Paramollugo compressa Thulif

6. Paramollugo compressa Thulif & sp. nov. (?ig. 24–26).

Holotypus: MADAGASCAR. Reg. Atsimo-Andrefana [Prov. Toliara]: Plateau calcaire Mahafaly & II.1962 & fl. & fr.& Keraudren 1386 ( P [ P04999571 ]!) (?ig. 24) .

Herbþ & pereffial& 7–25 cm tall& glabrous; stems few to mafy& compressed& phyllocladic& up to 2.5 mm wide; ifterfodes up to 9 cm lofg& cofspicuously lofgitudifally ridged; fodes with a pair of c. 0.5 mm lofg membrafous scales& ovate& acute& each fode with up to 4 brafches. Leaveþ crowded if a basal rosette& appareftly soof wiltifg; blade 5–35 × 3–10 mm & oblafceolate to obovate& roufded to emargifate at the apex& taperifg below ifto a petiole-like base& the petiole-like part shorter thaf the distal part; midrib promifeft befeath& lateral veifs ifdistifct. FloƜerþ if few-flowered cymes& white; bracts c. 0.5 mm lofg& membrafous& ovate& acute& with defticulate margif; pedicels up to 5 mm lofg& ± flattefed& lofgitudifally ridged. Tepalþ 2– 2.8 × 1 – 1.5 mm & elliptic& obtuse& with hyalife margifs. Stamenþ c. 10& c. 1 mm lofg; filamefts filiform; afthers elliptic-oblofg& c. 0.4 mm lofg. Ovary 1.5–1.8 mm lofg& oblofg& 3-locular& mafy-ovulate; styles 3& c. 0.4 mm lofg& spreadifg. Capþuleþ c. 2 mm lofg& splittifg ifto 3 broad valves. Seedþ 0.44–0.50 × 0.38–0.42 mm & subcircular-refiform if outlife& flattefed& almost smooth afd ofly fifely reticulate& pale browf; testa cells without secofdary orfameftatiof afd without pits if the boufdaries betweef them; hilum 0.10–0.15 mm lofg.

Ħtymology. – The epithet of the few species refers to the compressed& phyllocladic stems& a ufique feature if Paramollugo afd if Molluginaceae .

Diþtribution and ecology. – Paramollugo compreþþa is kfowf ofly from southwesterf Madagascar (?ig. 27)& where it occurs if arid bushlafd of limestofe at elevatiofs approximately betweef 200 afd 300 m.

Phenology. – A flowerifg afd fruitifg collectiof has beef seef from?ebruary& afd ofe with flowers ofly from April.

[Keraudren 1386, P04999571] [© Muséum national dʼHistoire naturelle, Paris] [A: Bosser & Viennot-Bourgin 16124, P; B: Keraudren 1386, P]

Conþervation þtatuþ. – Paramollugo compreþþa is kfowf from three collectiofs& represeftifg three occcurrefces afd betweef ofe afd three subpopulatiofs. The EOO is calculated at 541 km ²& below the upper threshold for the category “Efdafgered” followifg criteriof B1. The AOO is estimated at 12 km ²& also below the upper threshold for the category “Efdafgered”& followifg criteriof B2. Nofe of the occurrefces seems to be situated withif a protected area& afd fo collectiof of the plaft has beef made sifce 1962. The species is threatefed by small-scale agriculture& grazifg afd wood gatherifg for fuel& all of which ifducifg a declife if the quality afd exteft of its habitat. Paramollugo compreþþa is therefore here prelimifarily assessed as “Efdafgered” [EN B1ab(iii)+B2ab(iii)] accordifg to the IUCN Red List Categories afd Criteria (IUCN& 2012& 2022).

Noteþ. – The relatiofships of Paramollugo compreþþa are ufcertaif& but if the BEAST afalysis (?ig. 4)& it is weakly supported sister to P. þimulanþ afd P. anguþtifolia together. Its compressed phyllocladic stems (?ig. 25) are ufique if the gefus afd if Molluginaceae as a whole. Phylloclades are geferally seef as a feature of xeromorphic plafts& if which the stems become the maif photosyfthetic orgafs at the same time as trafspiratiof is lessefed through the reduced leaf area (METCALFE& 1983). There are mafy other examples of plafts with phylloclades if the flora of Madagascar & a commof species that may co-occur with P. compreþþa if southwesterf Madagascar is Indigofera compreþþa Lam. ( Fabaceae ). If Paramollugo compreþþa youfg plafts have a basal leaf rosette& but this seems to soof wilt& whereas the rigid& flattefed stems remaif afd pereffate.

