Mendoncia flagellaris (Baker) Benoist, Bull. Mus. Hist. Nat.

Magnaghi, Emily B. & Daniel, Thomas F., 2017, Systematics of Mendoncia (Acanthaceae: Thunbergioideae) in the Paleotropics, Proceedings of the California Academy of Sciences 64 (3), pp. 37-94 : 66-70

5. Mendoncia flagellaris (Baker) Benoist, Bull. Mus. Hist. Nat. (Paris). 31:387. 1925.

Monachochlamys flagellaris Baker, J. Linn. Soc., Bot. 20:217. 1883 (“1884”).

TYPE.— MADAGASCAR. Province unknown: Central Madagascar , Oct 1882 (fide specimen at K), Baron 1743 (syntypes: K-000393667 -photo!, P-00091104 !); Baron 1789 (syntypes: K-000393668 -photo!, P-00091103 !).

Mendoncia madagascariensis Radlk. , Abh. Naturwiss. Vereins Bremen 8:467. 1884. Monachochlamys madagascariensis (Radlk.) Baill. , Bull. Mens. Soc. Linn. Paris 2:826. 1890 (as “ madagascarica”).

TYPE.— MADAGASCAR. Antananarivo: Ost-Imerina, Andrangolaoka [as “Andrangolóaka;” ca. 19°02ʹ15.26ʺS, 047°54ʹ57.70ʺE], Nov 1880, Hildebrandt 3693 (syntypes: HBG-517341 -photo!, JE-00003605 -photo!, JE-00003606 -photo!, JE-00003607 -photo!, K-000393669 -photo!, K-000393671 -photo!, M-0186545 -photo!, M-0186546 -photo!, P-00493833 !, WU-035583 -photo!) .

Monachochlamys boivinii Baill. , Bull. Mens. Soc. Linn. Paris 2:826. 1890.

TYPE.— MADAGASCAR. Antsiranana: Nossi-be, cratère d’Ampombilava [13°23ʹ35.06ʺS, 048°14ʹ48.74ʺE], 1850 (fide specimen), Boivin s.n. (holotype: P-00493802 !)

Young stems glabrous (at least internodes) to mealy-glandular to sparsely (to densely) pubescent with antrorse to antrorsely appressed, straw-colored, unbranched, eglandular trichomes 0.1– 0.5 mm long (or rarely densely pubescent with erect to flexuose to recurved trichomes to 1 mm long, i.e., pilose), nodes sparsely to densely pubescent with stiff, straw-colored, mostly appressed, eglandular, unbranched trichomes to 1 mm long (sometimes deciduous on older stems). Leaves membranaceous to coriaceous, petioles to 27 mm long, glabrous to pubescent with erect to flexuose trichomes like those at nodes (or pilose), blades ovate to narrowly elliptic to broadly elliptic to subcircular to oblong to obovate to obcordate, 14–137 mm long, 7–102 mm wide, truncate to rounded to cuneate at base, acuminate to acute to rounded to truncate to retuse and often apiculate (with apiculum to 1.6 mm long) at apex, abaxial surface glabrous to sparsely pubescent with antrorse to antrorsely appressed trichomes 0.1– 0.5 mm long (or pilose), adaxial surface glabrous to mealy-glandular and occasionally with eglandular trichomes like those of abaxial surface (or pilose), foliar trichomes sometimes restricted to major veins. Inflorescences borne in leaf axils on young, mostly herbaceous, and leafy stems, solitary or opposite at nodes, consisting of axillary, pedunculate dichasia or of a pedunculate, bracteate raceme bearing pedunculate dichasia in axils of bracts, dichasia 1–4 (–8) per axil, peduncles to 40 (–55) mm long, glabrous to sparsely pubescent like young stems, racemes (when present) to 15 cm long, rachis pubescent like young stems, bracts of racemes subfoliose and reduced in size acropetally, 2–40 mm long, 0. 8–14 mm wide. Bracteoles green to greenish white, mostly deciduous in fruit, ovate to elliptic to oblong to obovate, 4.3– 11. 2 mm long, 2. 5–10 mm wide, rounded to cuneate to subtruncate at base, rounded to acute to acute-apiculate to subacuminate at apex, abaxial surface glabrous to mealy-glandular or sparsely to densely pubescent with straw-colored, unbranched, mostly antrorsely appressed, eglandular trichomes to 0.5 mm long (or pilose), basally often beset with a conspicuous pair of whitish, gall-like, bulbous protuberances up to 2.5 mm high and 1.5– 3 mm in diameter, adaxial surface mealy-glandular or ± densely beset with larger, sessile glands, distal margin and apex of adaxial surface inconspicuously but densely pubescent with flexuose, eglandular trichomes to 0.3 mm long. Calyx glabrous to mealy-glandular (or sometimes with a fringe of trichomes 0.1– 0.2 mm long, e.g., Baron 3749). Corolla white to white with purple markings near base of lower lip, 8–15 (–18) mm long, externally glabrous to mealy-glandular, internally glabrous to mealy-glandular (especially in throat near mouth), tube (3–) 5–12. 7 mm long, limb 4.5–29. 1 mm in diameter, upper lip 2–12. 5 mm long with rounded to emarginate lobes 1.4–11. 6 mm long and 1.3–11. 1 mm wide, lower lip 2.3–16. 7 mm long with rounded to emarginate lobes 1.7–14. 5 mm long and 2. 5–9 mm wide. Stamens 2.8– 5.9 mm long, ventral pair inserted up to 1 mm distal to dorsal pair, thecae 1.1–4. 4 mm long, pubescent at base with a tuft of bristles 0.1– 0.2 mm long, these sometimes extending toward apex of thecae and thecae sometimes minutely glandular dorsally, connective not extending or extending up to 0.5 mm beyond thecae, extension of connective triangular to oblong, glabrous to minutely glandular, staminode (if present) 1.3– 2 mm long, thecae (when present) up to 0.9 mm long. Pollen 4 (–5)- colpate, E = 25–32 µm, P = 23–27 µm, P:E = 0.812–0.979, C = 5.8–9.1, C: P = 0.224–0.362 µm, rugulae microverrucate to microgemmate. Style 3–8. 8 mm long, glabrous, stigma subcapitate with undulating margin/rim to unequally 2-lobed. Drupe ovoid-ellipsoid to ellipsoid to spherical to obovoid, 9. 5–20 mm long, 6–15. 7 mm diameter, glabrous to mealy-glandular. n = 19 ( Daniel 2006).

