District • Port Leven ; 1849; fl, fr; Boivin 2430; P • commune Daraina, Daraina, Bekaraoka-Sud ; 9Apr. 2008; fr; Bremer, Mouly, Ravelonarivo & Kainulainen 5122; S • commune Daraina, Daraina , forêt de Solaniampilana-Maroadabo; 12 Apr. 2008; fr; Bremer, Mouly, Kainulainen & Mariano 5161; S • forêt d’Andaingo, S de la rivière d’Andripatra, N de Vohemar ; 16 Mar. 1967; fl; Capuron 27443-SF; BR, P, TEF • vestige forestier d’Ambatrabe, S de Maintialaka, N de Vohemar ; 15 Mar. 1967; fl; Capuron 27458-SF; BR, P, TEF • S de Vohemar ; Mar. 1967; fl; Capuron 27477-SF; P, TEF • commune Daraina , Daraina , forêt d’Ankaramy , à 910 m du point côté 361, au 203°; 200 m a.s.l.; 22 Feb. 2004; fl; Nusbaumer & Ranirison 1201; BR, G n.v . • commune Nosibe, lieu de campement au lac Sahaka, près de la forêt d’Analabe ; 22 Feb. 2003; fl; Rabevohitra, Rabenantoandro & Razakamalala 4479; MO, P • commune Nosibe, Anjiabe, forêt littorale d’Analabe ; 24 Feb. 2003; fl; Rabevohitra, Rabenantoandro & Razakamalala 4531; MO, P • Maromikotro-Iloky , road to Daraina; 21 Feb. 2013; fl; Rakotonasolo, Randriamboavonjy, Letsara, Bone & Andriantiana 2143; BR, K, MO, P, TAN • commune Daraina, Daraina , forêt de Bekaroaka , partie nord , à 420 m du point côté 96, au 133°; 180 m a.s.l.; 10 Feb. 2004; fl; Ranirison & Nusbaumer 401; BR, G n.v . • commune Daraina , Daraina , forêt d’Ankaramy , à 1160 m du point côté 361, au 171°; 27 Feb. 2004; fl; Ranirison & Nusbaumer 491; BR, G n.v . • commune Daraina , Daraina , forêt d’Antsaharaingy , à 1080 m du point côté 41, au 303°; 42 m a.s.l.; 18 Apr. 2004; fr; Ranirison 685; BR, G n.v . • commune Nosibe , forêt littorale d’Analabe près du village d’Anaborano et du lac Sahaka; 21 m a.s.l.; 12 Jul. 2003; fr; Razakamalala, Rabevehitra & Rakotomamonjy 577; MO n.v., P online • fir. Tsarabaria , fkt. Manakana , forêt sublittorale E du village d’Ambondrobe; 13 Mar. 2004; fl; Razakamalala, Rabevohitra & Faralahy 993; BR, K, MO • fir. Tsarabaria, fkt. Manakana, forêt littorale d’Ambondrobe ; 20 May 2004; fr; Razakamalala 1317; MO n.v., P online • Vohemar ; 1840; fr; Richard s.n.; P • Vohemar; s.dat.; fr; Richard in Drake 47; P • aux environs de Vohemar ; s.dat.; fr; Richard 83; L, P • Vohemar ; 1840; fr; Richard 649; P • Port Leven; 1850; fl, fr; Vesco s.n.; P. – Toamasina Province, Alaotra-Mangoro Region, Ambatondrazaka District • lac Alaotra; s.dat.; fr; coll. ignot. (Herbier Jardin botanique de Tananarive) 3832 (MEN-75); P • lac Alaotra; s.dat.; fl; coll. ignot. (Herbier Jardin botanique de Tananarive) 3878 (ANK-206) ; P • lac Alaotra ; s.dat.; fl; coll. ignot. (Herbier Jardin botanique de Tananarive) 3923 (MEN-6) ; P • forêt d’Analamihilana ; 850 m a.s.l.; 27 Dec. 1944; fl; Cours 1996; BR, P • Antandrokomby; Jun. 1957; fl; Peltier J. & M. 964; P, TAN. – Andilamena District • fkt. Ampamoho, forêt d’Analalava, 10 km S d’Andilamena; 1006 m a.s.l.; 8 Dec. 2001; fl; Randrianaivo, Ratodimana, Razafindrabeaza, Razanatsoa, Rakotozanahary, Rabeabimanana & Tolozafy 814; K, MO, P.