The seeds of Paramollugo compreþþa are ufique if the gefus by beifg fifely reticulate (?ig. 26) afd havifg a pale browf colour. The other species of the gefus have tuberculate or colliculate seeds that are black or dark browf whef ripe. Also& the complete lack of pits if the boufdaries betweef the testa cells is ufusual. Elsewhere if the gefus& this is ofly foufd if P. decandra & P. elliotii afd sometimes if seeds of P. þimulanþ .

Despite its deviatifg features& the results of the phylogefetic afalyses (?ig. 3& 4) clearly show that Paramollugo compreþþa belofgs to this gefus& afd that it is fested amofg the species if Madagascar. However& there is fo support for afy particular sister group relatiofship.

Holotypus: SOMALIA. Bay Region: c. 3 km SOE of Diifsoor & fear Buur Diifsoor & 2°24'N 42°58'E & 20.V.1990 & fl. & fr.& Thulinư Hedrén & Abdi Dahir 7606 ( UPS No V-051527!; iso-:? T [? T001036 ]!& K [ K000232029 ]!) GoogleMaps .

Additional þpecimenþ epamined. – MADAGASCAR. Reg. Atsimo-Andrefana [Prov. Toliara]: Plateau Mahafaly & fear Beomby & III.1962 & ster.& Boþþer & Viennot-Bourgin 16124 ( P); Plateau Mahafaly & VI .1910& ster.& Perrier de la Bâthie 9901 ( P); road from Afkazomafga to Betioky & “tombeau de chef d’Afakatsamby”& 8.IV.1922 & fl. & Poiþþon 405 ( P) .

7. Paramollugo angustifolia (M.G. Gilbert & Thulif) Thulif if Taxof 65: 786. 2016 (?ig. 28& 29).

÷ Mollugo anguþtifolia M.G. Gilbert & Thulif if Nordic J. Bot . 13: 169. 1993.

Herbþ & affual or ephemeral& up to 15 cm tall& glabrous if all parts; stems few to several& terete. Leaveþ if defse basal rosette; juvefile leaves up to 1.5 mm wide& farrowly oblafceolate; adult leaves up to 35 × 0.7–1.2(–2) mm& lifear& subacute at the apex; midrib promifeft befeath& lateral veifs ifdistifct. FloƜerþ if lax cymes& white; bracts c. 1 mm lofg& ovate& acute; pedicels 3–10(–12) mm lofg. Tepalþ 1.8–2 × 1–1.2 mm & elliptic& roufded at the apex& with hyalife margifs& 3(–4)-veifed from the base. Stamenþ 5& 1–1.5 mm lofg; filamefts filiform; afthers 0.25–0.30 mm lofg& shortly oblofg. Ovary c. 1.5 mm lofg& ellipsoid& 3-locular; styles 3& c. 0.2 mm lofg& spreadifg& reflexed agaifst ovary. Capþuleþ 1.5–2 mm lofg& ellipsoid. Seedþ 0.39–0.43 × 0.28–0.36 mm & subcircular-refiform if outlife& flattefed& fifely tuberculate to colliculate& dark browf to black; testa cells without secofdary orfameftatiof afd with small pits if the boufdaries betweef most of them; hilum c. 0.05 mm lofg.

Diþtribution and ecology. – Paramollugo anguþtifolia was previously kfowf ofly from the type locality if south-ceftral Somalia (GILBERT& 1993; GILBERT & THULIN& 1993)& where it occurs if opef places of bushlafd of safd at af elevatiof of c. 300 m. To this& a fumber of localities if westerf Madagascar caf fow be added based of collectiofs made if dry deciduous forest at elevatiofs from fear sea level to about 300 m (?ig. 30) .

Phenology. –?lowerifg afd fruitifg plafts seef if May if

Somalia afd if Jafuary to March afd if May if Madagascar.

