Three New Species of Mendoncia (Acanthaceae) from Madagascar

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1 Three New Species of Mendoncia (Acanthaceae) from Madagascar Emily B. Magnaghi and Thomas F. Daniel* Department of Botany, California Academy of Sciences, 55 Music Concourse Drive, Golden Gate Park, San Francisco, California 94118, U.S.A. *Author for correspondence: ABSTRACT. Three species of Mendoncia Vell. ex Mendoncia in Madagascar and Africa (Magnaghi, Vand. (Acanthaceae) are newly recognized from 2010), which included many more collections than Madagascar: M. decaryi (Benoist) E. Magnaghi, M. were known to Benoist, have revealed the necessity delphina E. Magnaghi, and M. kely E. Magnaghi. for changes in the status of M. cowanii var. decaryi Each is based on a unique suite of morphological (which is herein elevated to species status) and the characters, and all are endemic to Madagascar. A key existence of two new species, which we describe to the six species of the genus known from below. Madagascar is provided. A lectotype is designated The three species newly treated here occur in for M. cowanii (S. Moore) Benoist var. decaryi northern, eastern, and southern Madagascar and are Benoist. endemic to the island. Each exhibits a unique Key words: Acanthaceae, IUCN Red List, Mada- combination of features that distinguishes it from gascar, Mendoncia. other Paleotropical members of the genus. Because each of these species is currently known from Mendoncia Vell. ex Vand. consists of about 80 relatively few localities in the northern and eastern species occurring worldwide, but species are con- wet-evergreen forests of Madagascar that have centrated in the Neotropics. Some 70 species occur in suffered extensive deforestation, we summarize moist to wet forests of the New World, from information useful toward assessing the conservation southeastern Mexico to Bolivia and Brazil, with the status of each, based on IUCN Red List categories, greatest number of species (35 to 40) recorded from criteria, and guidelines (IUCN, 2013). Google Earth Colombia. Species in the Old World occur throughout Pro (Google Inc., 2013) was used to plot the tropical Africa, from Liberia east to Kenya, and on collections known for each species and to estimate the islands of Madagascar and Mayotte. To date, there their extent of occurrence (EOO) and area of has been no recent comprehensive taxonomic account occupancy (AOO; grid size 4 km 2 ). Inadequacies in for the entire genus. our knowledge of these species, and thus possible Mendoncia is included in the Acanthaceae sources of error in our assessments, include our subfamily Thunbergioideae, where it is sister to understanding (1) that although the major herbaria Thunbergia Retz. and Pseudocalyx Radlk. (Scotland with holdings of Malagasy plant collections have been & Vollesen, 2000; Borg et al., 2008; McDade et al., consulted, the distributional data obtained from those 2008). The genus is distinguished from other sources are not necessarily complete; (2) that our Acanthaceae by the combination of its twining habit, knowledge of habitats is based primarily on informaenlarged and paired bracteoles subtending the tion provided on herbarium specimen labels, which is flowers, highly reduced calyx, 5-brevicolpate and often scanty and can result in imprecise comparisons spherical pollen, and 1-seeded, drupaceous fruits. of habitats; and (3) that no counts or estimates of Three species of Mendoncia have been reported numbers of mature individuals at a site are available. from Madagascar: M. cowanii (S. Moore) Benoist, M. Our attempts to estimate numbers of individuals for flagellaris (Baker) Benoist, and M. vinciflora Benoist. other species of Malagasy Mendoncia (e.g., M. Mendoncia cowanii is endemic to the wet tropical cowanii and M. flagellaris) were complicated by the forests of eastern Madagascar, and M. vinciflora is twining habit of these plants and by their inconspicrestricted to the wet montane forests in the north of uousness, except in the forest canopy. Considerable the island. Mendoncia flagellaris occurs in wet loss of habitat through time due to human activities in tropical forests throughout Madagascar and also on the lowland and montane, wet-evergreen forests of the island of Mayotte, in the Comoro Archipelago. northern and eastern Madagascar has been well Benoist (1944, 1967) recognized three varieties of M. documented (e.g., Preston-Mafham, 1991). If these cowanii: variety cowanii, variety coursii Benoist, and forests are treated as a single location (IUCN, 2013: variety decaryi Benoist. Recent taxonomic studies of 41 guidelines define location as a geographically or doi: / NOVON 23: PUBLISHED ON 16 JULY 2014.

