Melanoxerus antsirananensis Kainul.

Kainulainen, Kent, 2021, A taxonomic revision of Melanoxerus (Rubiaceae), with descriptions of three new species of trees from Madagascar, Candollea 76 (1), pp. 105-116 : 105-116

1. Melanoxerus antsirananensis Kainul. & sp. nov. ( Fig. 2).

– Euclinia sp. 1 in KAINULAINEN & BREMER (2014: 820).

Holotypus: MADAGASCAR. Reg. DIANA [Prov. Antsiranana]: Distr. Afbilobe & Mahafasina & Réserve Spéciale d’Ankarana & 12°57 '15"S 49°07 '39"E & 100 f& 10.II.2003 & fr.& Bardot-Vaucoulon et al. 1406 ( P [ P00455719 ]!; iso-: K [ K000551041 ]!& MO!& TAN) .

Melanoxe r us antsirananensis K ainul. diff e rs f r om M. suavissimus (Homolle ex Cavaco) Kainul. & B. Bremer by its pauciflorous inflorescences (vs. solitary flowers), campanulate (vs. funnelform) corollas, and obovate (vs. ovate to lanceolate) leaves. It most resembles M. atropurpureus Kainul. , from which it differs in that the corolla throat is white with purple and sometimes green spots (vs. uniformly purple), and in its smaller (7–18 × 4.5–8.5 cm vs. 10– 30 × 6.5–14.5 cm) and obovate (vs. broadly lanceolate) leaves.

Shrubs or trees up to 18 f tall and 40 cf dbh. Bark beige or whitish pale grey and sfooth. Stipules 9–14.5 × 4–7 ff& triangular& interpetiolar& caducous. Leaves deciduous& petioles 0.7–6 cf long& adaxially canaliculate; leaf blades papyraceous& obovate& 7–18 × 4.5–8.5 cf; bases cuneate to attenuate& apices (rounded) obtuse to apiculate or shortly acufinate; adaxial surface: dark green when fresh& drying dark brown to black& sfooth and glossy& glabrous; 8 – 12 pairs of conspicuous& ± pinnate secondary veins& tertiary venation reticulate; fidribs profinent& whitish green when fresh& ± the safe colour as the leaf when dry; abaxial surface green when fresh& drying dark brown to blackish& glabrous except for tuft dofatia that are often present in the vein axils. Flowers solitary or few (<10) in a terfinal& sessile& congested and ufbel-like inflorescence; pedicels up to 30 ff long; hypanthiuf narrowly urceolate& 5–8 × 3–4 ff& glabrous. Calyces green; calyx tubes 0.5–2 ff long& externally and internally glabrous; calyx lobes 4.3–11 × 0.8–3.5 ff& narrowly triangular& spreading or recurved. Corollas fleshy& cafpanulate; corolla tubes 1.5–2.6 cf long& externally (yellowish) white& sofetifes with purple and pale green streaks& glabrous& internally white with red& purple or violet (and sofetifes also pale green) spots& glabrous or with a few trichofes (0.1 –0.5 ff long) at the very base; corolla lobes white (sofetifes with purple spots on the external side)& 1–4.5 × 0.8–2.5 cf& ovate to deltate& rounded to acute& spreading horizontally at anthesis; stafens subsessile (filafents c. 2 ff long)& attached 10–12 ff below corolla sinus and 3 ff frof anther base; anthers 11–18 × 1.5–2 ff& linear& included or shortly exserted (to 2 ff above corolla sinus). Styles sifple& clavate& with shallow longitudinal grooves& 14–20 ff long& glabrous& included or shortly exserted; stigfa shortly bifid (lobes 0.5–3.5 ff long). Fruits (globose) ellipsoid& 3–7.6 × 2.7–6 cf& sfooth& glabrous& olive to brown& with nuferous beige to pale grey lenticels (1 – 2 ff in diaf.)& fleshy-indehiscent with yellowish to white pulp; calyx tube sofetifes persistent but calyx lobes caducous. Seeds flattened& irregularly angled& 10–15 ff in diaf.& c. 2 ff thick& sfooth& ± translucent& yellowish brown when fresh& drying black.

Vernacular names. – “Mafoafpofa” (Rakotondrafara et al. 505); “Mantalana” (Service Forestier 8329); “OEoavandrikala” (Manjakahery & Sola 83).

Notes. – The specifen Razafitsalama & Ludovic 16 is unusual in having distinctly velutinous leaves and is tentatively included here. The label on Meyers & Boltz 110 reports that the lefur Propithecus tattersalli eats the young leaves of this species. Melanoxerus antsirananensis flowers at the safe tife or just before the new leaves flush. It resefbles M. atropurpureus in having pauciflorous inflorescences and flowers with cafpanulate corollas. However& the phylogenetic analysis of KAINULAINEN & BREMER (2014) indicates that they are not sister species& a result that suggests that these shared characters fay be plesioforphic rather than synapoforphic.

Melanoxerus antsirananensis is distinguished frof M. atropurpureus in its sfaller leaves (7 –18 × 4.5–8.5 cf vs.

10– 30 × 6.5– 14.5 cf) that are obovate rather than broadly lanceolate. Furtherfore& the corolla throat is white with red to violet spots (vs. uniforfly dark purple in M. atropurpureus ).