– Moramanga District • on the left of Mangoro river; 920 m a.s.l.; 13 Jan. 2001; fl; Rakotonasolo 285; BR, K. – Mahajanga Province, Betsiboka Region, Maevatanana District • Ampanibe, près du confluent de l’Ikopa et du Betsiboka; Feb. 1899; fl; Perrier de la Bâthie 828; P. – Tsaratanana District • bord N du sentier vers Antaminbaribe, forêt d’Ampasindava, village le plus proche Ampasindava; 28 Apr. 1958; fr; Razafinarivo 19142-SF; BR, P, TEF. – Boeny Region, Ambato-Boeni District • RNI 7, Ankarafantsika, Bevasaka ; 100 m a.s.l.; Mar. 1933; fl; coll. ignot. 144-SF; P • forêt de Maroaboaly , village le plus proche Antafia , fir. Sitampiky ; 8 Mar. 1957; fr; coll. ignot. 17762-SF; P, TEF • Ampijoroa forestry station, forest across from lake Ravelobe ; 14 Feb. 1999; fl; De Block & Rakotonasolo 813; BR, MO, P, TAN • forêt d’Ankarokaroka, village le plus proche Antafia, fir. Sitampiky ; 21 Jan. 1958; fl; Tefamilia 19322-SF; TEF • RNI 7, Ankarafantsika, Tsaramandroso; 28 Sep. 1951; fr; Tsilizy 2974-RN; BR, P, TEF. – Mahajanga I District • environs de Majunga; 2–15 m a.s.l.; Dec. 1924; fl; Humbert 4048; BR, P • dunes de Majunga; Jan. 1921; fl; Perrier de la Bâthie 13459; BR, P. – Mahajanga II District • Mariarano, Ankatsabe, forêt d’Ankatsabe; 16 m a.s.l.; 20 Apr. 2007; fr; Rakotoarivelo, Miandrimanana, Randrianarivelo, Razakamalala & Beby 88; MO n.v., P online, TAN n.v. – Mitsinjo District • Tsiombikibo; 24 m a.s.l.; 6 May 2005; fr; Andriamahay & Rakotoarisoa 1055; K. – Soalala District • RNI 8, Namoroka, ca 38.5 km S of Soalala ; 130 m a.s.l.; 3 Feb. 2000; fl; Davis, Rakotonasolo & Wilkin 2534; BR, K, P, TAN • ca 3 km S of Soalala ; 5 m a.s.l.; 8 Feb. 2000; fl; Davis & Rakotonasolo 2546; BR, K, P, TAN • Soalala ; 12 Jul. 1977; fr; Rakotozafy 1925; TAN • fir. Andranomavo; 24 Feb. 1956; fl; Randriamiero 8646-RN; BR, P, TAN, TEF (as coll. ignot. 17476- SF). – Melaky Region, Antsalova District • Bemaraha; Aug. 1943; fr; coll. ignot. (Herbier Jardin botanique de Tananarive) 6191; P. – Maintirano District • forêt Analalava, 1600 m NE du campement Mokotra, fir. Maintirano; 24 Feb. 1955; fr; coll. ignot. 14803-SF; P, TEF • forêt d’Amboloando, village le plus proche Amboloando, fir. Maintirano; 30 May 1956; fr; coll. ignot. 16391-SF; BR, P, TEF. – Sofia Region, Antsohihy District • Anjimangirana , 3 km W of Anjimangirana (Analananbe) village; 290 m a.s.l.; 30 May 2000; fr; Rakotonasolo 172; BR, K, P, TAN • Anjimangirana , Analananbe, Andrafiborizina; 285 m a.s.l.; 30 May 2000; fr; Rakotonasolo 186; BR, K, TAN • Ankerika, Andohanakerika; 200 m a.s.l.; 3 Jun. 2000; fr; Rakotonasolo 199; BR, K, TAN. – Boriziny-Vaovao District • N of Port Bergé ; 70 m a.s.l.; 18 Jan. 1988; fl; Bisset M52; K • Mandritsara; Apr. 1974; fr; Morat 4469; P. – Mampikony District • Bongolava, Ampombimanangy, Bongolava forest , 6 km W of Ampombimanangy; 221 m a.s.l.; 14 Jun. 2008; fr; Andriamahay & Rakotoarisoa 2047; K. – Mandritsara District • Ambalafarihy, village le plus proche Ambalafarihy, fir. Antsirabe; 14 Mar. 1966; fr; Sajy 25847-SF; P, TEF. – Antananarivo Province, Analamanga Region, Andramasina District • Andramasina, Ambatolampy; Dec. 1958; fl; Bosser 12461; TAN. – Anjozorobe District • Analabe, N d’Antananarivo; Mar. 1953; fr; Bosser 5111; P • Analabe centre; Mar. 