2 188 Novon ecologically distinct area in which a single threaten- Indeed, for each of the three species of Mendoncia, ing event can rapidly affect all individuals of the some collection sites occur in areas receiving habitat taxon present ), then species will receive a higher protection while others do not. Those sites receiving threat category than they would if the forests were habitat protection could not be considered part of the treated as multiple locations. Although the IUCN same location as the sites without protection under (2013) guidelines do not define their use of the word the IUCN (2013) guidelines. Based on the distriburapidly, from an example provided therein the threat tional data (including whether some subset[s] of the from gradual loss and degradation of habitat in these plants occur in protected lands), we propose a forests would not qualify them as a single location. provisional conservation assessment for each taxon. KEY TO SPECIES OF MENDONCIA IN MADAGASCAR 1a. Stigma subcapitate (unlobed or lobes not evident); style 3 10 mm; drupes glabrous, flowers white (usually with purple markings in throat) M. flagellaris 1b. Stigma subequally 2-lobed, lobes mm; style mm; drupes glabrous or pubescent; flowers white (usually with purple markings in throat) to pink to purple. 2a. Bracteoles glabrous to sparsely pubescent; calyx glabrous; drupes glabrous. 3a. Leaves glabrous; drupes oblong to ellipsoid; flowers white with purple in throat M. delphina 3b. Leaves evenly and sparsely to densely pubescent; drupes spherical; flowers violet-purple.... M. vinciflora 2b. Bracteoles densely pubescent, calyx glabrous or pubescent, drupes pubescent. 4a. Leaves mostly broadly elliptic (length/width ); corollas mm, tube mm; style 9 10 mm; drupes ovoid to spherical M. kely 4b. Leaves mostly elliptic (length/width ); corollas mm, tube mm; style mm; drupes ovoid to subellipsoid. 5a. Corollas white with lavender at mouth, limb mm diam., upper lip mm with lobes mm, lower lip mm with lobes mm M. decaryi 5b. Corollas pink to dark pink, limb 6 20 mm diam., upper lip mm with lobes mm, lower lip 3 12 mm with lobes mm M. cowanii 1. Mendoncia decaryi (Benoist) E. Magnaghi, comb. apex, acumen/apicule mm, abaxial surfaces et stat. nov. Basionym: Mendoncia cowanii (S. densely pubescent with eglandular trichomes Moore) Benoist var. decaryi Benoist, Notul. Syst. mm, adaxial surfaces mealy-glandular, margins of the (Paris) 11: TYPE: Madagascar. pair connate from base to two thirds of the way toward Toamasina: Zahamena (réserve no. 3), 23 Mar. the apex during anthesis. Calyx consisting of a rigid, 1941 (fl., fr.), R. Decary (lectotype, entire, undulating annular or cupular ring surrounddesignated here, P ). Figure 1. ing nectar disc, mm high, densely pubescent with eglandular trichomes 2 3 mm; corolla white with Woody vines; young stems quadrate to quadratelavender at the mouth, mm, tube sulcate, evenly and densely to sparsely pubescent mm, apically ampliate, limb mm diam., with golden-brown antrorse eglandular multicellular upper lip 13.5 mm with rounded or emarginate lobes trichomes mm, sometimes pubescent with mm, lower lip mm with unicellular short trichomes mm underlying rounded or emarginate lobes longer flexuous trichomes mm. Leaves petiomm; stamens mm, inserted near the middle of late, petioles to 15 mm, blades mostly elliptic (length/ corolla tube, ventral pair inserted 3.8 mm distal to the width ), but sometimes varying to broadly dorsal pair, thecae mm, densely pubescent at elliptic or subobovate, mm, rounded base with a tuft of