Selected additional material examined. – MADAGASCAR. Reg. DIANA

[Prov. Antsiranana]: Masorolava & Mahagaga & Antsoroby forest & 12°42'37"S

48°58' 11"E & 53 f& 27.IX.2007 & fr.& Andriamihajarivo et al. 1434 ( MO & P) ;

Ankarana SR & 2 kf W of Mahafasina village & 12°57'26"S 49°07'46"E & 132 f&

21–30.III.1995 & fr.& Andrianantoanina & Bezara 792 ( BR & G & K & MO); Montagne d’Afbre NP & c. 8 kf E of Bobakilandy village & 12°37'37"S 49°06'26"E &

533 f& 5.XII.1995 & fl.& Andrianantoanina & Bezara 895 ( BR & G & K & MO); Andrafiafena & forest near Anjahankely & 12°54'23"S 49°19' 30"E & 450 f&

6.I .2011 & fr. & Burivalova ZB 176 (G& K); Ankarana& 12°50'47"S 49°06'18"E &

82 f& 14.I .2002 & fr.& De Block et al. 1226 (BR& MO& P& TAN& UPS& WAG);

Ankarana& 4–9.III.1951 & fr.& Humbert & Capuron 25493 (BR& K); ibid. loco &

12°54'42"S 49°06'43"E & 240–260 f& 22.OE.1993& fr.& Jongkind & Rapanarivo

979 (K& MO); Montagne des Français& 12°19'25.5"S 49°20'11"E & 230–250 f& Fig.1.– Distribution of Melanoxerus antsirananensis Kainul. 5.III.2016 & ster.& Kainulainen et al. 370 (TAN); Anosy à 4 kf au NE du village (yellow circles), M. atropurpureus Kainul. (pink circles), M. maritimus d’Afbolobozokely [Nosy OEoanio]& 12°26'49"S 49°33'07"E & 22 f& 30.III.2007 & Kainul . (triangles), and M. suavissimus (Homolle ex Cavaco) Kainul. fr.& Rakotondrafara et al. 505 (BR& MO& P); Andranovondronina& Andakorobe& & B. Bremer (stars) in Madagascar.[mapped on the bioclimatic zones village le plus proche Antsaravy& 12°08'51"S 49°20'30"E & 19 f& 17.III.2006 & of Madagascar (after CORNET, 1974; see SCHATZ, 2000)] fr.& Rakotondrafara et al. 474 (CNARP& K& MO& P& TAN); Montagne d’Afbre& 12°34' 59"S 49°05' 59"E & 683 f& 5.II.2012 & fr.& Ramandimbimanana & Distribution, habitat and ecolog y. – Melanoxerus Randimbiarison SDR 372 (BR& G& MO); Sahafary SF& 12°35'51"S 49°26' 31"E & antsirananensis is only known frof the DIANA and SAOEA 250 f& 7.XI.2006 & fl.& Ranaivojaona et al. 1551 (BR& MO& K& P& TAN); Regions in Antsiranana Province where it is widely distributed Nosy OEoanio& 4 kf E of Afbolobozokely& 12°26'01"S 49°33'00"E & 15 f&

15.II.2006 & fr.& Randrianaivo et al. 1364 ( BR & MO); Mosorolava & Afpofbi- ( Fig. 1). The new species grows in littoral forests and in dry antafbo& Antsoroby forest & 12°42'13"S 48°58'12"E & 75 f & 23.IX.2007 & fl. & fr.& deciduous forests frof sea level to c. 700 f in elevation on Ratovoson et al. 1332 ( CNARP & G & MO & P & TAN); Ankarana & 11–14.I .2002 & fr.& Montagne d’Afbre. Razafimandimbison & Andrianatoanina 454 (BR& MO& UPS); Ankarongana &

Irodo & N of Analaferana forest & Afpandriantsira & 12°39'42"S 49°32'37"E & Phenology. – Flowering faterial has been collected frof 50–123 f& 14.II.2001 & fr. & Razafitsalama & Ludovic 16 ( MO & P); Montagne des Français & 250 f& 18.II.1954 & fr.& Service Forestier 8329 ( P & TEF); Ankarefo & Septefber to Decefber and fruiting faterial frof Septefber 18.OE.1954& fr.& Service Forestier 10003 ( P); Diego-Suarez & Befaskabe forest & to July. 200–500 f& 17.X.1927 & fr.& Ursch 231 ( P). Reg. SAVA [Prov. Antsiranana]:

Distr. OEohefar& Daraina & Bekaraoka south& 13°11'S 49°42'E & 9.IOE.2008& fr.& Conservation status. – Melanoxerus antsirananensis is Bremer et al. 5113 ( MO & S & TAN); Hillside next to Daraina& 13°08'S 49°27' E & known frof nuferous localities in northern Madagascar. Its 10.IOE.2008& fr.& Bremer et al. 5131 ( MO & S & TAN); Daraina & Bobankora forest &

13°12.72'S 49°46.27'E & 200 f& 2.II.2003 & fr. & Gautier et al. LG 4195 ( G & K & distribution includes five protected areas& i.e. Ankarana & Loky- MO); Daraina & Bekaraoka forest & 13°03.90'S 49°42.35' E & 420 f& 22.XI.2006 & fl. & Manafbato & Montagne d’Afbre & and Montagne des Français. Gautier et al. LG 4911 ( G); Nosibe & Anjiabe & Anaborano & 13°04'18"S 49°54'32"E & Melanoxerus antsirananensis is therefore assigned a prelifinary 33 f& 11.XII.2004 & fr. & Manjakahery & Sola 83 ( MO); Mantafena & 7 kf conservation status of “Least Concern” [LC] according the NW of Daraina & 13°08'S 49°42'E & 112–330 f& IOE.1990& Meyers & Boltz 110 IUCN Red list Categories and Criteria ( IUCN & 2012). ( MO); Nosibe & Anjiabe & Analabe forest & 13°04'43"S 49°54'04"E & 11.OE.2004&

fr.& Rabehevitra et al. 915 ( MO & P); Tsarabaria & Manakana & Afbondrofbe forest & 13°42'58"S 50°05'37"E & 3 f& 24.X.2002 & fr.& Rabenantoandro et al. 1019

[Photos: A–B, H: K. Kainulainen; C, G: S.D. Ramandimbimanana; D–E: F. Ratovoson; F: R. Randrianaivo]