1953; fr; Bosser 5229; TAN • Anjozorobe, by the side of Mananara river; 23 Dec. 1999; fl; Rakotonasolo 119; BR, K, TAN. – Ankazobe District • Tampoketsa d’Ankazobe ; 29 Apr. 1943; fr; Decary 19311; BR, P • 30 km N of Ankazobe, Tampoketsa d’Ankazobe ; 1600 m a.s.l.; 18 May 1974; fr; Gentry 11822; MO, P, TAN • plateau du Tampoketsa entre la Mahazemba et le Bemarivo; Jun. 1905; fr; Perrier de la Bâthie 3626; P • Tampoketsa d’Ankazobe; 1500 m a.s.l.; s.dat.; fr; Perrier de la Bâthie 17315; P. – Antananarivo Renivohitra District • Antananarivo; s.dat.; fl; Hilsenberg & Bojer s.n.; BM • in monte Tananarivo; s. dat.; fl; Richard in Drake s.n.; P. – Manjakandriana District • forêt de la Mandraka; s.dat.; fl; d’Alleizette 10; P. – Itasy Region, Arivonimamo District • Antongona, environs d’Antananarivo ; Jan. 1956; fl; Bosser 8946; TAN • Antongona ; Jan. 1960; fl; Bosser 13460; TAN • Antongona , rochers à 40 km W d’Antananarivo; 1500 m a.s.l.; 16 Jan. 1960; fl; Leandri 2591; P • Antongona; 16 Jan. 1960; fl; Peltier J. & M. 1752; P, TAN. – Miarinarivo District • lac Itasy; Feb. 1964; fl, fr; Bosser 19183; P. – Vakinankaratra Region, Ambatolampy District • PK 35 at stream-crossing S of Behenjy on road from Antananarivo to Antsirabe ( RN 7 ); 1300–1400 m a.s.l.; 12 Jan. 1986; fl; Dorr, Barnett & Rakotozafy 4554; K, MO, P. – Antanifotsy District • Ambohimandroso, près d’Ambatolampy, bords de l’Onive; 1500 m a.s.l.; Dec. 1955; fl; Bosser 8781; P, TAN. – Antsirabe II District • Antsirabe; 1600 m a.s.l.; Jan. 1914; fl; Perrier de la Bâthie 3508; BR, P. – Betafo District • Mandoto; 6 Dec. 1939; fl; Decary 15245; BR, P. – Faratsiho District • Ramainandro, Ankaratra; Jan. 1955; fl; Bosser 7587; P, TAN. – Fianarantsoa Province, Ihorombe Region, Ihosy District • entrée d’Ihosy , face à la piste d’aviation; 570–631 m a.s.l.; 3 Jan. 1999; fl; Allorge 2236; P • bassin de la Sahambana , SE d’Ihosy, PK 21 de la route Ihosy-Ivohibe; 9 Feb. 1963; fr; Capuron 22606-SF; BR, K, P, TEF • Isalo PN , canyon des makis ; 9 Jan. 1999; fl; De Block, Rakotonasolo & Randriamboavonjy 586A; BR, K, MO, P, TAN • Isalo PN, beyond canyon des makis; 9 Jan. 1999; fr; De Block, Rakotonasolo & Randriamboavonjy 586B; BR, K, MO, TAN • road from Ihosy to Farafangana, 9 km before Ihosy, PK 26; 745 m a.s.l.; 30 Jan. 2006; fl; De Block, Tosh & Rakotonasolo 1942; BR, K, MO, TAN, UPS • environs d’Ihosy ; 1 Feb. 1957; fl; Descoings 2191; P, TAN • inselberg on RN 7 to Tulear; 22°10′12.9″ S, 46°22′39.4″ E; 812 m a.s.l.; 30 Jan. 2007; fl; Groeninckx, Rakotonasolo, Dessein & De Block 162; BR, MO, P, TAN, UPS • Ranohira ; s.dat.; fr; Homolle 1463; K, P • environs d’Ihosy ; 800–900 m a.s.l.; Mar. 1934; fr; Humbert 14471; P • haute vallée de la Menarahaka , E d’Ihosy ; 700–800 m a.s.l.; 28 Jan.