stout eglandular trichomes 0.16 to cordate at base, acute to acuminate at apex, abaxial 0.36 mm, sometimes ventral pair with dense surfaces bearing entangled multicellular flexuous pubescence along length of thecae, connective trichomes mm, sometimes with an understory projecting mm beyond apex of thecae, of shorter trichomes as well, sometimes with tufts of attenuate; pollen not seen; style mm, woolly trichomes forming domatia in axils of major trichomes on lower half of style mm, stigma veins, adaxial surfaces pubescent with antrorse subequally 2-lobed, lobes mm. Drupe dark eglandular multicellular trichomes mm (hirsute purple to black, subellipsoid, mm, to villous), trichomes often more dense on major pubescent with golden brown eglandular trichomes veins, margin ciliate. Inflorescence of 1 to 3 flowers mm. in leaf axils, flowers alternate or opposite at nodes, pedunculate, peduncles to 20 mm, pubescent-like Distribution and habitat. Mendoncia decaryi stems; bracteoles ovate to elliptic to broadly elliptic, (Benoist) E. Magnaghi is endemic to northern and mm, long acuminate to subapiculate at eastern-central Madagascar (former provinces Ant-

3 Volume 23, Number 2 Magnaghi & Daniel Mendoncia (Acanthaceae) from Madagascar Figure 1. Mendoncia decaryi (Benoist) E. Magnaghi. A. Habit. B. Close-up of trichomes on adaxial leaf surface. C. Drupe. D. Androecium on dissected corolla tube. E. Calyx and gynoecium. F. Stigma. A, B, and F from J. Miller 4202 (MO), with flowers from D. Du Puy & Du Puy MB 163 (K); C from J. Razafitsalama 473 (MO); D from Du Puy & Du Puy MB 163; E from the lectotype Decary siranana and Toamasina) where plants occur in by ca. 275 km, a northern subpopulation with an EOO primary lowland to montane, wet-evergreen forest at of 1355 km 2 and a southern subpopulation with an elevations between 300 and 1400 m. EOO of 489 km 2. The AOO consists of 52 km 2.Based on the EOO, this species would be considered IUCN Red List category and conservation data. At Vulnerable (VU) under IUCN criterion B1, and based least 17 collections from a minimum of 14 sites are on the AOO it would be considered Endangered (EN) known for Mendoncia decaryi. The population (total under IUCN criterion B2, in each case if appropriate distributional range) has an EOO of 19,447 km 2.Itcan knowledge of subcriterion b or c were available. Two of be divided into two disjunct subpopulations separated the sites for this species in the northern subpopulation

4 190 Novon occur in the Marojejy Nature Reserve, and at least specimen designated above as lectotype agrees best three in the southern subpopulation occur in the with the protologue and has the most intact flowering Zahamena National Park. Because of deforestation in and vegetative material. this region of Madagascar in combination with the reduced threat to M. decaryi afforded by its partial Representative specimens examined. MADAGASCAR. Antsiranana: N of Maroantsetra, ca. 8 km E of Sahavary, occurrence in two protected areas, we propose a Andranofotsy River, Du Puy & Du Puy MB 163 (K); provisional assessment of Vulnerable (VU) for the Andapa, Doany, Betsomanga, NW du Marojejy, 0.2 km N species. du camp I, Gautier et al (K, G, MO); Rés. Nat. Marojejy, N slopes of Ambatosoratra, Miller 4202 (MO, Phenology. Plants of Mendoncia decaryi have TAN); Andapa, Ambodiangezoka, Antanambe, foret ˆ de Betaolana, à 8.5 km NW d Ambodiangezoka, Rivière been collected in flower from February through April; Ambolokopatrika, Camp. I, Rakotomalaza & Ravelonarivo fruiting collections have been made from March