PHENOLOGY.— Flowering: October–March; fruiting: year-round especially during July– March.

DISTRIBUTION AND HABITATS.— Throughout northern and eastern Madagascar (Antsiranana, Fianarantsoa, Toamasina, Toliara) and known from a few localities in central and west-central portions of the island (Antananarivo, Fianarantsoa, Mahajunga); also on Mayotte in the Comoros Archipelago (EOO = 458,376 km 2; Fig. 12 ). Plants occur in lowland and montane, primary and secondary, moist to wet, evergreen forests; degraded secondary forests; gallery forests; dry deciduous forests; and coastal forests. Elev. 5–2000 m.

ILLUSTRATIONS.— Baker (1883:t. 26); Benoist (1967:5, Fig. 1 ); Figs. 11E, F and 13.

LOCAL NAMES.— “Mbarahanjelinala” (Razafitsalama et al. 136); “vahimbarongy” (Kotozafy et al. 372); “vahivarongy” (Kotozafy 125).

Morphological variation within and among the many collections of M. flagellaris is apparent in size, shape, texture, and pubescence of leaf blades; bracteolar size, shape, and protuberance presence/absence; inflorescence development (dichasia in leaf axils vs. dichasia borne in axils of bracts on elongate racemes from leaf axils); and drupe size and shape. Dorr et al. 3697 is atypical by having stems, leaves, peduncles, and abaxial surface of the bracteoles densely pubescent (pilose), as noted in the description above. Other collections (e.g., van der Werff et al. 12665) are somewhat intermediate between this collection and more typical ones by having more abundant trichomes, although shorter and antrorse to antrorsely appressed. The presence or absence of a pair of conspicuous, bulbous, and white protuberances on the basal portion of the abaxial surface of bracteoles is also variable. This feature, when present, is unique in the genus. Collections from throughout the geographic range of the species vary from having a pair of these on each immature and mature bracteole, having the protuberances when young but with the bracteoles becoming smooth at maturity, or having no protuberances on the immature or mature bracteoles. Some specimens show variation in this character on the same sheet. Several collections from northern Madagascar (e.g., Andrianantoanina & Rocsceohclher 259, Andrianantoanina & Solotiana 74, Birkinshaw 127, Nicoll & Abraham 679, Rahajasoa et al. 202, Schatz et al. 1345) have conspicuously spherical drupes (in dried state). In other characters (including presence/absence of bulbous basal protuberances on bracteoles) these plants show variation like that in non-spherical-fruited specimens.

Rakotomalala & Narison 227 (14°05ʹS, 048°24ʹE; MO), from the Réserve Spécial de Manongarivo in Antsiranana Province, appears similar to M. flagellaris in most characters, but differs in having dense, erect to flexuose trichomes to 2 mm long dispersed along the proximal third of the midvein on the abaxial leaf surface or forming domatia in the axils of the midvein and secondary veins. It is possible that all of these trichomes are associated with the domatia. This collection also has longer corollas (25 mm long) than otherwise observed in M. flagellaris , a pubescent calyx (with eglandular trichomes to 2 mm long), and a proximally pubescent style. The flower color (white with purplish markings) appears to conform to M. flagellaris . Although lacking flowers and the denser pubescence of Rakatomalala & Narison 227, Gautier et al. 3512 (13°59ʹS 48°26ʹE; K, MO) from the same region, is otherwise similar and has what appear to be incipient domatia in the vein axils. The disposition of these collections requires additional study, and they are not treated as pertaining to M. flagellaris here. Other collections from the region of the Manongarivo Reserve (e.g., McPherson & van der Werff 16410, Rakotomalala & Narison 103) more closely conform to M. flagellaris , into which species they are placed here.