–10 Apr. 1955; fl; Humbert 28554; BR, P • Ranohira, E d’Isalo; 750–800 m a.s.l.; Feb. 1955; fr; Humbert 28795; P. – Amoron’i Mania Region, Ambatofinandrahana District • Analandratehina, route vers Ambatomenaloha; 1615 m a.s.l.; 27 Apr. 2012; fr; Andrianaivoravelona, Ravaoarisoa, Lazasoa, Ralaivao & Manampisoa 613; BR, K, MO, P, TAN • environs d’Ambatofinandrahana ( Betsileo ); 1400–1500 m a.s.l.; 16 Jan. 1955; fr; Humbert & Capuron 28091; P • Itremo, gallery forest in Analamaizina ; 31 Nov. 2009; fl; Rakotonasolo, Moat, Rakotoarinivo, Rakotoarisoa, Bachman & Couch 1520; BR, K, P, TAN • Analabe, ca 1 km SE of Analabebiby; 16 Dec. 2011; fl; Randriamboavonjy, Rajaovelona & Andriamilanto 1011; BR, K, MO, P, TAN. – Ambositra District • Faliarivo, W d’Ambositra; 1600 m a.s.l.; 15 Jan. 1955; fl; Humbert & Capuron 28023; P. – Haute Matsiatra Region, Ambalavao District • RNI 5, Andringitra, Sendrisoa; 19 Dec. 1952; fl; Rakoto 4823-RN; P, TAN, TEF. – Toliara Province, Anosy Region, Amboasary-Atsimo District • forêt d’Ankirikirika (Ankiritry); 25 Jan. 1945; fl; Homolle E 1; P. – Betroka District • vallée de la Manambolo, bassin du Mandrare, au confluent de la Sakamalio, mont Morahariva; 1000–1200 m a.s.l.; Dec. 1933; fl; Humbert 13145; BR, P. – Taolagnaro District • RN 10, 16 km W of Manambaro ; 110 m a.s.l.; 21 Feb. 1975; fr; Croat 31950; K, MO, P, TAN • RNI 11, Andohahela , Fort-Dauphin , parcelle 3 , Ankazofotsy ; 100–250 m a.s.l.; 21–23 Jun. 1994; fr; Eboroke 823; BR, K, MO, P • Ambatoabo, Mahamavo , Ankazomena , subhumid forest of Andohahela ; 660 m a.s.l.; 25 Apr. 2009; fr; Rakotonasolo, Ratovoson, Iso Fiadana, Rasolondrainy & Constant Mbola 1244; BR, MO, P • RNI 11, Andohahela , parcelle 3 ; 100–300 m a.s.l.; 18 Feb. 1993; fr; Randriamampionona 124; BR, K, MO, P, WAG • RNI 11 , Andohahela , parcelle 3 ; 100–300 m a.s.l.; 8–10 Apr. 1993; fr; Randriamampionona 269; BR, K, MO, P, UPS, WAG • Andohahela, parcelle 3, Ankazofotsy; 15 May 2001; fr; Randriamampionona 1001; BR, CNARP n.v., MO, P, TAN. – Atsimo-Andrefana Region, Beroroha District • fkt. Beronono-Makay, déviation canyon vers le plateau Behetaheta; 531 m a.s.l.; 16 Jan. 2010; fl; Rakotovao, Razakamalala & Andriantiana 5105; MO n.v., P online, TAN n.v . • fkt. Antsoa, village le plus proche Antanamary, Androsy; 254 m a.s.l.; 9 Jan. 2011; fl; Razakamalala 5892; BR, MO, P, TAN. – Betioky District • Manasoa Tanosy ; 14 Jan. 1913; fl; Afzelius s.n.; P, S • Manasoa Tanosy ; 14 Jan. 1913; fl; Afzelius 258; S • Manasoa Tanosy ; 14 Jan. 1913; fl; Afzelius 264; S • road from Betioky to Ampanihy; 27 km from Betioky; 335 m a.s.l.; 6 Feb. 2007; fl; De Block, Dessein, Groeninckx & Rakotonasolo 2317; BR, MO, BR, TAN. – Morombe District • N of Toliara, forêt de Mikea, 23–25 road-km W of Vorehy ; 50 m a.s.l.; 12 Feb. 1998; fr; McPherson & Razafimandimbison 17411; K, MO • forêt de Mikea axe Ankilimihavotse-Ankindranoke, Basibasy; 0–50 m a.s.l.; 1 Feb. 2000; fl; Ranaivojaona et al. 308; MO n.v., P online, TAN n.v. – Sakaraha District • Mahaboboka, Marotsiraka, forêt proche du village d’Ambinanitelo ; 447 m a.s.l.; 20 Dec. 2010; fl; Andriamihajarivo, Razanatsima & Fagnarena Miandry 1828; P online • Mahaboboka , Marotsiraka , partie NW de la forêt d’Analavelona, près de la source, W de la rivière de Manasay; 1293 m a.s.l.; 16 Jan. 2012; fl; Andriamihajarivo & Fagnarea 1833; BR, MO, P, TAN • Mahaboboka , Marotsiraka Betsileo , village le plus proche Andombiry, forêt d’Analavelona le long de la rivière Manasay; 997 m a.s.l.; 1 Dec. 