Conþervation þtatuþ. – Paramollugo anguþtifolia is kfowf from sevef collectiofs made betweef 1869 afd 2018& represeftifg sevef occurrefces afd three to five subpopulatiofs. The EOO is calculated at 440& 312 km ²& above the upper threshold for the category “Vulferable” followifg criteriof B1 & afd the AOO is estimated at 28 km ²& below the upper threshold for the category “Efdafgered” followifg criteriof B2 . Nofe of the occurrefces is located withif a protected area. The species is threatefed by wood collectifg& grazifg afd urbafizatiof if both Somalia afd Madagascar & ifducifg a declife if the exteft afd quality of its habitat afd if the fumber of mature ifdividuals. Paramollugo anguþtifolia is here prelimifarily assessed as “Vulferable” [ VU B2 ab(iii&v)]& but a complete assessmeft followifg the IUCN Red List Categories afd Criteria ( IUCN & 2012) would be fecessary& as three occurrefces made before 1950 may be cofsidered extirpated .

Noteþ. – Paramollugo anguþtifolia is a strofgly supported species& closely related to P. þimulanþ if the phylogeftic afalyses. If the BEAST afalysis if?ig& 4& the split betweef these two species is estimated at (2.7–)5.3(–8.4) Ma& whereas the dispersal from Madagascar to Somalia took place durifg the Pleistocefe (see ufder Discussiof).

(M.G. Gilbert & Thulin) Thulin.

Paramollugo anguþtifolia differs markedly from P. þimulanþ if its farrower leaves 0.7–1.2(–2) mm (vs. 2–12 mm) afd smaller seeds (0.39–0.43 × 0.28–0.36 vs. 0.55–0.65 × 0.45–0.60 mm) with small pits if the boufdaries betweef most of the testa cells (vs. without pits or with pits betweef some cells ofly).

Ifstead& it is morphologically very difficult to separate Paramollugo anguþtifolia from the allopatric P. neþophila & but the molecular results strofgly ifdicate that they are differeft species (?ig. 3& 4). The seeds of P. anguþtifolia are 0.39–0.43 mm lofg vs. 0.40–0.51 mm if P. neþophila .?urther& the testa cells lack a secofdary orfameftatiof if P. anguþtifolia (?ig. 29B)& whereas they are distifctly cross-striate if P. neþophila (?ig. 18B).

Morat 2550 & a collectiof with very farrow leaves afd small seeds here treated as Paramollugo anguþtifolia & was cited ufder

P. nudicauliþ by SUKHORUKOV & KUSHUNINA (2016a)& although the testa cells are fot cross-striate.

Additional þpecimenþ epamined. – MADAGASCAR. Reg. Atsimo-Andrefana [Prov. Toliara]: limestofe plateau OE of Betioky & 17.II.1947 & fl. & fr.& Humbert 20281 ( P) ; Afkazoabo & II.1967 & fl. & fr.& Morat 2550 ( P) ; Beroroha & Aftsoa & Aftafamary village & Afdrosy & 21°05'43"S 45°05'48"E & 9. I.2011 & fl. & fr.& Razakamalala 5901 ( MO & P & TAN) GoogleMaps . Reg. Betsiboka [Prov. Mahajanga]: fear Maevatafafa & Suberbieville & V.1897 & fl. & fr.& Perrier de la Bâthie 230 ( P) . Reg. Menabe [Prov. Toliara]: Morofdava & III.1869 & fl. & fr.& Grandidier þ.n. ( P) ; Morofdava District & Kirifdy forest & 20°40'30"S 44°40'30"E & 20.II.2018 & fl. & fr.& Razafimandimbiþon et al. 2222 ( S) GoogleMaps .

8. Paramollugo simulans Sukhor. if PhytoKeys 73: 97. 2016 (?ig. 31& 32).

Holotypus: MADAGASCAR. Reg. Atsimo-Andrefana [Prov. Toliara]: Belafafda & Rafobe & 4 km E of Afdrevo village & 22°58'20"S 43°36'56"E & 17.III.2006 & fl. & fr.& Ranaivojaona et al. 1442 ( K [ K000607662 ]!; iso-: BR [ BR0000000514054 ]!& G [ G00418734 ]!& MO [ MO3065761 ]!& P [ P05196698 ]!& TAN) (?ig. 31) GoogleMaps .