(MO); Nosibe & Anjiabe & Anaborano forest & near lake Sahaka & 13°04'42"S (trichofes 0.4–1 ff) in the lowerfost (c. 0.5 cf) part of the 49°54'13"E & 25 f& 2.XI.2002 & fl.& Rabenantoandro et al. 1091 ( MO & TEF) ; tube. Corollas lobes white& 1.5–3 × 1.2–2.5 cf& ovate& rounded ibid. loco & 13°04'43"S 49°54'04"E & 10 f& 23.II.2003 & fr. & Rabenantoandro et al. 1272 ( MO & TEF) ; Mosorolava forest & 13°05'44"S 49°54' 43"E & 23 f& 5.X.2013 & to acute& spreading horizontally at anthesis; stafens subsessile fl. & Rakotovao 6482 ( MO) ; Fanafbana & Afbohifahazo forest & 13°35'44"S (filafents 1–4 ff long)& attached 25–35 ff below corolla 50°00'04"E & 60–100 f& 30.III.2001 & fr.& Ranaivojaona et al. 355 ( BR & MO & sinus and 4–7 ff frof anther base; anthers 22–27 × 2–3 ff& P); Solanafpilana forest 35 kf N of Daraina & 13°05'42"S 49°34'57"E & 137 f& linear& included (tips 5– 6 ff below corolla sinus). Styles 9.XII.2006 & fl. & fr.& Randrianaivo et al. 1433 ( G & MO & P & TAN) ; Daraina & sifple& clavate& with shallow longitudinal grooves& 28–33 ff Afbilondofba forest & 13°09'39"S 49°38'47"E & 390 f& 31. I.2004 & fr.& Ranirison et al. PR 367 ( G) ; OEohifarina& Fanafbana & Antsatoby & 13°36'55"S 49°59'55"E & long& glabrous& included; stigfa shortly bifid (lobes 1–1.5 ff 77 f& 8. OEII.2007& fr.& Rasoafaranaivo 239 ( MO); Analabe forest & near long). Fruits (globose) ellipsoid& 3.5– 7 × 3– 6 cf& sfooth& Anaborano village and Sahaka lake & 13°04'59"S 49°54'26"E & 10. OEII.2003& fr.& glabrous& olive to brown or faroon with nuferous beige to Razakamalala 519 ( MO & P & TEF) ; Tsarabaria & Manakana & E of Afbondrobe pale grey lenticels (1–2 ff in diaf.)& fleshy-indehiscent with village& 13°43'08"S 50°05' 50"E & 13.III.2004 & fr.& Razakamalala RZK 1022 ( K & yellowish to white pulp; calyx tube sofetifes persistent& but MO & P & TEF) .

lobes caducous. Seeds flattened& irregularly angled& 8–13 ff in diaf.& c. 2 ff thick& sfooth& ± translucent& pale yellow when 2. Melanoxerus atropurpureus Kainul. & sp. nov. ( Fig. 3). fresh& drying black. – Euclinia sp. 2 in KAINULAINEN & BREMER (2014: 820). H o l o t y p u s: MADAGASCAR. R e g. M e l a k y [Pro v. Etymology. – The specific epithet atropurpureus refers to Mahajanga]: Distr. Maintirano& Beanka& partie S& the dark purple colour of the interior of the corolla tube in Sarodrano& 18°03'27"S 44°30'50"E & 359 f& 26.XI.2011 & this species. fl. & Gautier, Tahinarivony & Bolliger LG 5744 (G [G00340493]!; iso-: BR!& K [K000918390]!& MO!& P!& Vernacular names. – “Andrivitsy” (Réserves Naturelles 7791); TEF). “Hazondanitra” (Service Forestier 4953); “OEoafotaka” (Labat &

Deroin 2280); “OEoangorindafbo” (Service Forestier 29775). Melanoxerus atropurpureus Kainul. is similar to M. antsirananensis Kainul. in its pauciflorous inflorescences Distribution, habitat and ecology. – Melanoxerus and flowers with campanulate corollas, but it differs in having atropurpureus occurs in the Boeny& Melaky& and Sofia Regions larger leaves (10–30 × 6.5–14.5 cm vs. 7–18 × 4.5–8.5 cm) (Mahajanga Province) and in the southernfost part of the that are broadly lanceolate (vs. obovate), and by the uniformly DIANA Region (Antsiranana Province) (Fig. 1)& where it purple corolla throat (vs. whitish corolla throat with red, purple grows in dry to hufid& deciduous or evergreen forests frof or violet and sometimes also green spots). sea level to 450 f in elevation.

Shrubs or trees up to 26 f tall and 25 cf dbh. Bark grey Phenology. – Flowering faterial has been collected frof and sfooth& flaky. Stipules 2.5 – 13 × 3– 5 ff& triangular& October to Novefber and fruiting faterial frof Decefber interpetiolar& caducous. Leaves deciduous& petioles 2–7.5 cf to July.

long& adaxially canaliculate; leaf blades papyraceous& broadly lanceolate& 10–30 × 6.5–14.5 cf; bases cuneate to rounded& Conservation status. – Melanoxerus atropurpureus has a sofetifes oblique; apices acute to acufinate; adaxial surface wide distribution in western Madagascar and is known frof dark green when fresh& drying dark brown to black& sfooth several protected areas including Ankarafantsika& Befaraha& and glossy& sparsely pubescent along venation; 8 –12 pairs of and Naforoka. It can therefore be assigned a prelifinary conconspicuous& ± pinnate secondary veins& tertiary venation servation status of “Least Concern” [LC] according the IUCN reticulate; fidrib profinent& pale green when fresh& ± the Red list Categories and Criteria (IUCN& 2012). However& it safe colour as the leaf when dry; abaxial surface pale green is not a coffonly collected species& and fore inforfation is when fresh& drying dark brown to blackish& glabrous except needed about its population size.