2016; fl; Andriamihajarivo, Rehary & Bruno 1992; MO n.v., P online, TAN n.v . • Sakaraha ; Feb. 1956; fr; Bosser 9138; BR, P, TAN • bassin de la Mananadabo dans le massif de l’Analavelona, N du Fiherenana ; 1000–1300 m a.s.l.; 13–15 Dec. 1962; fl; Capuron 22200-SF; BR, P, TEF • Lambomakandro ; 3 Mar. 1943; fl, fr; Decary 18896; BR, P • Zombitse-Isoky , Zombitse PN ; 612 m a.s.l.; 1 Feb. 2007; fr; Groeninckx, Rakotonasolo, Dessein & De Block 177; BR, G, MO, P, TAN • Mahaboboka , fkt. Marotsiraka Betsileo , forêt de Marokobay NE du village d’Ambinanintelo; 462 m a.s.l.; 20 Feb. 2011; fr; Randrianarivony, Rakotoarivony, Helison, Heriman & Rehary 245; MO n.v., P online, TAN n.v . • commune Mahaboboka , fkt. Marotsiraka Betsileo , Analavelona , forêt de Manasay ; 1335 m a.s.l.; 18 Jan. 2012; fl; Randrianarivony, Ramarosandratana, Andriamihajarivo & Makabe 411; MO n.v., P online, TAN n.v . • Mahaboboka, Ambinanintelo, forêt d’Analavelona, forêt d’Andohaniankokoky au bord de la rivière Ankokoky; 970 m a.s.l.; 5 Dec. 2015; fl; Randrianarivony, Rakotoarivony, Helison, Heriman, Rehary & Rebesa 801; MO n.v., P online, TAN n.v. – Menabe Region, Morondava District • Kirindy forest, Andranomena RS; 17 m a.s.l.; 21 Feb. 2018; fl; Atalahy, Razafindrahaja, Swenson & Razafimandimbison 115; P online, S n.v . • forêt de Kirindi, CFPF Morondava (forêt d’Andalandahalo), jardin botanique 2, ca 45 km NE of Morondava; 15 m a.s.l.; 20 Feb. 2000; fr; Davis, Rakotonasolo & Wilkin 2570; BR, K, P, TAN • Andranomena forest , ca 17 km before turn-off to Kirindi; 16 m a.s.l.; 18 Jan. 2007; fl; De Block, Rakotonasolo, Groeninckx & Dessein 2168; BR, MO, TAN • Kirindy forest , N part - Conoco 7; 16 m a.s.l.; 19 Jan. 2007; fl; De Block, Rakotonasolo, Groeninckx & Dessein 2193; BR, G, MO, P, TAN • RN 35 at Antsehase bridge/river; 197 m a.s.l.; 23 Jan. 2007; fr; De Block, Dessein, Groeninckx & Rakotonasolo 2244; BR, MO, P, TAN, UPS, WAG • Lamboukily , 14 km from camp de base in Kirindi; 42 m a.s.l.; 20 Jan. 2007; fl; Groeninckx, Rakotonasolo, Dessein & De Block 105; BR, G, MO, P, TAN, WAG • RN 35, ca 22 km from Morondava on road to Antananarivo, dunes des Belges; 19 m a.s.l.; 24 Jan. 2007; fl; Groeninckx, Rakotonasolo, Dessein & De Block 149; BR MO, P, TAN, WAG • commune Bemanonga, Andranomena Reserve ; 18 Mar. 2011; fl, fr; Kainulainen, Razafimandimbison, Razafindraibe & Wikström 170; BR, S • commune Bemanonga , fkt. Marofandilia , village le plus proche Kirindy, forêt de Kirindy ; 57 m a.s.l.; 12 Apr. 2013; fr; Razakamalala, Randrianaivo & Rakotondrasaona 7404; MO n.v., P online, TAN n.v . • commune Bemanonga, fkt. Marofandilia, village le plus proche Kirindy, forêt de Kirindy; 59 m a.s.l.; 21 Apr. 2013; fr; Razakamalala, Randrianaivo & Rakotondrasaona 7505; MO n.v., P online, TAN n.v. – Without locality • chiefly in Betsileo land; 1890; fl; Baron 150; K, P • s.dat.; fl; Baron 4596; BM, K, P • 1840; fl; Bernier s.n.; K • N de Madagascar; 1835; fl; Bernier in Guillemin 319; P • côte orientale; 1853; fl, fr; Boivin in Barbey-Boissier s.n.; G • s.dat.; fr; Boivin in Cosson 2430; P • s.dat.; fl; d’Alleizette s.n.; P • s.dat.; fr; Homolle 1419; BR, K, P .