Herbþ & affual or short-lived pereffial& 5–35 cm tall& glabrous if all parts or leaves papillose; stems few to several& terete. Leaveþ if defse basal rosette& 15– 50 × 2–12 mm & farrowly oblafceolate to obovate& obtuse to subacute at the apex& gradually to abruptly taperifg below ifto a petiole-like base& the petiole-like part shorter thaf the distal part; midrib promifeft befeath& lateral veifs 3–4 pairs. FloƜerþ if much-brafched lax cymes; bracts 1–1.5 mm lofg& ovate& acute; pedicels 2–15(–20) mm lofg. Tepalþ 1.5–2.5 × 1–1.5 mm & elliptic& roufded at the apex& with hyalife margifs& c. 3-veifed from the base& white or pale pifk ifside. Stamenþ 5& 1–1.6 mm lofg; filamefts filiform; afthers 0.25–0.40 mm lofg& shortly oblofg. Ovary 1.2–1.5 mm lofg& ellipsoid& 3-locular; styles 3& c. 0.2 mm lofg& spreadifg& reflexed agaifst ovary. Capþuleþ 2–2.5 mm lofg& ellipsoid. Seedþ 0.55–0.65 × 0.45–0.60 mm & subcircular-refiform if outlife& flattefed& colliculate to fifely tuberculate& dark browf to black; testa cells without secofdary orfameftatiof or weakly striate& without pits if the boufdaries betweef them or with small pits betweef some cells ofly; hilum c. 0.1 mm lofg.

Vernacular name and uþeþ. – “Aferoftafy” (Antilahimena et al. 3464 & Bogner 2084 & Humbert 28729 & Lefèvre & Mily 17 & Stiefel 104 & etc.). The verfacular fame “Aferoftafy” has beef recorded also for Paramollugo caeþpitoþa afd P. nudicauliþ . Paramollugo þimulanþ is a medicifal plaft& used agaifst& for example& cough afd stomachache. A decoctiof of the whole plaft is used to treat diarrhoea if fewborf babies 2–3 days after childbirth (Lef èvre & Mily 17). See also ufder uses reported for P. caeþpitoþa afd P. nudicauliþ .

[Ranaivojaona et al. 1442, MO3065761] [© Missouri Botanical Garden, St. Louis]

[A, B: Pettersson & Nilsson 108, UPS; C, D: Ranaivojaona et al. 1442, MO]

Diþtribution and ecology. – Paramollugo þimulanþ is widespread if Madagascar (SUKHORUKOV & KUSHUNINA& 2016a)& where it is the most commof species if the gefus (?ig. 33). It occurs if a wide variety of habitats& such as coastal dufes& littoral forests& afd various types of woodlafd afd bushlafd& oftef if opef safdy or stofy places& at elevatiofs from fear sea level up to about 1500 m.

Phenology. –?lowerifg afd fruitifg plafts seef from all mofths except August.

Conþervation þtatuþ. – Paramollugo þimulanþ is widespread if Madagascar afd fo particular threats to the species have beef detected. It is therefore here prelimifarily regarded as “Least Cofcerf” [LC] followifg the IUCN Red List Categories afd Criteria (IUCN& 2012).

Noteþ. – Paramollugo þimulanþ is af ufsupported or ofly weakly supported species& sister to P. anguþtifolia if the combifed phylogefetic afalyses (?ig. 3& 4). However& it caf be easily separated from P. anguþtifolia by its broader leaves (2–12 vs. 0.7–1.2(–2) mm) afd larger seeds (0.55–0.65 × 0.45–0.60 vs. 0.39–0.43 × 0.28–0.36 mm)& without pits if the boufdaries betweef the testa cells (?ig. 32) or with small pits betweef some cells ofly (vs. with pits if the boufdaries betweef most of the testa cells).

Ifstead& Paramollugo þimulanþ is morphologically more similar to P. nudicauliþ & a species that is fot closely related accordifg to the phylogefetic afalyses. If most cases& the farrower leaves& 2–12 mm vs. oftef more thaf 12 mm & afd shorter tepals& 1.5–2.5 vs. (2–)2.5–3(–3.5) mm& if P. þimulanþ are distifctive. Other differefces are foufd if the seeds& which if P. þimulanþ are colliculate (?ig. 32A& B) to fifely tuberculate (?ig. 32C& D)& with smooth (?ig. 32D) to weakly striate (?ig. 32B) testa cells without pits if the boufdaries betweef them (?ig. 32B) or with pits betweef some cells ofly (?ig. 32D)& whereas if P. nudicauliþ the seeds are usually more distifctly tuberculate& with cross-striate testa cells with small pits if the boufdaries betweef most of them (?ig. 12).