Two collections from Mt. Vato Vavy (21°24ʹS, 047°56ʹE) in eastern Fianarantsoa Province show similarities to M. flagellaris (e.g., white corollas with purplish markings), but appear to fall outside of the morphological variation of that species. Both collections represent fragments collected from the forest floor: Daniel et al. 9291 ( CAS) consists of dichasia from which the corollas have fallen, whereas Daniel et al. 9295 ( CAS), from a nearby site, consists of fallen corollas. Compared to M. flagellaris , the dichasia have mostly longer bracteoles (up to 16 mm long) and the corollas are larger in nearly all aspects (e.g., 24 to 34 mm long); thus both collections more closely resemble dichasia and corollas of M. vinciflora , which is currently known only from northern Madagascar. It remains unknown whether the corollas of the latter collection pertain to the dichasia of the former, and what characteristics the remainder of the plant(s) may have. Additional collections should resolve the identity or identities of these collections, which are not identified to species herein.

ADDITIONAL SPECIMENS EXAMINED.— MADAGASCAR. Antananarivo: Mandraka, D’Alleizette 1067 ( P) ; La Mandraka, R.N. 2, at p.k. 69, 18°55ʹS, 047°56ʹE, Dorr et al. 3697 ( MO) ; PK 64 entre Tananarive à Perinet , Keraudren-Aymonin & Aymonin 25314 ( P) ; Bongolava, W de Tsiroanomandidy , [ca. 18°52ʹ16.55ʹS, 45°42ʹ08.53ʺE], Morat 4645 ( P) ; Manankazo, NE de l’Ankazobe , [ca. 18°09ʹS, 047°14ʹE], Perrier de la Bathie 10247 ( MO, P) ; Forêt d’Andranomay, 13 km SE d’Anjozorobe et 2 km E d’Andranomay , 18°28ʹ48ʺS, 047°57ʹ18ʺE, Ranaivojaona 88bis ( MO) ; Ambohidratrimo, without collector, 435 ( P), s.n. ( P). Antsiranana: S d’Antsiranana, près de Joffre-Ville dans le PN de Montagne d’Ambre, campement de Ben Freed, 12°27ʹS, 049°13ʹE, Andrianantoanina & Rocsceohclher 259 ( BR, MO), 280, ( BR, MO) ; S d’Antsiranana, près de JoffreVille Parc Natl. de Montagne d’Ambre , campement de Chris, 12°35ʹS, 049°09ʹE, Andrianantoanina & Rocsceohclher 309 ( BR, MO) ; Montagne d’Ambre PN, 12°32ʹS, 049°07ʹE, Andrianantoanina & Solotiana 74 ( BR, CAS, MO) ; Montagne d’Ambre PN, 12°31ʹS, 049°09ʹE, Andrianantoanina et al. 123 ( CAS, MO) ; Nosy-Be, Réserve Intégrale de Lokobe , 13°24ʹ35ʺS, 048°19ʹ06ʺE, Antilahimena 95 ( MO) ; Réserve Naturelle Integrale de Tsaratanana, following Ramena River , Fok. Antsahabe, Fiv. Ambanja, 13°51ʹ50ʺS, 048°50ʹ53ʺE, Antilahimena 433 ( MO) ; Diana, Nosy Be, Lokobe Reserve Integral, 13°24ʹ37ʺS, 048°18ʹ33ʺE, Antilahimena & Devigny 250 ( CAS) ; Sahaenjika, Ampanavoana, Antalaha, Parc Masoala, 15°41ʹ33ʺS, 050°13ʹ30ʺE, Bernard et al. 154 ( CAS, MO) ; Nossi Be, Lokobe RNI, 5 km SE of Hellville , 13°25ʹS, 048°18ʹE, Birkinshaw 48 ( MO), 111 ( MO) 127 ( MO) ; Ambatosoratra, [ca. 14°32ʹS, 049°42ʹE], Cours 3327 ( P) ; Anjanaharibe, Cours 3719 ( P) ; Sous-préfecture d’Andapa, commune rurale de Doany, Fokontany de Betsomanga, versant NW du Marojejy, camp I, 14°25ʹS, 049°36ʹE, Gautier & Andriamparany 3887 (K, MO) ; R.N. du Marojejy, Guillaumet 4110 ( P) ; Province de Diego-Suarez des Roussettes au grand lac de la Montagne d’Ambre, Homolle s.n. ( P) ; Ankarana, forêt d’Analamalitso, S d’Arivorano-Nord, Humbert 19070 ( P) ; pentes E du Massif de Marojejy ( NE), W de la Rivière Manantenina, affluent de la Lokoho, Humbert 22550 ( P) ; contreforts W du Massif de Marojejy (nord-est), près du col de Doanyanala (limite des bassins de la Lokoho et de l’Andraronga), [ca. 14°19ʹS, 049°37ʹ20ʺE], Humbert 23153 ( P) ; partie W du Massif de Marojejy ( NE) de la callée de l’Ambatoharanana au bassin supérieur de l’Antsahaberoka, Humbert 31415 ( P) ; Massif de l’Anjanaharibe (pentes et sommet N), W d’Andapa (haute Andramonta , bassin de la Lokoho, NE), [ca. 14°36ʹS, 049°27ʹE], Humbert et al. 24655 ( BR, K, MO, P) route Andapa – Doany, vallée de l’Andranotsara, Jacquemin H565 J ( P) ; N of Andapa, Anjanaharibe-Sud