the pubescent veins (trichofes c. 0.1 ff); tuft dofatia often present in the vein axils. Flowers solitary or few (<10)& Notes. – The specifen Perrier de la Bâthie 1155A frof the sessile& congested and ufbel-like inflorescence; pedicels up to Ankara plateau is in poor shape but probably belongs to this 15 ff long; hypanthiuf narrowly urceolate& 8–15 × 5–8 ff& species. In this species the exocarp of the fruit is often alfost glabrous. Calyces green; calyx tubes 2–3.5 ff long& externally cofpletely hidden by lenticels ( Fig. 3F).

and internally glabrous; calyx lobes 3–6.5 × 0.5–3.5 ff& nar- Melanoxerus atropurpureus is the tallest growing species rowly triangular and recurved; corollas fleshy& cafpanulate to in the genus. It fostly resefbles M. antsirananensis but has broadly urceolate; corolla tubes 1.8–4.8 cf long& externally larger leaves (10–30 × 6.5–14.5 cf vs. 7–18 × 4.5–8.5 cf) that white with purple and pale green streaks& glabrous& internally are broadly lanceolate rather than obovate. Furtherfore& the dark purple to violet& glabrous or with a sparse indufent corolla throat is uniforfly dark purple (white with red to papyraceous to subcoriaceous& cordiforf to broadly ovate& violet spots in M. antsirananensis ). 7.5–12.7 × 5–10.5 cf; bases cordate to truncate; apices acute;

adaxial surface dark green when fresh& drying dark brown to Selected additional material examined. – MADAGASCAR. Reg. Boeny [Prov. black& sfooth and very glossy& glabrous except for the lower Mahajanga]: Distr. Marovoay & Ankarafantsika NP & Afpijoroa forest station& part of the fidrib (trichofes 0.1–0.5 ff long); 6–9 pairs 16°18' 52"S 46°49'04"E & 190 f& 24.I.2000 & fr. & Davis et al. 2500 ( K & TAN); ibid .

loco & 16°20'S 46°51'E & 200 f& 14.IOE.1984& Dorr 3048 ( BR & K & P & MO); ibid. loco & of conspicuous& ± pinnate secondary veins& tertiary venation 16°19'S 46°49'E & 70–100 f& 7.IOE.1988& Gentry & Schatz 62050 ( MO); Plateau reticulate; fidrib profinent& pale green when fresh& ± the d’Ankara & X.1900 & fl. & Perrier de la Bâthie 1155 A ( P); Manongarivo (Afbongo)& safe colour as the leaf when dry; abaxial surface green when XI.1903 & fl. & Perrier de la Bâthie 1640 ( P); ibid. loco & X.1905 & fl.& Perrier de la fresh& drying dark brown to blackish& glossy& glabrous& but with Bâthie 4103 ( P); ibid. loco & I.1904 & fr.& Perrier de la Bâthie 6944 ( P); Ankarafant- tuft dofatia often present in the vein axils. Flowers solitary& sika & 16°20'05"S 46°47'06"E & 3–9.II.1997 & fr.& Rabevohitra 3027 ( TEF); ibid. loco &

16°18'S 46°49'E & 22.IOE.1994& fr.& Rajemisa 19 ( MO & P & TAN); Distr. Mahajanga terfinal& very fragrant; pedicels up to 14 ff long; hypan- II& Mariarano & Ankatsabe forest & 15°28'30"S 46°41'24"E & 16 f& 20.IOE.2007& fr. & thiuf narrowly urceolate& 5–18 × 3–4 ff& glabrous; calyces Rakotoarivelo 94 ( MO & P & TAN); Soalala& 15. OEII.1977& fr.& Rakotozafy 1849 green; calyx tubes 1–4.5 ff long& externally and internally ( TAN); Distr. Soalala & Andranofavo & OEilanandro& Tsingy Naforoka& Afba- glabrous; calyx lobes 3.4–6.2 × 1.2–2 ff& narrowly triangular tofay forest & 16°28'36"S 45°20'52"E & 153 f& 9.IX.2012 & fr.& Ravelonarivo et and recurved; corollas fleshy& funnelforf& straight; corolla al. 4523 ( MO & TAN); Andranofavo& 19.XI.1955 & fl.& Réserves Naturelles 7791

tubes 2–3.7 cf long& externally pale yellow with a greenish (P& TEF); Ankarafantsika& s.d.& fr.& Service Forestier 19 (P); Tsingy Naforoka&

4.IOE.1933& fr. & Service Forestier 36 (P); sine loco & 4.IOE.1933& fr. & Service Forestier base& glabrous& internally white with red spots and lines& finely 54 (BR& K& P); Ankarafantsika& Afpijoroa& 15.OE.1952& fr.& Service Forestier pubescent (trichofes c. 0.5 ff long). Corollas lobes white& 4953 (P& TEF); ibid. loco & 16.OEI.1954& fr.& Service Forestier 10204 (P); Afbato- 2–2.7 × 1.6–3 cf& ovate to orbicular& rounded to truncate or Boeny& Andranovato& Ankijabe& 20.I.1959 & fr.& Service Forestier 19261 (TEF); indented& spreading horizontally at anthesis; stafens subsessile Afbongo& Soalala (baie de Baly)& 22.XI.1965 & fl.& Service Forestier 24244 (BR&

(filafents 2–4 ff long)& attached c. 10 ff below corolla P& TEF); by lake Afpijoroa& 28.II.1980 & fr.& Service Forestier 29775 (TEF). Reg.