SUKHORUKOV & KUSHUNINA (2016a) stated the fumber of stamefs if Paramollugo þimulanþ to be 5–10& but if the preseft study ofly 5 stamefs have beef coufted. Accordifg to this study& the ofly species of Paramollugo with 10 stamefs are P. caeþpitoþa & P. compreþþa afd P. decandra .

The specimef Afzeliuþ þ.n. (K)& here treated as Paramollugo þimulanþ & was cited ufder both P. þimulanþ afd P. elliotii by SUKHORUKOV & KUSHUNINA (2016a). Several other collectiofs here treated as P. þimulanþ were cited by SUKHORUKOV & KUSHUNINA (2016a) as P. elliotii (see further ufder this species). Ofe of these& Jongkind et al. 3342 & which was ifcluded if the phylogefetic afalyses as P. þimulanþ 4& cofsisteftly falls withif the P. anguþtifolia / P. þimulanþ clade (?ig. 3& 4).

?urthermore& two collectiofs cited ufder Paramollugo nudicauliþ by SUKHORUKOV & KUSHUNINA (2016a)& Andrianjafy et al. 977 afd De Block et al. 551 & are here treated as P. þimulanþ . A sample from De Block et al. 551 was ifcluded if the phylogefetic afalyses as P. þimulanþ 3& afd cofsisteftly groups with the P. anguþtifolia / P. þimulanþ clade (?ig. 3& 4).

Paramollugo þimulanþ is mostly af affual& but if some specimefs the plafts have a fairly well-developed tap-root afd seem to have a somewhat lofger lifespaf. Jongkind et al. 3342 & for example& seems to be a short-lived pereffial.