Réserve Speciale, 14°46ʹ48ʺS, 049°28ʹ42ʺE, Malcomber et al. 2675 ( MO) ; WNW of Andapa, NW of village of Ambodisatrana , 14°32ʹS, 049°26ʹE, McPherson 17167 ( MO) ; Manongarivo Massif, above millage of Ambodisakoana, E of Ankaramy , 14°05ʹS, 48°20ʹE, McPherson & van der Werff 16410 ( MO) ; Réserve Naturelle de Marojejy, N slopes of Ambatosoratra , 14°32ʹS, 049°41ʹE, Miller 4206 ( MO) ; RNI Ankarana, 12°51ʹS, 049°05ʹE, Nicoll & Abraham 679 ( CAS, MO) ; ( Massif ), M’Tsaratanana, [ca. 13°57ʹS, 048°52ʹE], Perrier de la Bathie 2071 ( P) ; M’Zaratanana (Massif Tsaratanana), Perrier de la Bathie 15324 ( P) ; Ambohitalana, Antalaha, [ca. 14°54ʹS, 050°16ʹ30ʺE], PN Madagascar 3382 ( P) ; Sambava, PN Madagascar (Sazy) 8837 ( P) ; Réserve Speciale d’Ankarana, 12°55ʹS, 049°07ʹE, Rahajasoa et al. 202 ( MO) ; Réserve Speciale de Manongarivo, E d’Ankaramibe, Bekolosy , 14°03ʹ05ʺS, 048°17ʹ07ʺE, Rakotomalala & Narison 103 ( MO) ; Fiv. Andapa, Fir. Ambodiangezoka, Fok. Antanambe, Forêt Betaolana, 8.5 km NW d’Ambodiangezoka , le long de la rivière Ambolokopatrika, Campement 1, 14°32.3ʹS, 049°26.3ʹE, Rakotomalaza & Ravelonarivo 1907 ( MO) ; Andapa, Marojejy NP, near piscine naturelle, 14°26ʹ26ʺS, 49°46ʹ44ʺE, Rakotonasolo et al. 1870 ( CAS) ; Forêt de Besanatribe , Ambanja, Rakotozafy 331 ( P) ; Adnapa, Marojejy Natl. Park, 14°26ʹS, 49°43ʹE, Ranarivelo & Ravelonarivo 783 ( CAS) ; Andapa, Marojejy, ca. 15 km from Manantenina village , 14°26ʹ50ʺS, 49°45ʹ37ʺE, Randriamboavonjy et al. 899 ( CAS) ; Fiv. Ambanja, Forêt Classée d’Ambato, 40 km N d’Ambanja , près du village de Bevoay, piste vers Ankify, 13°27ʹS, 048°32ʹE, Randrianaivo et al. 257 ( MO) ; Sous-préfecture de Vohemar , commune rurale de Daraina, forêt d’Antsahabe, 13°12.88ʹS, 049°31.83ʹE, Ranirison & Nusbaumer 1116 ( MO) ; Réserve Naturelle integrale 12, Marojejy, N d’Andapa , aux environs du sommet de l’Est, 14°29ʹS, 049°38ʹE, Rasoavimbahoaka 15 ( MO) ; Réserve Speciale d’Anjanaharibe-Sud, Andapa , 14°36ʹS to 14°49ʹS, 049°23ʹE to 049°32ʹE, Rasoavimbahoaka & Ravelonarivo 203 ( MO) ; Préfecture d’Antalaha, Sous-préfecture d’Andapa, commune rurale de Bealampona, SW d’Andapa, Réserve Speciale d’Anjanaharibe-Sud , village d’Andranotsarabe , suivant la route Nationale d’Andapa–Bealanana de la piste à W, Ambatomainty , camp No. 2, 14°44ʹ42ʺW, 049°27ʹ42ʺE, Ravelonarivo & Rabesonina 477 ( CAS, MO) ; Fiv. d’Andapa, Fir. de Bealampona, Fok. de Befingotra , Réserve Speciale Anjanaribe-Sud, environs 37 km SW de l’ancien village de Mandritsarahely, piste vers Ranomafana, Ravelonarivo & Rabesonina 741 ( MO) ; Fiv. d’Andapa, Réserve Special d’Anjanaharibe Sud, aux environs du sommet, 14°46ʹ15ʺS, 049°29ʹE, Ravelonarivo et al. 17 ( MO) ; Préfecture d’Andapa, commune rurale de Bealampona, village de Mandritsarahely, Réserve Speciale d’Anjanaharibe-Sud, suivant la piste vers Ranomafana , 5.5 km SW de Befingotra, campement no. 1, 14°45ʹ03ʺS, 049°30ʹ03ʺE, Ravelonarivo et al. 420 ( MO) ; Tsaratanana RN 1, Fok. Beangona, 8 km de Beangona, Antsahamanara, 14°02ʹ50ʺS, 048°47ʹ09ʺE, Razakamalala et al. 36 ( CAS, MO). Fianarantsoa: col de Sakavalana, [ca. 20°44ʹ48.41ʺS, 046°04ʹ32.54ʺE], Alluaud s.n. ( P) ; ca. 5 km NW of the main entrance to Ranomafana NP on the road to Vohiparara , 21°14ʹ43ʺS, 047°23ʹ48ʺE, Almeda 8031 ( CAS) ; Iakora, Begogo, Bekora, forêt de Sahalava S du village de Androizaha, 23°32ʹ09ʺS, 046°32ʹ21ʺE, Andrianjafy et al. 740 ( MO) ; environs d’Ivohibe , Armand s.n. ( P) ; forêt basse au PK298 (sud Ambositra), Cremers 3639 ( P) ; Ranomafana Natl. Park, along road and Namorona River near cascade overlook ca. 5 km NW of main park entrance toward Vohiparara parcel, 21°15ʹS, 047°24ʹE, Daniel et al. 9131 ( CAS) ; Ranomafana Natl. Park, trail from Vatoharanana research camp to Valo research camp, 21°18ʹS, 047°25ʹE, Daniel et al. 9157 ( CAS) ; Ranomafana Natl. Park, Talatakely trail system S of Namorona River , 