Description

Shrub or small tree, 1–5(–7) m tall; pubescence whitish to tawny; leaves sometimes caducous. Young shoots brown or dark brown but rapidly becoming corky and then fawnish or orange brown, rarely glabrous but usually sparsely to densely covered with appressed or erect hairs; older branches corky, grey, greyish brown or reddish brown, somewhat flaking, glabrescent or glabrous. Petioles 4–10(–15) mm long, rarely glabrous, usually sparsely to densely covered with appressed or erect hairs. Leaf blades broadly obovate, broadly elliptic, orbicular, obovate, elliptic, ovate, rarely narrowly elliptic, 2–12 × 1–8.5 cm, subcoriaceous, drying dark or blackish brown above, somewhat paler below or not discolorous, upper surface glossy, moderately to densely covered with minute to short erect hairs sometimes combined with sparse long appressed hairs associated with higher order venation, or, glabrous but then usually at least midrib or midrib and secondary veins pubescent, lower surface moderately to densely covered with long erect hairs to woolly or glabrous except for midrib and secondary veins; margin ciliate; base cuneate, obtuse, rounded, rarely truncate, subcordate or somewhat unequal; tip rounded, obtuse, acute, more rarely acuminate or retuse; hair tuft domatia often present; 5–10 secondary veins on each side of midrib, impressed above and raised below; midrib and secondary veins densely to sparsely covered with long appressed hairs on both surfaces or rarely glabrous; higher order venation visible on both surfaces. Stipules triangular with needle-like awn, often somewhat keeled, outer surface rarely glabrous but usually with at least a few appressed hairs along the base and/or along the midrib, or, moderately to densely covered with appressed hairs all over, inner surface glabrous with row of colleters at the base; sheath 1.5–4 mm long; awn 0.5–3 mm long. Inflorescences sessile but leaves of first order bracts often fallen (giving the impression of short peduncle), 1.5–5 × 2–8 cm, with (15–)25–numerous flowers; inflorescence parts densely covered with ± long appressed, spreading or erect hairs; first order axes 0.3–3 cm long; higher order bracts often displaced 1–4 mm above branching point, with stipular parts reduced or absent, foliar parts narrowly triangular and vaulted or more rarely ovate, 2–4 mm long; bracteoles at base of ovary and opposite or on pedicel somewhat below ovary and subopposite, stipular parts absent, foliar parts narrowly triangular to linear, ca 1(–1.5) mm long, tips acute. Flowers sessile to shortly pedicellate, pedicels 0–2 mm long. Calyx densely covered with appressed, spreading or erect hairs outside, inner surface densely covered with appressed hairs; tube 0.2–0.5 mm long; lobes triangular, 0.5–1 mm long, tips acute. Corolla tube 5–7.5 mm long, inner surface moderately covered with long erect hairs in upper half; lobes 3–4.5 mm long. Anthers 3–4 mm long, completely exserted from corolla tube at anthesis; filaments ca 1 mm long; connective often drying reddish brown. Ovary ca 1 mm long, densely covered with appressed, spreading or erect hairs; per locule 1–4 ovules pendulous from small placenta attached to upper half of septum. Style and stigma 11–13 mm long, exserted for 5–7 mm; style densely covered with long erect hairs in upper half (pubescence visible at anthesis); stigmatic lobes 4.5–5 mm long, upper 2.5–3 mm fusiform; papillate grooves along fusiform part and running further down for ca 2 mm. Fruits spherical, 3.5–6 mm in diam., wrinkled when dried, moderately to densely covered with appressed, spreading or erect hairs; fruit wall thin, with sclerified vascular bundles; pyrenes hemispherical, 3–5 mm in diam., papyraceous, with rounded base and tip and small opening in upper half adaxially. Seeds (1–)2–6 per fruit, 2.5–4 × 2–3.5 mm.