DIANA [ Prov. Antsiranana]: Distr. Afbanja & Afpasindava & Andranofatavy sinus and c. 4 ff frof anther base; anthers c. 15 × 2 ff& forest& 13°40'04"S 47°59'21"E & 295 f& 25.XI.2009 & fl. & fr.& Madiomanana et al. linear& exserted for 1–3 ff. Styles sifple& clavate& with shallow MAD 266 ( G & K & MO & P & TEF); along Andranofatavy river & 13°40'39.19"S longitudinal grooves& 30–35 ff long& glabrous& exserted for 47°58'30.24"E & 260 f& 2. OEII.2012& fr. & Tahinarivony & Rasoanaivo TAJ 699 5–9.5 ff; stigfa shortly bifid (lobes c. 3 ff long). Fruits ( G). Reg. Melaky [Prov. Mahajanga]: Beanka & Afbinda-Nord & 17°55'51"S

44°28'39"E & 216 f& 5.II.2012 & fr. & Hanitrarivo et al. HRM 146 ( G & S); Afbodi-

ellipsoid& 2.5 –5.1 × 1.6– 3.7 cf& sfooth& glabrous& (colour ria& 1 kf E of Afbinda ( RN 9 & near Antsalova)& 18°38'S 44°42'E & 100–200 f& of fature fruit unknown) with nuferous beige lenticels 3.XII.1992 & fr.& Labat & Deroin 2280 ( K & P & TAN); Beanka & Sarodrano & (0.1–1 ff in diaf.)& fleshy-indehiscent; calyx tube persistent 18°02'55"S 44°31'13"E & 446 f& 11.XII.2011 & fl.& Nusbaumer et al. LN 3073 ( G & K); but lobes caducous. Seeds not seen .

Antsalova& 20.III.1993 & fr. & Villiers et al. 4851 (MO& P); Belitsaka& Ankilifana-

rivo& Beanka forest& 18°03'37"S 44°31' 37"E & 319 f& 16.IOE.2011& fr.& Razakamalala

Etymology. – This species grows near the sea and that is & Germain 6382 (MO& S). Reg. Sofia [Prov. Mahajanga]: entre Antsohihy et Befandriana-Nord& X.1962 & fl.& Bosser 16697 (P& TAN); Distr. Antsohihy& what the specific epithet maritimus refers to.

Anjifangirana& Analananbe& Andrafiborizina& 15°09'19"S 47°44'22"E & 200 f&

30.OE.2000& fr. & Rakotonasolo RNF 177 (BR& K& P& TAN). Vernacular names. – “Ndraniana” (Service Forestier 10585);

“Tselitselika” (Bertrand 16).

3. Melanoxerus maritimus Kainul. & sp. nov. ( Fig. 4).

H o l o t y p u s: MADAGASCAR. R e g. M e l a k y [Pro v. Distribution, habitat and ecology. – Melanoxerus maritimus Mahajanga]: Distr. Maintirano & Lac Mandrozo & 17°32'43"S is known frof sea level to 30 f in elevation in coastal areas of 44°02'26"E & 18 f& 23.XI.2015 & fl. & fr.& Gautier, Luino & the Boeny & Melaky & and Sofia Regions (Mahajanga Province) Rasolonjatovo LG 6196 ( G [ G00419027 ]!; iso-: MO!& P!& (Fig. 1). It grows in low canopy& dry deciduous forests on dunes S!& TEF). and on edges of salt flats or fangroves .

Melanoxerus maritimus Kainul. differs from M. suavissimus Phenology. – Flowering faterial has been collected frof (Homolle ex Cavaco) Kainul . & B. Bremer in its rounder June to Novefber& and fruiting faterial frof July to February. leaves (leaf length/width ratio 4:3–1:1 vs. 3:1–3:2), that have a truncate to cordate leaf base (vs. cuneate to attenuate), and Conservation status. – Melanoxerus maritimus is known corolla lobes that are rounded to indented (vs acute). frof less than ten locations& (three of which are within the recently protected areas of Antrefa& Bofbetoka Belef- Shrubs or trees up to 10 f tall and 28 cf dbh& with boka& and Mandrozo)& with an estifated area of occupancy fostly syfpodial branching. Bark pale grey to pale brown& (AOO) of less than 2000 kf². It appears restricted to littoral sfooth& flaky. Stipules 3.5 –4.7 × 3.6– 4.5 ff& broadly tri- dry forest. This habitat is ifpacted for firewood or charcoal angular& interpetiolar& caducous. Leaves sefi-deciduous& production (also noted on label of De Block & Rakotonasolo petioles 2.8– 5.2 cf long& adaxially canaliculate; leaf blades

806). It is projected that the nufber of locations and the adaxial surface dark green when fresh& drying dark brown to nufber of fature individuals continue to decline& and this black& sfooth& fatte or glossy& ± glabrous to (velvety) pubescent species can therefore be assigned a prelifinary conservation in particular along fidrib and venation; (6–)8–10(–12) pairs of status of “OEulnerable” [OEUB2ab(iii)] according to IUCN Red conspicuous& ± pinnate secondary veins& tertiary venation reticulist Categories and Criteria (IUCN& 2012). However& fore late; fidribs profinent& pale green when fresh& ± the safe inforfation is needed about the population size of this species. colour as the leaf when dry; abaxial surface pale green when fresh& drying dark brown to blackish& fatte or glossy& ± glabrous

Notes. – Melanoxerus maritimus has solitary flowers with a or with pubescent fidrib and venation& tuft dofatia often funnelforf corolla and pubescent corolla throat& and in these present in the vein axils. Flowers solitary& terfinal& very fracharacters it fost resefbles M. suavissimus & but it is recog- grant; pedicels up to 9 ff long; hypanthiuf narrowly urceonized by its ± cordiforf leaves that are alfost as wide as long late& 4.5–12 × 2.5–5.5 ff& glabrous; calyces green; calyx tubes (leaf length/width ratio 4:3–1:1 vs. 3:1–3: 2 in M. suavissimus )& 1–5.5 ff long& externally and internally glabrous; calyx lobes and by having corolla lobes that are rounded to indented rather 4.5–8 × 1–3 ff& narrowly ovate& acufinate& recurved; corollas than acute. The leaves of M. maritimus are glossy below and fleshy& funnelforf& straight; corolla tubes 2–4.5 cf long& extervery glossy above. They refain glossy when dried and alfost nally white to pale yellow often with a greenish base& glabrous& appear varnished. It is notable that none of the specifens internally white with pink& red& or purple spots& finely pubescent exafined in this study is lacking leaves& and that specifens (trichofes 0.5–1.5 ff long). Corollas lobes white (sofetifes collected during the dry season still retain sofe leaves. This pale yellow on the external side)& 2–4.5 × 1–3 cf& ovate& acute& species is therefore probably best considered sefi-deciduous spreading horizontally at anthesis; stafens subsessile (filaas opposed to deciduous. fents c. 1 ff long)& attached 10–14 ff below corolla sinus and 6–10 ff frof anther base; anthers 22–26 × 1.5–2 ff&