21°16ʹS, 047°26ʹE, Daniel et al. 9239 ( CAS) ; Ranomafana Natl. Park, Vohiparara trail system N of Namorona River (ca. 1.5 km N of cascade overlook along Hwy. 25), Daniel 9271 ( CAS) ; Ranomafana Natl. Park, Vohiparara trail system N of Namorona River (ca. 1.5 km N of cascade overlook along Hwy. 25), 21°14ʹS, 047°23ʹE, Daniel et al. 9276 ( CAS) ; Vondrozo, P. de Farafangana , [ca. 22°49ʹS, 047°19ʹ30ʺE], Decary 5311 ( P) ; Massif di l’Kongo – P. de Farafangana , Decary 5691 ( MO, P) ; Fort Carnot, Prov. de Farafangana , [ca. 21°51ʹ30ʺS, 047°26ʹ30ʺE], Decary 5804 ( P) ; Ranomafana Natl. Park, trail to Vatoharanana from Talatakely, 21°17ʹS, 047°25ʹE, de Nevers 11373 ( CAS) ; Ranomafana Natl. Park, trail between Vatoharanana and Takatakely parcels, 21°15ʹS 047°27ʹE, Fritsch 1554 ( CAS) ; Ranomafana Natl. Park, along trail between Vatoharanana and Takatakely parcels, just N of A 1250 m trail marker, 21°15ʹS, 047°27ʹE, Fritsch 1556 ( CAS) ; Bassin de la Matitanana, Cascade d’Ankitso, Forêt de Tsianovoha , Heim s.n. ( P) ; Réserve Speciale de Pic d’Ivohibe , forest of Marovitsika, [22°31ʹ30ʺS, 046°59ʹE], Hoffmann et al. 218 (K) ; Chaine du Vohibory (W d’Ivohibe ), Humbert 3126 ( P) ; Pic d’Ivohibe (Bara), Humbert 3309 (A, P, K, MO) ; haute valeé de la Rienana (bassin du Matitanana), Humbert 3457 (K, P), 3564 (A, P) ; Akimba, near Ankerana, 30 km from Ambositra, E slope of watershed mountains, 20°37ʹS, 047°19ʹ30ʺE, Jongkind & Rapanarivo 908 ( MO) ; 10 km S d’Ambalavao, Keraudren-Aymonin & Aymonin 24621 ( P) ; PN de Ranomafana, parcelle #3, Talatakely, piste AA, 21°15ʹS, 047°27ʹE, Kotozafy 125 ( BR, MO) ; PN de Ranomafana, parcelle #3, Vatoharanana , 21°15ʹS, 047°27ʹE, Kotozafy & Randriamanantena 302 ( MO) ; Parc Natl. de Ranomafana , aux environs de Vohiparara, 21°14ʹS, 047°23ʹE, Kotozafy et al. 372 ( BR, MO) ; Andringitra, Camp I, ca. 45 km S of Ambalavao, E bank of Iantara River, along Ambalamanenjana–Ambatomboay trail, edge of Andringitra Reserve , 22°13ʹ20ʺS, 047°01ʹ29ʺE, Lewis et al. 759 ( BR, MO) ; Andringitra, Camp III, ca. 40 km. S of Ambalavao, Andringitra Reserve, along tributary of Sahavatoy River , 22°13ʹ22ʺS, 046°58ʹ18ʺE, Lewis et al. 944 ( MO) ; between Fianarantsoa and Ifandiana, Route National 45, Parc Natl. Ranomafana, trail S from Cabine de Recherche to Vato camp, 21°15ʹS, 047°27ʹE, Malcomber 1047 ( MO) ; S of Farafangana, in Forêt Classée near Manombo Reserve , 23°03ʹ16ʺS, 047°40ʹ28ʺE, McPherson & Rabenantoandro 18434 ( CAS, MO) ; bord de l’Ihovika, bassin de Matitanana, Perrier de la Bathie 10267 ( P) ; Fiv. & Fir. Ivohibe, Réserve Speciale d’Ivohibe , 8 km E d’Ivohibe, 22°29ʹS, 046°58ʹ06ʺE, Rakotomalaza et al. 1447 ( MO) ; Mahazony, Ambalavao , Ankotsobory, [ca. 22°13ʹ30ʺS, 046°55ʹ30ʺE], Rakotonao 9178 ( P) ; Sendrisoa, Ambalavao , [ca. 22°02ʹ27ʺS, 047°00ʹ34ʺE], Rakotonao 9988 ( P) ; Ivohibe, dans la RNI d’Andringitra, NO d’Ambarongy, SO du campement #3, 22°13ʹ22ʺS, 046°58ʹ18ʺE, Rakotovao 270 ( MO) ; Ivongo, Farafangana , Ankarenambe, Rakotovao 613 ( P) ; proche de Behazy-Sud Antomilahala, Farafangana , Rakotovao 7485-RN ( P) ; Fiv. Ivohibe, Fir. Ivohibe, limite N de la Réserve Speciale d’Ivohibe le long de la rivière Ifefitany, 7.5 km ENE d’Ivohibe, Campement 1, 22°28ʹ12ʺS, 046°57ʹ36ʺE, Rakotovao et al. 847 ( MO) ; Ivohibe, Ivongo, Ambahatsy, RNI Andringitra, au 2e campement, forêt coupee par la riviere d’Iatara, 22°13ʹ40ʺS, 047°00ʹ13ʺE, Razafindrabe et al. 98 ( MO) ; Localité: R N V Canton: Antambohobe District : Ivohibe, Razafindrakoko 3518-RN ( P) ; 7 km W of Ranomafana, on slopes above Namorona River , Duke Univ. Primate Center study site, 21°16ʹS, 047°25ʹE, Schatz 2597 ( MO) ; 7 km W of Ranomafana, just S of Namorona river, Duke Primate center study site, 21°16ʹS, 047°25ʹE, Schatz & Miller 2444 ( CAS, MO). Mahajanga: road from Antsohihy to Bealanana, 21–23 km E of Antsahabe , [ca. 14°44ʹ7.95ʺS, 048°32ʹ46.63ʺE], Gentry 11738 ( MO) ; Réserve Speciale d’Anjanaharibe-Sud, 11 km WSW Befingotra , 14°44ʹ30ʺS, 049°26ʹ30ʺE, Lewis et al. 1309 ( MO) ; Sous-préfecture de Befandriana-Nord, commune rurale de Matsoandakana, Quartier de Belalona, SW d’Andapa, Réserve Speciale Anjanaharibe-Sud, village d’Anjiamazava, suivant la route Nationale d’Andapa – Bealanana, piste vers le N approchant le sommet de Bevitsika, 14°42ʹS, 049°27ʹE, Ravelonarivo & Rabesonina 558 ( CAS, MO). Toamasina: Moramanga , Savoka, Ambodilongotra, village Coby André Daubercies, Abraham 26779 ( P) ; Phelps Dodge project site, ca. 15 air-km NE of Moramanga, ca. 11 km E of Antanambao, Ambatovy-South , 18°52ʹ06ʺS, 048°18ʹ23ʺE, Andriatsiferana et al. 2162 ( MO); PN de Masoala, sur la route d’Ambanizana à Analambolo (25 km N d’Ambanizana), ca. 6 km NE d’Ambanizana, Fiv. Maroantsetra , 15°34ʹ07ʺS, 050°00ʹ14ʺE, Aridy et al. 44 ( MO); 3 km ESE of Ankosy, on path from Ankosy to Antenina, 500 m from edge of RNI Zahamena, to 1000 m inside RNI, Birkinshaw et al. 623 ( MO); Ambohidavak’ely (D-43), District d’Ambatondrazaka, Cours 515 ( P); Moramanga , route d’Anjiro, Cours 807 ( P); District d’Ambatondrazaka, Cours 953 ( MO, P) ; Ambodimanga à Antanambao , Cours 2787 ( P), 2792 ( P) ; de Didy à Brickaville, Cours 4832 ( P), Cours 4844 ( P), Cours 4874 ( P) ; S de Moramanga, Decary 6988 ( P), 6994 ( P) ; Fanovana, [18°55ʹ30ʺS, 048°32ʹE], Decary 17981 (K, MO, P) ; moyennevallée du Mangaro, Decary 18475 ( P) ; Lac Alaotra-D43, [ca. 17°29ʹ16.28ʺS, 048°34ʹ2.81ʺE], Jardin Botanique de Tana 3901 ( P) ; Soanierana – Antasibe, Lam & Meeuse 825 ( P) ; bords de torrents entre Sandrangato et Anosibe (S de Moramanga), Leandri 1679 (K, MO, P) ; East Coast, Betampona Réserve Naturelle Integrale, 40 km NW of Toamasina , [ca. 17°54ʹS, 049°13ʹE], Lewis & Razafimandimbison 688 ( MO) ; Maroantsetra, trail E of village of Hiaraka, ESE of Maroantsetra, across bay, on NW coast of Masaola Peninsula , [15°29ʹS, 049°54ʹE], Lowry et al. 4045A ( MO) ; Maroantsetra, Masoala Peninsula, Antalavia, ca. 22 km S of Ambanizana, along trail NE from “village” to Ampatra river , 15°46ʹS, 050°01ʹE, Lowry 4330A ( CAS, MO) ; W of Vavatenina, Réserve Naturelle Integrale Zahamena, forest of Amboditamenaka, 17°44ʹS, 049°00ʹE, Malcomber et al. 2518 ( MO), 2575 ( MO) ; NE of Moramanga, at nickel mining exploration site, Ambatovy , 18°51ʹ25ʺS, 048°17ʹ50ʺE, McPherson 17475 ( MO) ; near Andasibe , forest of Mantadia, beyond graphite mine, 18°55ʹS, 048°25ʹE, McPherson & van der Werff 16472 ( MO) ; near Andasibe , forest of Mantadia, beyond graphite mine, 18°55ʹS, 048°25ʹE, McPherson & van der Werff 16539 ( MO) ; Forêt d’Analamazoatra, Perrier de la Bathie 10239 ( P) ; Analamazoatra, Perrier de la Bathie 10242 ( P) ; Riv. Anove – Coli Est, Perrier de la Bathie 10256 ( P) ; Phelps Dodge project site, ca. 15 air-km NE of Moramanga, ca. 11 km E of Antanambao, Ambatovy, Antsahalava River east, 18°50ʹ54ʺS, 048°17ʹ56ʺE, Rakotomalaza et al. 1102 ( MO) ; fiv. Vivatenina, commune: Ambodimangavalo, 10 km W d’Antevibe-Sahandrazana, adjacente du Parc Natl. de Zahamena , 17°32ʹS, 48°48ʹE, Rakotonandrasana et al. 714 ( MO) ; Moramanga, commune: Ambohibary, fok.: Ampitambe, Ambatovy, NE de Moramanga, environ 22 km de Moramanga , 18°50ʹ57ʺS, 048°17ʹ18ʺE, Rakotovao et al. 1321 ( MO) ; district: Moramanga, commune: Ambohibary, fokontany: Ampitambe, Ambotovy, 22 km NE de Moramanga , 18°51ʹ21ʺS, 048°18ʹ22ʺE, Rakotovao et al. 1407 ( MO) ; Alaotra Mangoro, Moramanga, Andasibe, 18 km N from Andasibe , 18°48ʹ49ʺS, 48°25ʹ48ʺE, Ranarivelo & Ranaivojaona 322 ( CAS) ; Maroanbehy, Tamatave, Randriamahavita 9064-RN ( P) ; Parc Natl. de Zahamena , Antanandava, 2 km SW du village d’Ankosy, au sommet de la Forêt d’Ampangalambolosy, dans le parc, 17°29ʹ38ʺS, 048°43ʹ50ʺE, Randrianasolo et al. 130 ( MO) ; district: Fenerive-Est. Tampolo Forestry Station , along the “Grand Layon,” ca. 1 km. from office, 