PJ1956 (G); Andranotsarabe, Befingotra, Bealampona, through September. Andapa, Rasoavimbahoaka & Rastefanonirina 287 (BR, MO, TAN); SW d Andapa, Rés. Spéc. Anjanaharibe Sud. Etymology. The epithet is taken from the name of Ambodisatrana, Ravelonarivo 145 (MO). Toamasina: rés. Benoist s variety, Mendoncia cowanii var. decaryi, in nat. 3 (Zahamena), Decary (P , K , photo); Betampona, rés. nat. 1, Decary honor of Raymond Decary, who collected the type. (P, TAN , photo); Majunga, 9.6 km NW of Discussion. Benoist (1944, 1967) distinguished Ambohitsaratelo-Bebao (NW of Tsiroanomandidy), Dorr 3538 (MO, TAN); Ampitanonoka à Fotsialanana, près du Mendoncia cowanii var. decaryi from the nominate Lac Alaotra, Herb. Station Agricole du L. Alaotra 2441 variety by its slightly larger leaves ( cm (MO); W of Vavatenina, Rés. Nat. Intégrale Zahamena, forest of Amboditamenaka, Malcomber et al (MO, vs cm) and longer and denser trichomes TAN); Bongolava W de Tsiroanomandidy, Morat 4776 (P); (specific measurements not provided). Based on Vavatenina, Miarinarivo, Fokontany Anamborano, limite several other morphological characters, M. cowanii entre Vavatenina et Toamasina II, à 500 m d Ifasina, var. decaryi is here elevated to specific rank as M. Rakotondrafara et al. 238 (MO); RNI Tamatave, Rakotoniana 2875 (P); Parc Nat. Zahamena, Andranofantsona, decaryi. The primary characters that distinguish it Manakambahiny I, Ambodimangavalo, Ratovoson et al. 273 from M. cowanii are summarized in the key above. (MO); Vavatenina, Ambodimangavalo, sect. 2, environs More subtle characters include cauline pubescence, Parc Azhamena, foret ˆ Ambinanin Antsahabesahona, vers 16 venation relief, and texture of the leaves. Trichomes km SE d Ambarifotsy, Rivière Ihofika, Razafitsalama 473 on the young stems tend to be more conspicuous and (MO). longer (up to 3 mm vs. up to 2 mm) in M. decaryi than in M. cowanii var. cowanii, and trichomes on the 2. Mendoncia delphina E. Magnaghi, sp. nov. TYPE: abaxial surfaces of bracteoles and leaf blades are Madagascar. Toliara: NW of Taolagnaro (Ft. Dauphin), réserve intégral no. 11 (Andohahela), erect to flexuose and do not obscure the surface (vs. parcelle I, E boundary, S, E, 250 densely wooly and obscuring the surface of bracteoles 500 m, Oct (fl.), S. Malcomber, H. in M. cowanii var. coursii). In M. decaryi the firstvan der Werff, B. Gray, S. Raparanivo & B. through fourth-order foliar veins are all prominently Randriamanpionona 1665 (holotype, MO raised and visible; in M. cowanii, the first- and ; isotype, K, not seen, TAN, not seen). second-order veins are prominently raised and Figures 2, 3B D, F. visible, but the subsequent orders are much less so. The leaf texture of M. decaryi appears to be more brittle, and the leaves are thinner than in M. cowanii, Diagnosis. Mendoncia delphina E. Magnaghi differs from M. vinciflora Benoist by its glabrous (vs. pubescent) which tends to have thicker and more coriaceous leaves, oblong to ellipsoid (vs. spherical) drupes, and white leaves. The geographic distribution of M. decaryi is corollas with purple markings (vs. violet-purple). restricted to the east-central part of the island in and near the Zahamena National Park, near Lac Aloatra, Woody vines; young stems quadrate to quadrateand in the north and near the Marojejy National Park. sulcate, internodes glabrous, nodes often pubescent It partially overlaps that of M. cowanii, which is with sparse trichomes mm. Leaves petiolate, widespread in wet-evergreen forests in northern and petioles to 14 mm, blades elliptic (length/width 2.1 eastern Madagascar. 