Distribution

Known throughout Madagascar. Present in: Antsiranana II and Nossi-Be Districts, Diana Region, and Vohemar District, Sava Region (Antsiranana Province); Ambatondrazaka, Andilamena and Moramanga Districts, Alaotra-Mangoro Region (Toamasina Province); Maevatanana and Tsaratanana Districts, Betsiboka Region, Ambato-Boeni, Mahajanga I, Mahajanga II, Mitsinjo and Soalala Districts, Boeny Region, Antsalova and Maintirano Districts, Melaky Region, and, Antsohihy, Boriziny-Vaovao, Mampikony and Mandritsara Districts, Sofia Region (Mahajanaga Province); Andramasina, Anjozorobe, Ankazobe, Antananarivo Renivohitra and Manjakandriana Districts, Analamanga Region, Arivonimamo and Miarinarivo Districts, Itasy Region, and, Ambatolampy, Antanifotsy, Antsirabe II, Betafo and Faratsiho Districts, Vakinankaratra Region (Antananarivo Province); Ihosy District, Ihorombe Region, Ambatofinandrahana and Ambositra Districts, Amoron’i Mania Region, and, Ambalavao District, Haute Matsiatra Region (Fianarantsoa Province); Amboasary-Atsimo, Betroka and Taolagnaro Districts, Anosy Region, Beroroha, Betioky, Morombe and Sakaraha Districts, Atsimo-Andrefana Region, and, Morondava district, Menabe Region (Toliara Province) ( Fig. 23B ).

Habitat and phenology

Dry deciduous or semi-deciduous forest or forest remnants, dry littoral forest, dry scrub, more rarely subhumid forest; on sandy soil (laterite or white sand) or on rocks; elev. 0–1650m. Flowers: (November–) December–March; Fruits: February–September.

Vernacular names and uses

Lengohazo (Humbert 14471); tarety (coll. ignot. 14803-SF); voatoana (Randriamiero 8646-RN). The species is used for firewood (coll. ignot. 14803-SF) and wood for construction (coll. ignot. 14803-SF, coll. ignot. 17762-SF, Razafinarivo 19142-SF).

Provisional IUCN assessment

Least Concern. This assessment is based on 153 herbarium collections collected between 1840 and 2018. The species occurs throughout Madagascar and the EOO is estimated to be 504911 km 2, far exceeding the upper limit of any threat category under criterion B1. The AOO is estimated to be 456 km 2, which complies with the Endangered category under criterion B2. The number of locations far exceeds 20, which is higher than the upper limit for any threat category under subcriterion ‘a’ of criterion B2. The species is present in several protected areas, notably: Andohahela (parcelle 3); Ankarafantsika; Andringitra, Analamerana, Bemaraha, Isalo, Kirindi, Loky-Manambato, montagne des Français, and Zombitse-Vohibasia. Throughout Madagascar natural vegetation is impacted by anthropogenic action, e.g., slash-and-burn agriculture, burning for grazing, logging for construction wood, firewood, and wood for the production of charcoal, resulting in a decrease in the quantity and quality of the habitat of P. cinerea . However, the species is very widespread and, while certainly threatened locally by destruction of its habitat, its large distribution area covers many other localities with more intact natural vegetation. Because of the above-mentioned facts, the species is assessed as Least Concern.

Notes

1. Paracephaelis cinerea is very variable in leaf shape, size and pubescence. Specimens with densely pubescent lower leaf surfaces are found mostly in the Central Highlands and in northern Madagascar (Analamera, forêt d’Orangea, Daraina), much less along the west coast. Specimens with less pubescent leaves have the same distribution range but were also frequently collected along the west coast and in south-eastern Madagascar (Andohahela). A number of specimens have (almost) completely glabrous leaves (with at most a few hairs on the midrib on upper and/or lower leaf surfaces). They are restricted to the Diana and Sava Regions. Examples are Vesco s.n., Capuron 27477-SF, Richard 83, and Razakamalala et al. 993. Other specimens have large, densely pubescent, almost orbicular leaves and relatively large inflorescences. They are most common in the Diana, Sava, and Boeny Regions and differ strikingly from P. cinerea specimens with smaller, less pubescent leaves. Leroy annotated one of these specimens (Capuron 24670-SF) as “ Paracephaelis diegoensis ”. Other examples of this large-leafed form are Andrianantoanina & Rabeharinosy 705, Rakotonasolo et al. 2143, Ranirison & Nusbaumer 491, and Capuron 23073-SF. But except for the large, orbicular leaves, no characters differentiate these specimens from other P. cinerea representatives. The morphological diversity of P. cinerea is the largest in the Diana and Sava Regions in northern Madagascar, where one may encounter all forms described above. It is also there that caducous specimens were collected (e.g., De Block et al. 1174 from Analamera; De Block et al. 1291 from baie de Sakalava).

2. Bremekamp (1934) transferred Pavetta cinerea to the genus Tarenna , giving it the name Tarenna cinerea . However, this name was already in use for a species from Thailand ( Craib 1932); therefore, the new combination was illegitimate. Bremekamp (1939) renamed the species Tarenna spodia .

3. Capuron (1973) erroneously used the name “ Tarenna sericea ” when referring to Tarenna cinerea (A.Rich.) Bremek in his unpublished Révision des Rubiacées de Madagascar et des Comores. He placed this species in the monospecific section Cinereae within Tarenna .