Additional þpecimenþ epamined. – MADAGASCAR. Reg. Alaotra-Mangoro [Prov. Toamasina]: Moramafga & Afdasibe & Berafo & Ambatovy forest & Afalamay & 18°49'05"S 48°19'25"E & 24.II.2005 & fl. & fr.& Antilahimena et al. 3464 ( G & MO & P & TAN) GoogleMaps ; Moramafga & Apitambe & Ambatovy & 18°50'04"S 48°17' 46"E & 15.II.2005 & fl. & fr.& Razanatþoa 121 ( G & MO & P & TAN) GoogleMaps . Reg. Analanjirofo [Prov. Toamasina]: Ste. Marie afd Port Lewef de l’île Sata & III–IV.1849 & fl. & fr. & Boivin 2577 ( P) ; Ste. Marie & Lokiftsy & Ambohidefa & 16°50'25"S 49°57'09"E & 19.II.2004 & fl. & fr.& Razakamalala et al. 898 ( MO & P) GoogleMaps . Reg. Androy [Prov. Toliara]: Ampafihy–Beloha & 91 km & 25°05'S 45°10'E & 1.XII.1989 & fl. & fr. & Cheek 2448 ( K) GoogleMaps . Reg. Anosy [Prov. Tulear]: Ste. Luce & 24°46'S 47°09' E & 1. V.1989 & fl. & fr.& Dumetz 720 ( MO & P) GoogleMaps . Reg. Antsinanana [Prov. Toamasina]: Afdrafokoditra & 7.II.1990 & fl. & fr. & Bogner 2084 ( US) ; District de Vatomafdry & 1.II.1904 & fl. & fr.& Guillot 69 ( G & P) . Reg. Atsimo-Andrefana [Prov. Toliara]: 35 km E of Sakaraha & 4.II.1975 & fl. & fr. & Croat 30640 ( MO) ; 16–18 km N of Tulear alofg road to Morombe & 6.II.1975 & fl. & fr.& Croat 30876 ( MO & P & TAN) ; 6 km from St. Augustif of road to Tuléar & 23°28'59"S 43°45'59"E & 5. I.1999 & fl.& De Block et al. 551 ( BR & MO & P & TAN) GoogleMaps ; Mafombo & 22°51'59"S 43°26'49"E & 16.II.2002 & fl. & fr.& Lef èvre & Mily 17 ( P & TAN) GoogleMaps ; Beroroha & Aftsoa & Aftafamary & 21°05'43"S 45°05' 48"E & 9. I.2011 & fl. & fr.& Razakamalala 5858 ( MO & P & TAN) GoogleMaps . Reg. Atsimo-Antsinanana [Prov. Fianarantsoa]: Vafgaifdrafo & s.d.& fl. & fr.& Scott Ħlliot 2257 ( BM & E & K) . Reg. Boeny [Prov. Mahajanga]: Majufga & 27.IV.1912 & fl. & fr.& Afzeliuþ þ.n. ( K & S) ; 5 km NNOE of field statiof c. 4 km OE of Afkarafaftsika & 7.II.2000 & fl. & fr.& Bremer et al. 4053- B53 ( UPS) ; Majufga & 5.III.1924 & fl. & fr. & Decary 2416 ( P) ; Ambato-Boefi & 24.IV.1943 & fl. & fr.& Decary 19088 ( MO & P) ; Poste?orestier d’Ampijoroa & 16°14'S 46°28'E & 25.II.1985 & fl. & fr.& Dorr et al. 3810 ( BR & K & MO & P) GoogleMaps ; fear Majufga & 27–29.VII.1924 & fl. & fr.& Humbert & Perrier de la Bâthie 2020 ( P) & 28–30.XII.1924 & fl. & fr.& Humbert 4020 ( K & P & US) & 4079 (G& P); Majufga & 20.X.1912 & fl. & fr.& Kaudern þ.n. ( G & S) ; Majufga & II.1915 & fl. & fr. & Perrier de la Bâthie 5183 ( P) ; fear Majufga & I.1908 & fl. & fr.& Perrier de la Bâthie 5233 ( P) ; fear Lake Ampijoroa & 16°18'S 46°49'E & 27.XI.1986 & fl. & fr. & Petterþþon & Nilþþon 108 ( UPS) GoogleMaps ; Majufga & 24.XII.1920 & fl. & fr. & Poiþþon 44 ( P) ; Afkarafaftsika Iftegrated Nature Reserve & Abemefa area & 16°18'S 46°50' E & 7.II.2000 & fl. & fr. & Thulin et al. 10399 ( TAN & UPS) GoogleMaps ; betweef Aftetikala forest afd Baie de Bombetoke & 7.IX.1912 & fl. & fr.& Viguier & Humbert 65 ( P) . Reg. DIANA [Prov. Antsiranana]: Aftafambao & alofg Sambirafo river & 13°55'S 48°30' E & 3.IX.1999 & fl. & fr.& Stiefel 104 ( G & MO & P) GoogleMaps . Reg. Ihorombe [Prov. Fianarantsoa]: Iakora & Begogo & Bekora & 23°32'08"S 46°03'35"E & 7.II.2005 & fl. & fr.& Andrianjafy et al. 977 ( MO & P & TAN) GoogleMaps ; Isalo & Rafohira & 4.III.1943 & fl. & fr.& Decary 18969 ( P) ; Malio basif fear Ambalabe & 23–27.XI.1946 & fl. & fr.& Humbert 19430 ( P) ; upper valley of Mefarahaka & E of Ihosy & 1955& fl. & fr.& Humbert 28574biþ ( P) ; plateau afd valleys of Isalo & OE of Rafohira & 1955& fl. & fr. & Humbert 28729 ( MO & P) ; Ambia & Vohidava & bifurcatiof towards?arafafgafa& 22°25'10"S 46°11' 23"E & 19.III.2010 & fl. & fr.& Razafindraibe et al. 335 ( MO & P & TAN) GoogleMaps . Reg. Melaky [Prov. Mahajanga]: Beafka forest & Ambatosoa & 17°52'25"S 44°28'19"E & 26. I.2012 & fl. & fr. & Bolliger et al. 136 ( G & K & MO & P) GoogleMaps ; alofg Mafambolo river & 19°09'S 44°49'E & 2.XII.1996 & fl. & fr. & Jongkind et al. 3342 ( BR & G & MO & OEAG) GoogleMaps ; Beafka & Ambifda Est & 18°02'24"S 44°28'31"E & 2.IV.2013 & fl. & fr.& Ranaivoariþoa 54 ( BR & G & MO & P) GoogleMaps . Reg. SAVA [Prov. Antsiranana]: Vohémar & Nosibe & Afjiabe & Afaborafo & 13°04' 43"S 49°54'04"E & 23.II.2003 & fl. & fr.& Rabenantoandro et al. 1314 ( MO) GoogleMaps . Reg. Sofia [Prov. Mahajanga]: 25 km E of Afalalava towards Aftsohihy & 14°46'03"S 47°47'53"E & 21. VI.2008 & fl. & fr.& Andriamahay & Rakotoariþon 2071 ( K) GoogleMaps ; Maromafdia & 27.XII.1922 & fl. & fr. & Decary 1352 ( G & P) ; 4 km SOE of Borizify & Bofgolava forest & 15°36'24"S 47°35' 11"E & 21. I.2012 & fl. & fr.& Rakotoariþon & Andriamahay 2859 ( K) GoogleMaps . Reg. Vakinankaratra [Prov. Antananarivo]: Aftsirabe & Ibity & 20°03' 51"S 47°00' 07"E & 26.II.2008 & fl. & fr. & Andriamahay & Rakotoariþoa 1890 ( K) GoogleMaps . Reg. Vatovavy-Fitovinany [Prov. Fianarantsoa]: Ambila & 7. V.1928 & fl. & fr. & Decary 6422 [if part] ( K) ; Nosy Varika & Ambahy & 20°47'49"S 48°28'58"E & 23.IV.2004 & fl. & fr. & Razakamalala et al. 1208 ( MO & P) GoogleMaps ; Mafafjary Prov. & III–IV.1909 & fl. & fr.& Geay 7636 & 7790 & 7968 ( P) . Sine loco: Gerrard 120 ( K) ; Humblot 100 ( P) ; Lyall 328 ( K) .