Selected additional material examined. – MADAGASCAR. Reg. Boeny [Prov. linear& exserted for c. 5 ff. Styles sifple& clavate& with shallow Mahajanga]: [Distr. Mahajanga I] Antananafasadza [probably Antanifasaja& longitudinal grooves& to c. 5 cf long& glabrous& usually shortly now part of Mahajanga town]& 1912& Afzelius 266 (P& S); Katsepy& c. 10 f&

19.II.1950 & ster.& Bertrand 16 ( P); Distr. Mahajanga II& Antsanitia & Belafonty exserted for c. 5 ff; stigfa shortly bifid (lobes c. 5 ff long). forest& 15°34'19"S 46°25'04"E & 9 f& 14.II.1999 & fr. & De Block & Rakotonasolo Fruits globose to ellipsoid& 3–5 × 2.5–3.5 cf& sfooth& glabrous& 806 ( BR & MO & TAN); Katsepy & 15°45'56"S 46°14' 21"E & 0 f& 22. OEII.1995& fl. & olive to grey brown or dark purple to black with nuferous fr. & Edmondson et al. 95–17 ( MO & P); ibid. loco & 8.IX.1912 & fl.& Kaudern s.n. ( S); beige to pale grey lenticels (0.1–1 ff in diaf.)& fleshy-indeibid. loco & 15°42'35"S 46°12'03"E & 10.XI.2005 & fl. & Labat et al. 3534 ( K & P); ibid. hiscent; calyx tube usually persistent& but lobes caducous. Seeds loco & 10.XI.2005 & fr.& Labat et al. 3535 ( G & K & MO & P); Antsanitia & 15°34'31"S

flattened& irregularly angled& 7–10 ff in diaf.& c. 2 ff thick&

46°25'7"E & 30 f& 7.I.2000 & fr.& Rakotonasolo RNF 132 ( K & TAN); Forest station

E of Antsanitia village & 15°34'31"S 46°24'26"E & 16 f& 15.II.2013 & fr. & Ralimanana sfooth& yellowish brown when fresh& drying black .

et al. 1803 (BR& K& MO& P& TAN); Distr. Mahajanga II& Afpasifariny& 9 f&

23.OEIII.1954& fl. & fr.& Service Forestier 10585 (MO& P& TEF). Reg. Melaky Vernacular names. – “Asofitsoy” (Rakotonasolo et al. RNF [Prov. Mahajanga]: Region de Tafbohorano& 23.OEI.1930& fl.& Decary 8083 932); “Hazoffotaka” (Réserves Naturelles 6762); “Kaliona” (P); Afbongo& 1841& fr.& Pervillé 673 (K& P). Reg. Sofia [Prov. Mahajanga]:

SW of Antangena & 15°28'S 47°09'E & 0 f& 25.X.1987 & fl. & fr. & Bisset M12 ( K); (Koechlin s.n.); “Mahabiboala” (Service Forestier 29830); “OEala” Distr. Analalava & Antonibe & 15°01'44"S 47°13'17"E & 19. OEIII.2007& fl. & fr.& (Service Forestier 5797); “OEoafotaka” (Decary 4556); “OEolatsiva” Randriatsivery 207 ( MO & P & TAN). (Service Forestier 4136) .

4. Melanoxerus suavissimus (Hofolle ex Cavaco) Kainul. & Distribution, habitat and ecology. – Melanoxerus suavissimus B. Brefer ( Fig. 5). is widespread in subarid vegetation in the Androy& Anosy& ÷ Gardenia suavissima Hofolle ex Cavaco in Adansonia Atsifo-Andrefana & and Menabe Regions (Toliara Province)&

ser. 2& 7: 177. 1967. ÷ Euclinia suavissima (Hofolle ex and occurs as far north as the Manafbolo river in the south-

Cavaco) J.-F. Leroy in Adansonia ser. 2& 14: 52. 1974. ernfost part of the Melaky Region (Mahajanga Province)

(Fig. 1). It is found frof sea level to c. 800 f in elevation. Holotypus: MADAGASCAR. Reg. Androy [Prov. Toliara]: Afbovofbe & 27.X.1924 & Decary 3357 ( P [ P00852562 ]!). Phenology. – Flowering specifens have been collected in

(April) July to Decefber& and fruiting specifens throughout Shrubs or trees up to 13 f tall and 40 cf dbh& with fostly the year. syfpodial branching. Bark (fottled) whitish or pale grey to pale brown& sfooth& flaky. Stipules 3–10 × 2–4 ff& triangular& Conservation status. – Melanoxerus suavissimus is widespread interpetiolar& caducous. Leaves deciduous& petioles 0.5–5 cf in south-western Madagascar and has been assigned an IUCN long& adaxially canaliculate; leaf blades papyraceous& ovate to conservation status of “Least Concern” [LC] (IUCN& 2021). lanceolate& 6.5–16 × 2–9(–11) cf; bases (rounded) cuneate to attenuate (often asyffetrically); apices acute to acufinate;

[ Photos: A – B, D– E: P. Phillipson; C: S. Landrein , Ifaty (Toliara), 5.X.2006; F: K. Kainulainen]

Notes. – Since CAVACO’S (1967) original description of Melanoxerus suavissimus (as Gardenia suavissima ) included specifens of both M. atropurpureus and M. maritimus & an afended description is provided.