4. In the past, specimens of P. cinerea were annotated by different authors. Leroy annotated Capuron 24670-SF as “ Paracephaelis diegoensis ” and Sajy 25847-SF as “ Paracephaelis calyptroides ”. Homolle annotated Humbert 4048 and Perrier de la Bâthie 13459 as “ Tarenna cinerea var. majungensis ”. Baker annotated Baron 4596 as “ Tarenna heterophylla ”. Attempting to classify the morphological variation within P. cinerea in an early phase of this revision, De Block annotated Capuron 23309-SF as “ Paracephaelis cinerea subsp. diegoensis ”. None of these names were validly published.

5. Bojer invalidly published the name “ Pavetta glauca ” for Pavetta cinerea ( Bojer 1837: 172) .

6. The specimen Baron 4596 is present in BM, K and P. All duplicates contain a mixture of material from P. cinerea and Helictosperma malacophylla .

7. One specimen, Boivin s.n., from Île Sainte Marie, collected in 1849 with hardly any label data, is discarded for the provisional IUCN assessment. It is the only specimen of P. cinerea from the Analanjirofo Region and may be wrongly localized.

Figures

Fig. 1. Flowers, fruits and stipules of Paracephaelis Baill. A–D. Inflorescences. E–H. Fruits. I–J. Stipules. K–L. Infrutescences. A, E, I. P. cinerea (A.Rich. ex DC.) De Block. B. P. pauciflora De Block sp. nov. C, G. P. saxatilis (Scott Elliot) De Block. D. P. sericea (Arènes) De Block. F. P. ranirisonii De Block sp. nov. H. P. tiliacea Baill. J–K. P. capitulifera De Block sp. nov. L. P. russata De Block sp. nov. Photographs: P. De Block (B–E, G, I–L); S. Dessein (A, H); L. Gautier (F).

Fig. 2. Overview of the placentation in Paracephaelis Baill. A–K. Abaxial views of placenta and ovules. A’. Adaxial view of placenta and ovules with zone of attachment of the placenta indicated. A–C. Typical placentation type of Paracephaelis, with many ovules arranged at the periphery of the placenta.D–E. Idem, with fewer ovules. F–H. Idem, but ovules restricted to the top of the placenta. I. Typical placentation type in P. cinerea (A.Rich. ex DC.) De Block, P. comorensis De Block sp. nov. and P. trichantha (Baker) De Block. J–K. Idem, with fewer ovules. Drawn by Mr A. Fernandez. From Perrier de la Bâthie 13452 (P. tiliacea Baill., A, A’); Pervillé 633 (P. tiliacea Baill., B); Capuron 23451-SF (P. longipedicellata De Block sp. nov., C); McPherson et al. 14875A (P. saxatilis (Scott Elliot) De Block, D); Gautier et al. 4188 (P. gautieri De Block sp. nov., E); De Block et al. 1280 (P. bardotiae De Block sp. nov., F); De Block et al. 1048 (P. russata De Block sp. nov., G–H); Capuron 23073-SF (P. cinerea (A.Rich. ex DC.) De Block, I); Barthelat et al. 732 (P. comorensis De Block sp. nov., J); and Labat & Pascal 2887 (P. comorensis De Block sp. nov., K).

Fig. 8. Paracephaelis cinerea (A.Rich. ex DC.) De Block. A. Flowering and fruiting branches. B. Flower. C. Flower bud. D. Longitudinal section through ovary and calyx. E. Bracteole, ovary and calyx. F. Longitudinally opened flower with style and stigma. G. Anther. H. Fruit. I. Seed.Adapted from Drake del Castillo (1897: pl. 421).

Fig. 23. Distribution maps of Malagasy species of Paracephaelis. A. P. aristata De Block sp. nov. (black dot), P. bardotiae De Block sp. nov. (blue dot), P. capitulifera De Block sp. nov. (green dot), P. gautieri De Block sp. nov. (red dot), and P. grandifructa De Block sp. nov. (yellow dot). B. P. cinerea (A.Rich. ex DC.) De Block (black dot). C. P. longipedicellata De Block sp. nov. (black dot), P. orientalis De Block sp. nov. (blue dot), P. pauciflora De Block sp. nov. (green dot), P. ranirisonii De Block sp. nov. (red dot), and P. seyrigii De Block sp. nov. (yellow dot). D. P. russata De Block sp. nov. (black dot), P. sambavensis De Block sp. nov. (blue dot), P. saxatilis (Scott Elliot) De Block (green dot), P. sericea (Arènes) De Block (red dot), and P. tiliacea Baill. (yellow dot).