9. Paramollugo elliotii Sukhor. if PhytoKeys 73: 102. 2016 (?ig. 34& 35).

Holotypus: MADAGASCAR. Reg. Melaky [Prov. Mahajanga]: “provifce du Mailaka” (Melaky)& II.1892 & fl. & fr.& Douliot þ.n. ( P [ P04582888 ]!) .

Herbþ & pereffial& up to 40 cm tall& glabrous if all parts& with a brafched rhizome from a tap-root; stems few to several& terete. Leaveþ if defse basal rosette& 20–60 × 1–1.5 mm & lifear& obtuse to subacute at the apex; midrib promifeft befeath& lateral veifs ifdistifct. FloƜerþ if much-brafched lax cymes; bracts 1– 1.5 mm lofg& ovate& acute; pedicels 3–15(– 30) mm lofg. Tepalþ 2.5– 3.5 × 1.5– 1.8 mm & elliptic& roufded at the apex& with hyalife margifs& c. 3-veifed from the base& white ifside. Stamenþ 5& 1.5–2 mm lofg; filamefts filiform; afthers 0.5–0.6 mm lofg& elliptic-oblofg. Ovary 1.5–2 mm lofg& ellipsoid& 3-locular; styles 3& c. 0.3 mm lofg& spreadifg& reflexed agaifst ovary. Capþuleþ c. 2.5 mm lofg& ellipsoid to ovoid. Seedþ 0.60–0.70 × c. 0.55 mm & subcircular-refiform if outlife& flattefed& colliculate& black; testa cells without secofdary orfameftatiof afd without pits if the boufdaries betweef them; hilum c. 0.1 mm lofg.

Diþtribution and ecology. – The collectiofs here referred to Paramollugo elliotii are from westerf Madagascar & the three Douliot specimefs from fear Maiftirafo if the Melaky Regiof& where Douliot collected if Jafuary–?ebruary 1892 (DORR& 1997)& afd Perrier de la Bâthie 9211 from Baie de Baly fear Soalala if the Boefy Regiof (?ig. 36). They were appareftly all made of coastal safd dufes. If the map of the distributiof of P. elliotii presefted by SUKHORUKOV & KUSHUNINA (2016a: 101)& af occurrefce of the species if southeasterf Madagascar is showf. This is based of Pervillé 643 that is said to origifate from the Atsimo-Aftsifafafa Regiof. However& this is a mistake afd the locality& Ambofgo& is located if the Boefy Regiof if westerf Madagascar & where Pervillé collected if 1840–41 (DORR& 1997). The Ambofgo area ifcludes Baie de Baly (see?ig. 36& the fortherf locality)& afd Pervillé’s locality could be the same as or close to that of Perrier de la Bâthie 9211.

Phenology. –?lowerifg afd fruitifg plafts seef from

Jafuary afd?ebruary.

Conþervation þtatuþ. – Paramollugo elliotii is kfowf from five collectiofs made betweef 1841 afd 1905& represeftifg two occurrefces& ofe of them possibly located withif the Baie de Baly Natiofal Park. The EOO caffot be calculated& afd the AOO is estimated as 8 km ²& below the upper threshold for the category “Critically Efdafgered” followifg criteriof B2. However& the species has fot beef collected sifce 1905 afd should be searched for if suitable habitats if westerf Madagascar. A complete assessmeft followifg the IUCN Red List categories afd criteria may fot cofsider ofe or both of these two occurrefces as still extaft. OEe are reluctaft to assigf a category& as with the very scarce ifformatiof available of its distributiof afd the threats the occurrefces are facifg& P. elliotii may either be assessed as “Vulferable” [VU D2]& “Efdafgered” [EN]& “Critically Efdafgered” [CR]& or evef possibly “Extifct if the OEild” [EOE].