17°17ʹS, 049°23ʹE, Randrianasolo et al. 605 ( BR, CAS, MO) ; sous-préfecture: Vavatenina, commune: Ambodimangavalo, fokontany: Manakambahiny, Parc Natl. de Zahamena , 17°29ʹ30ʺS, 048°54ʹ32ʺE, Randrianjanaka et al. 652 ( MO) ; Moramanga, Andasibe , Menalamba, entre Analamay et Ambatovy, 18°49ʹ47ʺS, 048°18ʹ41ʺE, Razafindrabe et al. 127 ( MO) ; fivondronana: Vivatenina, commune: Ambodimangavalo, fokontany: Manakambahiny I. Andranofantsona 10 km S de Manakambahiny I, au bord du rivière Ihofika , 17°39ʹ17ʺS, 048°49ʹ10ʺE, Razafitsalama et al.136 ( MO) ; Vatomandry, Ambalabe, Ambinanindrano II, NE du Tobin’I Foara , 19°09ʹ26ʺS, 048°34ʹ47ʺE, Razanatsima & Ranaivojaona 36 ( MO) ; Vatomandry, Ambalabe, Ambinaninandro II, E de Toby Foara , 19°09ʹ46ʺS, 048°34ʹ59ʺE, Razanatsima et al. 66 ( MO) ; Alaotra Mangoro, Moramanga , Beforona, Forêt Ankeniheny, 19°07ʹ50ʺS, 048°32ʹ10ʺE, Razanatsima et al. 297 ( MO) ; Alaotra Mangoro, Moramanga , Lakato, village Manasamena, forêt corridor Ankeniheny, Razanatsima et al. 422 ( MO) ; Alaotra Mangoro, Moramanga , Lakato, village Manasamena, forêt corridor Ankeniheny, 19°04ʹ34ʺS, 048°30ʹ40ʺE, Razanatsima et al. 475 ( MO) ; Nosy Mangabe, in the Bay of Antongil, 5 km S of Maroansetra , 15°30ʹS, 049°46ʹE, Schatz & Carlson 2920 ( MO) ; Nosy Mangabe, 5 km from Maroansetra in Bay of Antongil, 15°30ʹS, 049°46ʹE, Schatz & Gentry 2177 ( CAS, MO) ; Masoala Peninsula, just S of Ambanizana village , 15°36ʹS, 049°57ʹE, Schatz et al. 1345 ( MO) ; Nosy Mangabe, 5 km from Maroansetra in the Bay of Antongil , 15°30ʹS, 49°46ʹE, Schatz et al. 2380 ( MO) ; Perinet , Moramanga, Service Forestier de Madagascar 1214 ( P) ; Analamazaotra, Perinet, Service Forestier de Madagascar 1414 ( P) ; Analamazaotra, TAN 3746 (K, MO, P) ; Ranomafana , 19°06ʹS, 048°42ʹE, van der Werff 12665 ( MO) ; near Andasibe , forest of Mantadia, beyond graphite mine, 18°55ʹS, 048°25ʹE, van der Werff et al. 13773 ( MO) ; Masoala Peninsula, S of village of Ambanizana in Andranobe River watershed, 15°40ʹ24ʺS, 049°57ʹ51ʺE, Vasey & Behasy 388 ( MO) ; Province d’Andovoranto, District de Moramanga , Forêt d’Analamazaotra, près du col d’Amboasary, Viguier & Humbert 984 ( P). Toliara: Fort Dauphin , Elliot 2532 ( P) ; Massif de Beampingaratra ( SE), du col de Bevava au sommet de Bekoho, [ca. 24°26ʹS, 046°53ʹ30ʺE], Humbert 6417 ( P) ; bassin supérieur du Mandrare (sud-est), col et sommet de Marosoui, [24°32ʹS, 046°48ʹE], Humbert 6599 ( P) ; Massif du Kalambatitra (centre-sud), Mont Beanjavidy, [23°22ʹS, 046°27ʹE], Humbert 12043 ( P) ; Massif de l’Ivakoany (centre-sud), Humbert 12217 ( P) ; Fort Dauphin, Eminiminy , RN #11 Andohahela, Camp 3, 13.5 km NW d’Eminiminy, 24°35ʹS, 046°44ʹE, Rakotomalaza 504 ( MO). Province undetermined: Ankeramadinika, Academie Malgache s.n. ( P) ; without locale, Baron 1345 ( P), 1743 ( P), 1789 ( P), 2828 (K, P), 2877 ( P), 3214 ( P), 3580 (K, P), 5149 ( P), Chapelier s.n. ( P), Commerson s.n. ( P), without collector 345 ( P) ; central Madagascar , Baron 1463 ( P), 2828 (K, P), 2920 (K, P), 3667 (K), 4196 (K, P) ; chiefly from NW Madagascar , Baron 3749 (K) ; forêt N de la route de Nickelville sur les bords Ankotréano, Cours 2079 ( MO, P) ; S de Mangabe, Cours 2505 ( P) ; près Samalahaza, Dequaire 27869 ( P), 27963 ( P) ; central Madagascar , Andrangaloaka, Parker s.n. (K) ; route Antsirabe à Fianarantsoa , Forêt d’Ambasofiterahana, Keraudren 245 ( P) ; Forêt de Fotsialanana , Rakotozafy 672 ( P) ; Sahamalaza, Forêt de l’Est aux confines du pays Sihanaka, TAN 2827 ( P). MAYOTTE ( French island in Comoros Archipelago). Grande Terre: Combani, Bajoni, [ca. 12°47ʹ39.06ʺS, 045°09ʹ01.24ʺE], Barthelat & Mchangama 1738 ( CAS, K) ; Dzomougne, Réserve Forestière d’Hachiroungou, [ca. 12°43ʹ01.85ʺS, 045°06ʹ16.05ʺE], Barthelat et al. 568 ( CAS, MO, P) ; Accoua, Réserve Forestière d’Hachiroungou, [ca. 12°43ʹ03.20ʺS, 045°03ʹ54.22ʺE], Barthelat et al. 597 ( MO).