3.2), mm, caudate to acuminate at In the protologue of Mendoncia cowanii var. apex, acute to cuneate at base, surfaces glabrous. decaryi, Benoist (1944) cited three collections of Inflorescence of 1 to 2 flowers in leaf axils, flowers Decary (16712, 16741, 16924) but did not designate alternate or opposite at nodes, pedunculate, peduna specimen of any of them as the type. From among cles to 40 mm, glabrous to pubescent with sparse these collections at P (where Benoist worked), the golden brown, appressed trichomes mm;

5 Volume 23, Number 2 Magnaghi & Daniel Mendoncia (Acanthaceae) from Madagascar bracteoles ovate to elliptic, mm, of the type specimen, near Taolagnaro (Fort Dauacute at apex, rounded at base, abaxial surface phin). glabrous or with a few eglandular trichomes to 0.1 mm, adaxial surface mealy glandular, woolly at apex, Discussion. Mendoncia delphina is distinguished margins connate from base to 2/3 of the way toward from other Malagasy Mendoncia by a combination of apex; calyx consisting of a rigid, entire, undulating floral features, geographic distribution, and phenolannular or cupular ring surrounding nectar disc, to ogy. The corolla resembles that of M. flagellaris in 1.4 mm high, glabrous; corolla purple to white with color (white with purple markings in the throat), but purple throat, mm, tube mm, the stigma is subequally 2-lobed as in M. cowanii. apically ampliate, limb mm in diam., upper lip Flowering occurs in October and November, which mm with rounded lobes contrasts with M. cowanii (February to April) but is mm, lower lip mm with rounded lobes similar to that of M. flagellaris (October to February) mm; stamens mm, inserted near The distribution of M. delphina overlaps that of both the middle of corolla tube, ventral pair inserted 0.5 M. cowanii and M. flagellaris but appears to be mm distal to dorsal pair, thecae mm, densely restricted to southern Madagascar. pubescent at base with a tuft of stout eglandular Pollen of relatively few species of Mendoncia have trichomes mm, connective projecting been studied to date. Pollen of both M. delphina and mm beyond apex of thecae; pollen 5-brevicolpate; M. kely E. Magnaghi were available for study, and we style mm, stigma subequally 2-lobed, lobes examined unacetolyzed grains from samples of each mm. Drupe (possibly immature) ovatespecies (e.g., Raj, 1961; Muller et al., 1989; Daniel, using SEM (Fig. 3). Like that reported for other elliptic, 13.7 mm, 8.5 mm diam., glabrous. 1998), pollen of these species is spherical, 5- Distribution and habitat. Mendoncia delphina is aperturate, brevicolpate, and rugulate over the endemic to extreme southeastern Madagascar (former interapertural surface. Endoapertures, which are province of Toliara) where plants occur in lowland, usually not visible in SEM preparations of Mendoncia wet-evergreen forests near Taolagnaro (Fort Dauphin) and were not evident in our preparations of these at elevations between 250 and 500 m. species, possibly accompany each colpus. Although 5-aperturate pollen appears to be common among IUCN Red List category and conservation data. Mendoncia in both the Old World and the New Mendoncia delphina is known only from two sites ca. World, grains with three to six apertures have been 26 km apart. In order to calculate an approximate reported for the genus (e.g., Muller et al., 1989). EOO, a 1-km 2 grid was centered on each site, and Pollen of M. delphina appears to differ from that of M. these two grids were linked at their outermost kely by having microverrucate (vs. smooth) rugulae (eastern and western) corners to calculate a total (Fig. 3E, F). EOO of ca. 32 km 2. The AOO is 8 km 2. Based on either the EOO or the AOO, this species would be Paratype. MADAGASCAR. Toliara: Préfecture de Ft. Dauphin, Foret ˆ Manantantely, N. Dumetz 1395 (MO). considered Critically Endangered (CR) under IUCN criterion B1 or B2 if appropriate knowledge of 3. Mendoncia kely E. Magnaghi, sp. nov. TYPE: subcriteria b or c were available. The northernmost Madagascar. Antsiranana: contreforts occidenof the two known collection sites for this species taux du massif de Marojejy (NE) près du col de occurs in the Andohahela National Park. The other Doanyanala (limite des bassins de la Lokoho et collection site occurs outside of the present de l Andraronga), N d Andapa, m, boundaries of the park and is undoubtedly at greater foret ˆ ombrophile sur latérite de gneiss, 25 Jan. risk of habitat loss, as noted above. Based on these 25 Feb (fl.), J.-H. Humbert data, we propose a provisional assessment of (holotype, P ; isotypes, K, MO). Endangered (EN) for M. delphina. Figures 3A, E, 4. Phenology. Plants of Mendoncia delphina have been collected in flower in October and November; a Diagnosis. Mendoncia kely E. Magnaghi differs from M. cowanii (S. Moore) Benoist by its mostly smaller and broadly fruiting collection was made in November. elliptic (vs. mostly elliptic) leaves, shorter ( vs. 16 Etymology. The epithet derives from the Latin 28 mm) bracteoles, and shorter (13 19 vs mm) corollas. delphinus (dolphin), referring primarily to the shape of the leaves, which have a beaked apex and loosely Woody vine; young stems quadrate to quadrateresemble a dolphin, and secondarily from the locality sulcate, densely and evenly pubescent with light

6 192 Novon Figure 2. Mendoncia delphina E. Magnaghi. A. Habit. B. Calyx, nectary, and gynoecium (partially submerged in nectary). C. Close-up of stigma. D. Androecium on dissected corolla tube; note staminode lacking anther. A D from the type S. Malcomber et al (K, MO). brown to golden retrorse to erect eglandular trichomes major veins, margin ciliate. Inflorescence of 1 to mm. Leaves petiolate, petioles to 18 mm, flowers in leaf axils, flowers alternate or opposite at blades mostly broadly elliptic (length/width ), nodes, pedunculate, peduncles to 30 mm, pubescentbut sometimes varying from broadly ovate to elliptic, like young stems; bracteoles sublanceolate to ovate to mm, acute-apiculate to acute to subcircular, (4 ) mm, acute to subacuminate (to retuse) at apex, rounded at base, subacuminate to apiculate at apex, abaxial surfaces abaxial surfaces 6 densely pubescent with golden- densely pubescent with woolly, golden-brown, brown multicellular eglandular trichomes mm, eglandular trichomes to 1.5 mm, adaxial surfaces adaxial surfaces pubescent with antrorse eglandular mealy-glandular. Calyx consisting of a rigid, entire, trichomes mm, trichomes usually denser on undulating, annular or cupular ring surrounding

7 Volume 23, Number 2 Magnaghi & Daniel Mendoncia (Acanthaceae) from Madagascar Figure 3. Pollen of Mendoncia delphina E. Magnaghi and M. kely E. Magnaghi. A. Mendoncia kely, subequatorial/apertural view. B. Mendoncia delphina, equatorial/interapertural view. C. Mendoncia delphina, equatorial/apertural view. D. Mendoncia delphina, polar view. E. Mendoncia kely, close-up of rugulate surface (note smooth rugulae). F. Mendoncia delphina, close-up of surface (note microverrucate rugulae). A, E from the paratype O. Andrianantoanina et al. 11 (MO); B D and F from the type S. Malcomber et al (MO).