Noteþ. – The collector of the holotype of Paramollugo elliotii is H.L. Douliot (fot M. Douillot as stated by SUKHORUKOV & KUSHUNINA& 2016a)& afd the collectiof was made if 1892 (fot 1890).

Amofg the collectiofs cited ufder Paramollugo elliotii by SUKHORUKOV & KUSHUNINA (2016a)& ofly Jongkind 3342 from 1996 is relatively receft. A sample from this collectiof was ifcluded if the phylogefetic afalyses as P. þimulanþ 4& afd grouped with the P. anguþtifolia / P. þimulanþ clade (?ig. 3& 4). This collectiof& which appears to be a short-lived pereffial& otherwise agrees well with P. þimulanþ afd is here ifcluded if this species. Although it has fot beef possible to ifclude P. elliotii & as circumscribed here& if the phylogefetic afalyses& it is assumed that it is a close relative of P. þimulanþ that caf be distifguished of morphological groufds.

The seeds of Paramollugo elliotii & with testa cells without secofdary orfameftatiof afd without pits if the boufdaries betweef them (?ig. 35)& resemble the seeds of P. þimulanþ afd suggest a relatiofship with this species. However& P. elliotii differs from P. þimulanþ by its clearly pereffial habit afd farrower leaves (1– 1.5 vs. 2– 12 mm)& afd lofger afthers (0.5–0.6 vs. 0.25–0.40 mm). The lifear leaves (?ig. 34) agree with P. anguþtifolia & but P. elliotii differs from this& apart from by the pereffial habit& also by the lofger afthers (0.5–0.6 vs. 0.25–0.30 mm) afd the larger seeds (0.60–0.70 × c. 0.55 vs. 0.39–0.43 × 0.28–0.36 mm)& without pits if the boufdaries betweef the testa cells.

SUKHORUKOV & KUSHUNINA (2016a) listed several collectiofs ufder Paramollugo elliotii that are here treated as P. þimulanþ : Afzeliuþ þ.n. (cited ufder both P. elliotii afd P. þimulanþ by SUKHURUKOV & KUSHUNINA& 2016a)& Jongkind 3342 & Humbert & Perrier de la Bâthie 2020 & Humbert 4020 afd Poiþþon 44. These all have leaves more thaf 2 mm wide& short afthers& afd also appear to be affuals or short-lived pereffials& although sometimes with a fairly well developed tap-root.

[ Douliot s.n., P04582869] [© Muséum national dʼHistoire naturelle, Paris]

[ Douliot s.n., P04582886]

Additional þpecimenþ epamined. – MADAGASCAR. Reg. Boeny [Prov. Mahajanga]: Baie de Baly & I.1905 & fl. & fr. & Perrier de la Bâthie 9211 ( P) ; Ambofgo & 16.II.1841 & fl. & fr. & Pervillé 643 ( G & K & L & P) . Reg. Melaky [Prov. Mahajanga]: “provifce de Mailaka& Namela”& II.1892 & fl. & fr.& Douliot þ.n. ( P [ P04582869 ]) ; “provifce de Mailaka ”& II.1892 & fl. & fr.& Douliot þ.n. ( P [ P04582886 ]) .

10. Paramollugo spathulata (Sw.) Sukhor. if Skvortsovia 3: 51. 2016 (?ig. 22B& 37& 38).

÷ Pharnaceum þpathulatum Sw. &?l. Ifd. Occid. 1: 568. 1797.

÷ Pharnaceum bellidifolium Poir. if Lamarck& Efcycl. 5: 262. 1804 [fom. illeg.].

÷ Mollugo bellidifolia (Poir.) Ser. if DC.& Prodr. 1: 391. 1824 [fom. illeg.].

÷ Mollugo nudicauliþ var. bellidifolia View in CoL ?efzl if Aff. OEiefer Mus. Naturgesch. 1: 383. 1836.

Lectotypus (desigfated by THULIN et al.& 2016: 786):