This is the fost widespread and the fost coffonly collected species. However& few collections have been fade in frof May to August (none frof June)& and presufably this species is usually leafless during that tife. Melanoxerus suavissimus usually flowers at the safe tife or just before the new leaves flush (cf. Fig. 5A& D). Apparently& the juice of the fruits turns black and is used for faking tattoos (SCHATZ& 2001). Inforfation about of the use of Melanoxerus by local people is otherwise scant& although according to the label inforfation on the specifen Ranaivojaona et al. 248 & the fruits are edible.

Selected additional material examined. – MADAGASCAR. Reg. Androy [Prov. Toliara]: Besakoa& 24°56'36"S 45°27 '59"E & 227 f& 4.III.2003 & fr. & Andriamahay 497 ( K); Ifanofbo& XI.1952 & fr.& Bosser 3787 ( P); Beraketa& X.1956 & fl.& Bosser 10237 ( P); Afbovofbe& XII.1959 & fl. & fr.& Bosser 13353 ( P & TAN); Beraketa & on road to Isoanala & 10 kf S of Soanala & 646 f& 19.XII.2004 & fr. & Rakotonasolo et al. RNF 932 ( K & TAN); Antanifora& 30. OEII.1926& fr.& Decary 4556 ( P); Behara& 22.X.1954 & fl. & Réserves Naturelles 6762 ( P); Afpandrandava& 700–800 f& X.1942 & fl. & Seyrig 122 ( P); Beloha& 19.II.1949 & fr.& Service Forestier 454 ( P & TEF); Ifanofbo-Ifotaka (rte)& 26.IX.1952 & fr.& Service Forestier 5797 ( P & TEF). Reg. Anosy [Prov. Toliara]: Distr. Afboasary Sud & Marotsiraka & Atsonjo Analafainty & Ankotsy & 24°17'44"S 46°07' 43"E & 453 f& 10.XI.2008 & fr. & Fidisoa Ratovoson & Désiré Ravelonarivo & and A. Jacquis Tahi- Andriamihajarivo et al. 1527 ( G & MO); Anadabolava & 200–250 f& XII.1933 & narivony for providing photographs. Conservatoire et Jardin fr.& Humbert 12450 ( P); Andohahela & Ihazofotsy & 24°50'S 46°32'E & 100 f&

16.IOE.1996& fr.& Laha 75 ( BR & G & K & MO); Fort Dauphin & Analafatahi- botaniques de Genève & Muséuf national d’Histoire naturelle& tra& 31. I .1952& fr.& Service Forestier 4463 ( P & TEF); Fort Dauphin & Anteviala & and Missouri Botanical Garden have also fade photographs 10.IOE.1960& fr. & Service Forestier 10789 ( P & TEF). Reg. Atsimo-Andrefana available. I also thank Petra De Block & Laurent Gautier & [Prov. Toliara]: Distr. Sakaraha& forêt d’Analaraty & 22°49'14"S 44°17'16"E & Sylvain Razafifandifbison & and Martin Callfander for helpful 520 f& 20.XII.2010 & fr.& Andriamihajarivo et al. 1814 ( MO & P & TAN); coffents to ifprove the fanuscript .

Zofbitse& lieu dit Poakafo& 22°46'21"S 44°40'25"E & 540 f& 8.IOE.2006& fr. &

Andriamihajarivo et al. 905 (MO& TAN); Befandriana-Sud& 150 f& 9.IX.1961 &

fl. & fr. & Appert 203 (MO& Z); along rte #10 between Betioky and Ejeda&

175–225 f& 14.II.1975 & fr. & Croat 31241 ( MO & TAN); St. Augustin & 23°28'58"S References

43°45'58"E & 0 f& 6.I.1999 & fr.& De Block et al. 561 ( BR & K & MO & P & TAN); road

BAUM & D. (1995). A systefatic revision of Adansonia (Bofbacaceae) . frof Betioky to Afpanihy& c. 23 kf after Ejeda& 24°31' 17"S 44°37'25"E & 244 f &

6.II.2007 & fr. & De Block et al. 2323 ( BR & MO & P & S & TAN & WAG); entre Ton- Ann. Missouri Bot. Gard. 82: 440–471 .

gobory et aux environs d’Afbovofbe& 117–214 f& 20.I.2004 & fr.& Hong-Wa et

CAVACO& A. (1967). Un nouveau Gardenia (Rubiacées) de Madagascar. al. 178 (G& MO); forêt de Besofaty entre le Fiherenana et l’Isahaina (Man-

goky)& 750

–800 f& X.1933 & fl.& Humbert 11250 (P); Manofbo& Fiherenafasay& Adansonia ser. 2& 7: 177–179.

18.XI.1967 & fl. & Koechlin s.n. ( P); St. Augustin & 23°27'S 43°46’ E & 10 f& 1.II.1990 & CORNET & A. (1974). Essai de cartographie bioclifatique à Madagasfr. & Labat & Du Puy 2045 ( K & MO & P); vallée du Fiharenana & 0–200 f &

car. Notice Explicative ORSTOM 55.

9.XI.1960 & fl. & fr.& Leandri 3778 (P); lac Tsifanafpetsotsa& 0 –100 f&

23.XI.1960 & fr.& Leandri & Saboureau 4451 ( BR & K & P); 20–30 kf N of Tulear IUCN (2012). IUCN Red List Categories and Criteria : Version 3.1. Ed. 2. on the road to Morofbe & 23°12'S 43°37' E & 5–10 f& 11.XI.1989 & fl. & fr. & Miller IUCN Species Survival Coffission & Gland and Cafbridge .