8 194 Novon Figure 4. Mendoncia kely E. Magnaghi. A. Habit with flowers. B. Bracteoles after corolla has fallen, showing gynoecium. C. Drupe, with calyx indicated by arrow. D. Corolla dissected to reveal androecium. E. Close-up of pubescent abaxial leaf surface with denser trichomes on major veins. A, E from the type J.-H. Humbert (K, P, MO); B, D from C. Rakotovao et al (MO); C from O. Andrianantoanina et al. 221 (MO). nectar disc, mm high, densely pubescent 19 mm, externally glabrous, tube mm, throat with light brown to golden eglandular trichomes to 1 with a central cluster of glandular trichomes at base mm; corolla pink proximally and white distally, white of lower lip lobe, apically ampliate, limb with purple spots, or purple with white spots, 13.2 mm diam., upper lip mm with rounded lobes

9 Volume 23, Number 2 Magnaghi & Daniel Mendoncia (Acanthaceae) from Madagascar mm, lower lip mm with rounded reproductive parts of Mendoncia kely compared to M. lobes mm; stamens 4 5 mm, cowanii. inserted near the middle of the corolla tube, ventral pair inserted mm distal to the dorsal pair, Discussion. Specimens of Mendoncia kely have filaments mealy glandular below thecae and onto sometimes been identified as M. cowanii, which corolla tube, thecae mm, pubescent at base flowers at the same time and shows similarities in leaf with a tuft of stout eglandular trichomes extending texture and pubescence. Characters noted in the key toward apex, connective projecting 0.2 mm beyond above distinguish it from that and other species. apex of the thecae; pollen 5-brevicolpate; style When flowers are present, the smaller size of the mm, stigma subequally 2-lobed, lobes mm. corolla readily distinguishes M. kely from M. cowanii. Drupe green (immature) to brown (mature), ovoid to Flowers of M. kely resemble those of M. flagellaris in spherical, mm mm, surface size, but their stigmas, unlike the capitate stigmas of pubescent with appressed light brown to golden M. flagellaris, are subequally 2-lobed like those of M. trichomes to mm. cowanii. Although the leaf shape in M. kely is variable as noted above, most of the leaves are Distribution and habitat. Mendoncia kely is relatively small (up to 75 mm long and 45 mm wide) endemic to Madagascar, where it occurs in both the and broadly elliptic (length/width ratio ). northern and southeastern portions of the island Leaves of M. cowanii are also variable, but they tend (former provinces of Antsiranana and Toliara). Plants to be larger (up to 138 mm long and 77 mm wide) and occur in lowland to montane, wet-evergreen forests at more narrowly elliptic (length/width ratio ). elevations between 250 and 1200 m. The bracteoles of M. kely are smaller ( vs mm) than those of M. cowanii and, unlike those of IUCN Red List category and conservation data. that species, do not appear to be filled with fluid. Mendoncia kely is known from 15 collections Paratypes. MADAGASCAR. Antsiranana: Roussettes representing at least nine sites from northern to SF, S, E, O. Andrianantoanina & R. Bezana southern Madagascar. The population of this species 213 (BR, MO); Fivondronana Antsiranana II, Parc Nat. was treated as consisting of three disjunct subpop- Montagne d Ambre, 70 km SW d Antsiranana et 9 km W ulations: a northern subpopulation (four sites; EOO du village de Marovato Scama (Anivorano), camp. 2 Andasibe, S, E, O. Andrianantoanica. 80 km ), which is about 180 km distant from the na & R. Bezana 777 (BR, MO); S d Antsiranana, près de central subpopulation; a central subpopulation (four Joffre-Ville, à la station Roussette, S, E, O. sites; EOO ca. 356 km 2 ); and a southern subpop- Andrianantoanina & B. Rocsceohclher 364 (BR, MO); près ulation (1 site; EOO estimated to be 1 km 2 ), which is d Antsalaka, Montagne d Ambre Parc Nat., S, about 1130 km from the central subpopulation. The E, O. Andrianantoanina, B. Rocsceohclher & G. Rajaonarivelo 221 (BR, MO); Sava, Andapa, Anjialavabe, 2 total EOO is ca. 58,850 km, and the AOO is 36 Ankiakabe, Tsaralanto, S, E, P. km 2. Based on