Figures

FIGURE 12. Map of Madagascar showing distributions of Mendoncia decaryi and M. delphina, and map of Madagascar and Mayotte showing distribution of M. flagellaris.

FIGURE 1. Phylogenetic relationships among genera of Acanthaceae sub- family Thunbergioideae and Avicennia. OW = Old World, NW = New World. Adapted from Borg et al. (2008).

FIGURE 11. Mendoncia cowanii (A–D) and M. flagellaris (E, F). A. Mature drupe. B. Corolla limb showing discoloration due to insect visitation. C, D. Flowers with corolla tube encased by fluid-filled bracteoles. E. Immature drupes. F. Flower and bracteoles, each with a pair of prominent, white, and basal protuberances. (A–D, photos by T. Daniel of Daniel et al. 11000; E, photo [cropped] of Razanatsima et al. 422, copyright by Aina Razanatsima on Tropicos [www.tropicos.org/Image/100126998], CC by-NC-ND 3.0; F, photo by T. Daniel of Daniel et al. 9239).

FIGURE 13. Mendoncia flagellaris (A–G, J, Daniel et al. 9239; H, I, Razanatsima et al. 422). A. Habit. B. Fertile node with multiple dichasia. C. Dichasium showing a flower and paired protuberances on bracteoles. D. Unopened bracteoles from above showing paired protuberances. E. Corolla split open to show androecium. F. Stamen showing two thecae bearing basal bristles, poricidal dehiscence, and extension of connective distal to pores. G. Apex of style and stigma. H. Fresh drupe (from photo) showing sequentially from top: distal portion of peduncle, calyx, and drupe. I. Dried drupe showing sequentially from top: distal portion of peduncle with flared apex, calyx with undulations, nectar disc, and drupe. J. Seed. Drawn by Jenny Speckels (A–G) and Becca Berezuk H–J.