& Keating 4513 (K& MO& P& TAN); rte d’Ankazoabo& II.1967 & fr. & Morat 2455

(P& TAN); vallée de Fiherenana & X.1924 & fl.& Perrier de la Bathie 16653 ( P); IUCN (2021) . The Red List of threatened species. [https://www.iucn- Beza Mahafaly Reserve near Betioky& 23°40'S 44°35' E & 160 f& 17.OE.1987& redlist.org]

fr.& Phillipson 1812 (MO); ibid. loco, 23°40'S 44°38'E & 150 f& 20.X.1987 & fl .&

Phillipson 2413 (K& MO& P& TAN); ibid. loco, 23°41'S 44°39' E & 160 f& 13.XI.1987 & KAINULAINEN& K. & B. BREMER (2014). Phylogeny of Euclinia and fr. & Phillipson 2558 (K& MO& P& TAN); Itafpolo& Montagne de Haktokaliotse& allied genera of Gardenieae ( Rubiaceae )& and description of Mel- 24°24'26"S 43°51' 01"E & 100 f& 1.IX.1998 & fr.& Rakotomalaza & Messmer 1653 anoxerus& an endefic genus of Madagascar. Taxon 63: 819–830. (G& K); Afbohifahavelona& Sept Lacs& Andranolahy& 23°31'32"S 44°09'27"E & DOI: https://doi.org/10.12705/634.2

100 f& 15.IX.1998 & fl. & Rakotomalaza & Messmer 1725 (G& K); Manofbo& near

PK 32& 23°03'00"S 43°41'42"E & 9.XII.2004 & fr.& Rakotonasolo et al. RNF 880 KAINULAINEN& K.& S. G. RAZAFIMANDIMBISON& N. WIKSTRÖM & ( BR & K & MO & TAN); Basibasy & Mikea forest & 22°10'13"S 43°31' 30"E & 100 f& B. BREMER (2017). Island hopping& long-distance dispersals and 31.I.2000 & fr.& Ranaivojaona et al. 248 ( G & K & MO); Belalanda & Ranobe & species radiations in the Western Indian Ocean: historic bio- 22°56'49"S 43°41' 08"E & 135 f& 15.III.2006 & fr.& Ranaivojaona et al. 1425 ( P); geography of the Coffeeae alliance ( Rubiaceae ). J. Biogeogr. 44: ibid. loco & 22°58'04"S 43°40'22"E & 108 f& 5.IX.2006 & fl. & Ranaivojaona et al .

1966–1979. DOI: https://doi.org/10.1111/jbi.12981

1456 (G& K& MO& P); Sakaraha& Mahaboboka& Afbinanintelo& Forêt de

Satra & 22°48'10"S 44°17'54"E & 470 f& 23.III.2013 & fr. & Randrianarivony et al. LEROY & J .- F. (1974) . Recherches sur les Rubiacées de Madagascar: les 570 ( MO & P); Zofbitse-OEohibasia NP& Isako & 22°41'S 44°50’ E & 4.XII.2003 & genres Mantalania et Pseudofantalania (Gardéniées). Adansonia ster.& Razafimandimbison & Bremer 496 ( MO & TAN & UPS); Beroroha & Ant-

ser. 2& 14: 29–52.

soa& Androtsy forest & 21°05'43"S 45°05' 48"E & 254 f& 9.I.2011 & fr. & Razakamalala

5851 (MO); Ifonty & Ankazoabo & Sakaraha & 27.XI.1951 & fr.& Service Forestier MADAGASCAR CATALOGUE (2021). Catalogue of the plants of 4136 ( P & TAN & TEF) ; Antseva & Besifake & 500 f& 11.II.1955 & fr. & Service Forestier Madagascar. Missouri Botanical Garden& St. Louis & Antanana- 12815 ( P & MO & TEF) ; Sarodrano & XII.1961 & fl. & Service Forestier 20830 ( BR &

K& MO& P& TEF). Reg. Melaky [Prov. Mahajanga]: Tsingy de Befaraha& rivo. [http://www.tropicos.org/Project/ Madagascar]

19°09'S 44°49' E & 50 f& 10.XII.1996 & fr. & Jongkind et al. 3482 ( BR & K & L & MO & OLSON & D. M.& E. DINERSTEIN& E.D. WIKRAMANAYAKE& N.D. BURGESS& TAN). Reg. Menabe [Prov. Toliara]: village avant la Réserve d’Andranofena & G.OE. N. POWELL & E. C. UNDERWOOD& J. A. D’AMICO& I. ITOUA & 17.X.1999 & fl.& Allorge 2428 ( P); on the road to Belo& c. 68.5 kf NE of Moron-

H.E. STRAND& J.C. MORRISON& C.J. LOUCKS& T.F. ALLNUTT& dava& 19°55'06"S 44°36'00"E & 5 f& 22.II.2000 & fr.& Davis et al. 2587 ( BR & K &

MO & TAN); Kirindy Mitea NP& 20°53'14"S 43°55'22"E & 20 f& 21.III.2011 & fr . & T.H. RICKETTS& Y. KURA& J.F. LAMOREUX& W.W. WETTENGEL& Razafimandimbison & Kainulainen 1205 ( MO & S); N de Tsifafana env. 2 kf & P. HEDAO & K.R. KASSEM (2001). Terrestrial ecoregions of the 25.OE.1980& fr.& Service Forestier 29830 ( TEF). world: a new fap of life on Earth. BioScience 51: 933–938 .

SCHATZ& G.E. (2000). Endefisf in the Malagasy tree flora. In: Ackowledgements LOURENÇO& W.R. & S.M. GOODMAN (ed.)& Diversity and Endemism in Madagascar: 1–9. Méfoires de la Société de Bio- I thank the curators of G& K& MO& P& S& TAN& TEF& and UPS géographie& Paris.

for providing access to their collections& and Laurent Gautier&

Mitia Hanitrarivo& the late Jean-Noël Labat& Sven Landrein& SCHATZ& G.E. (2001). Generic tree flora of Madagascar. Royal Botanic Peter Phillipson& Landy Rita Rajaovelona& Solotiana D. Rafan- Gardens& Kew.

difbifanana& Richard Randrianaivo& Monique Randriatsivery&