the AOO, M. kely would be classified Antilahimena, Randriamandimby, Tianina & Patrio as Endangered (EN) under IUCN criterion B2 if 4587 (CAS); Montagne d Ambre, T. Daniel & M. appropriate knowledge of subcriteria b or c were Butterwick 6742 (CAS), T. Daniel, H. Ranarivelo & A. Razafimanantsoa (CAS); WNW of Andapa, NW of available. Several collection sites from the northern Ambodisatrana, S, E, G. McPherson subpopulation occur in Montagne d Ambre National (MO); Sava, Andapa, Ambodivohitra, Tsaralanto, 12 km Park, at least one site in the central subpopulation SW de Anjialavabe et N d Anjialavahely, S, occurs adjacent to the Marojejy Nature Reserve, and E, C. Rakotovao, J. Ihasotra & J. Rabevazaha 2740 (MO); Montagne d Ambre, partie central, S, the sole site in the southern subpopulation is in the E, S. Ramandimbimanana et al. SDR047 (G); Andohahela National Park. In view of its broad Antalaha, Andapa, Bealampona, Mandritsarahely, SW distribution (though apparently fragmented based on d Andapa, Rés. Spéc. Anjanaharibe-Sud, vers Ranomafaknown collections) and occurrence in at least two na, 5.5 km SW de Befingotra, camp. 1, S, protected lands, we propose a provisional assess E, D. Ravelonarivo, B. Lewis & R. Rabesonina 451 (MO); Montagne d Ambre, partie central, S, ment of Vulnerable (VU) for M. kely E, S. Trigui, M. Razanajatovo & S. Ramandimbimanana Phenology. Flowering collections of Mendoncia SMT220 (G), SMT306 (G). Toliara: Andohahela RNI, Parcelle 1, Itrotroky River, NW of Eminiminy, Ft. kely have been made in January and February; Dauphin, S, E, O. Andrianantoanina, H. van fruiting collections are known from July to November. der Werff, C. Hemingway, M. van Bergen, S. Rapanarivo, S. Malcomber, P. Rakotomalaza & B. Randriamampionona Etymology. The epithet is taken from a Malagasy 11 (BR, G, MO). expression for small (kely ray or kely reny) and refers Acknowledgments. This study is derived, in part, to the diminutive size of many of the leaves and from a Master of Science thesis earned at San

10 196 Novon Francisco State University by the first author. Our Daniel, T. F Pollen morphology of Mexican 2007 studies in Madagascar were funded by the Acanthaceae: Diversity and systematic significance. Proc. Calif. Acad. Sci. 50: Lewis and Elise Rose Memorial Fund at CAS. We are Google Inc Google Earth Pro, ver ,www. grateful to H. Ranarivelo and P. Ranirison for google.com/enterprise/mapsearth/products/ assisting with field activities in Madagascar. FA- earthpro.html., accessed 24 March NAMBY (a Malagasy non-governmental organization) IUCN Guidelines for Using the IUCN Red List permitted studies at Andrafiamena. Sarah Adler Categories and Criteria, ver. 11., org/documents/redlistguidelines.pdf., accessed 4 skillfully rendered the illustrations, and S. Serata February assisted with SEM. The following herbaria generously Magnaghi, E. B Systematics and Distribution of Old made specimens available for our studies: BR, CAS, World Mendoncia (Acanthaceae: Thunbergioideae). M.S. G, K, MO, TAN, and P. Thesis, San Francisco State University, San Francisco. McDade, L. A., T. F. Daniel & C. A. Kiel Toward a Literature Cited comprehensive understanding of phylogenetic relationships among lineages of Acanthaceae s. l. (Lamiales). Benoist, R Contribution à la connaissance des Amer. J. Bot. 95: Acanthacées africaines et malgaches. Notul. Syst. (Paris) Muller, J., M. Schuller, H. Straka & B. Friedrich : Palynologia Madagassica et Mascarenica. Fam. 182: Benoist, R Acanthacées. Pp in H. Humbert Acanthaceae. Trop. Subtrop. Pflanzenwelt 67: (editor), Flore de Madagascar et des Comores (Plantes Preston-Mafham, K Madagascar: A Natural History. Vasculaires). Museum National d Histoire Naturelle, Facts on File, Oxford. Paris. Raj, B Pollen morphological studies in the Borg, A. J., L. A. McDade & J. Schönenberger Acanthaceae. Grana Palynol. 3: Molecular phylogenetics and morphological evolution of Scotland, R. W. & K. Vollesen Classification of Thunbergioideae (Acanthaceae). Taxon 57: Acanthaceae. Kew Bull. 55: