Psychotria palifera C.M. Taylor
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|Psychotria palifera C.M. Taylor|
Holotypus: MADAGASCAR. Reg. Analanjiforo [Prov. Toamasina]: Ambatondrazaka, Réserve Naturelle Intégrale de Zahamena , 15–20 km au SE d’Ambarifotsy, Rivière d’Ihofika, 17 °39'46"S 48°59'05"E, 560 – 630 m, 27. V GoogleMaps .2003, Andrianjafy et al. 368 (MO-5789365!; iso-: TAN).
Psychotria palifera C.M. Taylor is distinguished from its congeners by its relatively large stipules with a distinctive shape in bud, along with the combination of its obovate, domatiate leaf blades that are subtruncate to shallowly retuse at the apex, relatively short, fasciculate, rounded inflorescences that often become overtopped by stem growth from both axils, and well developed lobed calyx limbs; the stipules of this new species are fused around the stem in the basal part into a tubular sheath and in the upper portion free, obovate, acute to acuminate or bidenticulate, and with dense longitudinal veins or fibers.
Shrubs and small trees, collected in flower variously at 6–12 m tall, branched; stems weakly flattened becoming terete, glabrous. Leaves opposite; petiole 5–30 mm, glabrous; blade obovate, 5–18 × 2.8–11.5 cm, at base cuneate to obtuse, at apex subtruncate to shallowly retuse then abruptly acuminate with tip 1–3 mm, drying papyraceous to chartaceous, on both surfaces glabrous and weakly shiny; secondary veins 6 to 12 pairs, looping to interconnect near margins at least in apical part of blade, without intersecondary veins or usually with 1 intersecondary vein present between pairs of secondary veins, with regularly developed crypt domatia, adaxially venation plane or costa and sometimes secondary veins prominulous, abaxially costa and secondary veins prominent and remaining venation plane. Stipules fused around stem, caducous, glabrous on both surfaces, sheath portion cylindrical, 3–12 mm, free upper portion ovate, 5–15 mm, with dense longitudinal fibers or veins, acute to acuminate or bidenticulate. Inflorescences terminal, cymose, with cymes borne on fasiculate axes, the group of axes sessile to subsessile, glabrous to densely pilosulous with trichomes 0.1–0.2 mm; flowering-bearing portion rounded-corymbiform, 3–6 × 4–8 cm, branched to 2 or 3 orders, c. 30- to 60-flowered; bracts reduced or broadly triangular, 0.1–0.5 mm, acute; pedicels 0.5–1 mm. Flowers all pedicellate in dichasial cymes of 5 to 9, 5-merous; hypanthium obconic, c. 0.8 mm, glabrous; calyx limb 1–1.2 mm, glabrous, denticulate or lobed to 1/3 of its length, lobes triangular, acute to obtuse; corolla funnelform, yellow, externally glabrous, tube c. 3.5 mm, c. 1.5 mm in diam. near middle, internally densely pilose in upper part with trichomes c. 1 mm, lobes ligulate to triangular, c. 2 mm, obtuse, adaxially shortly galeate, abaxially smooth; stamens inserted in upper part of corolla tube, filaments not seen, anthers c. 0.8 mm, included or partially exserted, positioned with tips at top of corolla tube; style c. 4 mm, stigmas c. 1 mm. Infructescences similar to inflorescences or often markedly overtopped by growth from subtending axils. Fruits subglobose, c. 5 mm diam., glabrous, red, with calyx limb 1.5–4 mm, lobed for 1/4–1/2 of its length; pyrenes 2, hemispherical, adaxially plane, abaxially smooth; endosperm densely deeply ruminate on both surfaces.
Etymology. – The profile shape of the stipules in bud resembles a spade, and the specific epithet refers to that implement.
Habitat, distribution and phenology. – Psychotria palifera has been collected in humid evergreen forest at 560–1100 m in central eastern Madagascar (Toamasina), with flowers in March and with fruits in May and June .
Conservation status. – Psychotria palifera is known from ten specimen collections representing nine unique occurrences in humid evergreen forest at 560–1100 m elevation. The EOO of the species is 2,716 km ², within the limits for “Endangered” under IUCN Red List Criterion B1; and the AOO is 36 km ², also within the limits for “Endangered” under Criterion B2 ( IUCN, 2012). Six of the collection sites are within two protected areas: Zahamena PA (two sites), with a good protection loss of this site would significantly decrease the EOO for this species as well as the AOO. Forested areas without formal protection in the vicinity of these protected areas are subject to degradation by small-scale slash and burn agriculture and fire to create habitat for cattle farming as well as resource exploitation including logging, hunting and mining ( GOODMAN et al., 2018). All three of the unprotected collecting sites would be subject to such threats. The northernmost two collections are within Zahamena PA, and constitute a single “location” (sensu IUCN, 2012, 2019). Three collections are from the northernmost parcel of Ankeniheny-Zahamena Corridor adjacent to Zahamena PA, and constitute a second location. One collection is from forêt d’Ambatoaragnana, Ankeniheny- Zahamena Corridor PA and constitutes a third location. Each of the three widely separated unprotected sites constitutes a separate location, giving a total of six locations. Three locations are in generally well-protected areas, but the other three are in vulnerable areas, one of which may already be lost, causing a decline in EOO and AOO, extent and/or quality of habitat, number of locations and number of mature individuals. Thus, the Red List status of P. palifera is assessed as “Vulnerable” [VU B1ab(i,ii,iii,iv,v)+2ab(i,ii,iii,iv,v)].
[Rasoanindriana et al. 169] [Photo: S.E. Rakotoarisoa]
level; and Ankeniheny-Zahamena Corridor PA (4 sites), with the interior of the reserve well protected but parts close to villages seriously impacted by exploitation of timber and, to a lesser extent, shifting cultivation (C. Birkinshaw, pers. comm.). The remaining three collection sites are in unprotected areas: one to the east of Ankeniheny-Zahamena Corridor PA and c. 4 km west of Betampona PA (forêt secondaire de Sondrimaro); one c. 200 m outside Ankeniheny-Zahamena Corridor in secondary forest (Ambodivoromborogna Forest); and the third c. 11 km east of the southernmost parcels of Ankeniheny- Zahamena Corridor PA. This distant collection is from 1975 in an area with no protection, near a major roadway; it very likely that the population no longer exists at this collection site. The
Notes. – Psychotria palifera is characterized by the combination of its relatively robust habit for its genus; glabrous vegetative structures; medium-sized obovate leaf blades with small domatia in the abaxial axils and the apices truncate to shallowly retuse; distinctive large stipules; relatively short, fasciculate, corymbiform inflorescences that often become overtopped by growth from both subtending axils; welldeveloped lobed calyx limbs; medium-sized fruits; pyrenes that are smooth abaxially; and endosperm that is reticulate on both surfaces. The specimens dry with a dark brown color, which is also seen in the sapwood of cut stems. The stipules of P. palifera are unusually well developed and distinctive in form: these are fused in their basal portions into a cylindrical sheath, then the upper parts are ovate, venose or fibrose, and initially pressed flat together with their margins rolling back. These stipules also vary markedly in size, from smaller ones on terminal buds of stems with young leaves to those that are mature and dehiscing. These stipules are similar in form to those of P. hamifera , newly described above. Only one fully developed flower has been seen, and was not dissected in order to preserve it; it agrees with the long-styled form of distylous Psychotria species , but whether P. palifera is distylous cannot be determined. The infructescences are relatively short and have a characteristic form that is commonly seen in Psychotria , with the flowers borne in groups on several axes, and these axes then arising from a node that either bears developed leaves, or bears an internode that is hardly developed (1–2 mm long) and then another node that bears bracts and the axes. This arrangement was analyzed in detail by TAYLOR (2020), who noted that it is part of continuous variation in inflorescence form in this genus, and its interpretation as pedunculate vs. subsessile or fasciculate depends on the degree of development of the leaf-like structures borne from the subtending node: if these are small they are considered bracts and the inflorescence pedunculate, while if they are enlarged and green they are considered leaves and the inflorescence sessile to subsessile and fasciculate. In P. palifera the inflorescences seen are subtended by developed leaves, and are also overtopped or enclosed by leafy stems that develop form one or usually both of the axils of the subtending leaves. The development of a reduced internode just above the leaves that subtend the inflorescence is variable in P. palifera , so it is sometimes evident and sometimes apparently did not develop. The calyx limb appears to enlarge as the fruit develops, to almost twice its size at anthesis, but the flowers are not well enough documented to confirm this. This species agrees with BREMEKAMP (1963) ’s Mapouria Group VII. A fruiting specimen is is chosen as the type because most of the characters that contrast it with the similar species Bremekamp treated are in the fruits and seeds.
Psychotria palifera is similar to P. imerinensis (Bremek.) A.P. Davis & Govaerts and P. manampamihensis (Bremek.) A.P. Davis & Govaerts , which both can be separated by their shorter ligulate to ovate stipules, 2–5 mm long, and their shorter, truncate to denticulate calyx limbs, 0.5–1 mm long, and they also often differ by their narrower leaf blades, 1.2–6 cm wide.
Paratypi. – MADAGASCAR. Reg. Analanjirofo [Toamasina]: Ambatondrazaka, RN Zahamena, 17°39'46"S 48°59'05"E, 560 GoogleMaps – 360 m, 27. V .2003, Andrianjafy et al. 371 ( CNARP, MO, P, TEF); Vavatenina, à 10 km au N d’Anamborano, 17°42'04"S 49°00'48"E – 17°42'04"S 49°00'48"E, 620 m, 12. VI .2004, Andrianjafy et al. 436 ( CNARP, MO, P, TEF); PN Zahamena, 17°33'30"S 48°53'35"E GoogleMaps , 800 m, 5. V .2003, Rakotonandrasana 688 ( CNARP, MO, TEF); sine loco, 17°41'27"S 48°59'52"E GoogleMaps , 13.VI.2001 GoogleMaps , Randrianjanaka GoogleMaps et al. 611 (CNARP, MO, P, TEF); Miarinarivo-Anamborano, Ambinanisavaharina, 17°41'15"S 49°00'02"E, 580–620 m, 14. VI .2001, Ratovoson et al. 492 ( CNARP, MO, P, TEF); Atsinanana, along rte #2, 41 km E of Perinet, [18°55'00"S 48°37'00"E] GoogleMaps , 600 m, 1.III.1975, Croat 32639 ( MO); Toamasina II, Sahambala, Sahavongo, forêt d’Ambodivoromborogna, 18°01'41"S 49°05'37"E GoogleMaps , 608 m, 20. V .2017, Ralaijaona & Syde 159 ( K, MO, P, TAN); forêt d’Ambatoaragnana, 18°01'31"S 49°04'59"E GoogleMaps , 675 m, 21. V .2017, Rasoanindriana et al. 169 ( K, MO, P, TAN); Ambodirafia, proche Lohanifotsy, 17°52'54"S 49°09'56"E GoogleMaps , 452 m, 20. VI .2017, Rasoanindriana et al. 201 ( BR, K, MO, P, TAN) .
Holotypus: MADAGASCAR. Reg. Haute Matsiatra [Prov. Fianarantsoa]: 2 km W de la gare Andrambovato, côté riv. Tatamaly , 21°30'07"S 47°24'06"E, 1075 m, 13.X.2000, Rakotovao & Randiantafika 1011 (MO-04804424!; iso-: TAN) GoogleMaps .
abaxially weakly ridged pyrenes, and endosperm that is ruminate on both surfaces.
Shrubs and small trees, collected in flower and fruit variously at 3–7 m tall, branched; stems glabrous, weakly flattened becoming subterete. Leaves opposite; petiole 4–42 mm, glabrous; blade obovate, 4–7 × 3–11 cm, at base acute to obtuse, at apex retuse for 1/5–1/4 of its length with sinus broadly rounded and sometimes costa terminating in an apiculate tip to 1 mm, drying papyraceous to chartaceous, glabrous on both surfaces; secondary veins 8 to 11 pairs, weakly looping to interconnect, without interescondary veins or with reticulated tertiary veins extending between some pairs of secondary veins, without domatia, adaxially costa prominulous and remaining venation plane or secondary veins sometimes thickened, abaxially costa and secondary veins prominent, loosely reticulated tertiary venation plane to prominulous, and remaining venation plane and not visible. Stipules fused around the stem for their entire length, caducous, externally glabrous, 2–4.5 mm, interpetiolarly truncate or rounded to obtusely angled, adaxially pilosulous with trichomes 0.8–1 mm, entire to bidenticulate. Inflorescences terminal, cymose, pedunculate, glabrous to moderately pilosulous with trichomes 0.1–0.3 mm, subtended by 1 reduced internode to 1 cm, at top with truncate to rounded stipules 1.5–3 mm, without leaves or with foliaceous bracts up to 2 cm; peduncles 0.6–4.5 cm; branched portion roundedcorymbiform, 2 –5.5 × 4– 9 cm, branched to 2 or 3 orders, c. 35- to 70-flowered; bracts triangular to rounded, 0.2–1 mm, aristate to bilobed or fimbriate; pedicels 0.5–2.5 mm. Flowers all pedicellate in dichasial cymes of 5 to 7, 5-merous, distylous; hypanthium ellipsoid to obconic, c. 1 mm, glabrous to puberulous; calyx limb 1–1.2 mm, glabrous, truncate to denticulate; corolla funnelform, yellow, externally glabrous, tube 4.5–5 mm, 1.2–1.5 mm diam. near middle, internally in upper part with densely pilosulous ring c. 1 mm wide with trichomes c. 0.3 mm, lobes triangular to narrowly triangular, 1.5–2 mm, at tip acute, adaxially galeate, abaxially with rounded thickening; stamens in short-styled form inserted in upper part of corolla tube, filaments c. 1 mm, anthers c. 1.5 mm, partly to fully exserted, in long-styled form filaments c. 0.3 mm, anthers c. 1 mm, included with tips positioned at top of corolla tube; style in short-styled form 2–3 mm, stigmas 0.5 –1 mm and included, in long-styled form style 5.5–6 mm, stigmas 0.8–1 mm and exserted. Infrutescences similar to inflorescences. Fruits ellipsoid to subglobose, 5–6 mm diam., glabrous, red; pyrenes 2, hemispherical, adaxially with 3–4 weak rounded longitudinal ribs; endosperm densely deeply ruminate on both surfaces.
Etymology. – This elegant, tall, distinctive new species named for one of its collectors, Dr. Sylvain Razafimandimbison, a leading specialist in Malagasy Rubiaceae and a native of Madagascar.
Habitat, distribution and phenolog y. – Psychotria razafimandimbisonii has been collected in humid evergreen forest at 400–1100 m in central eastern Madagascar (Fianarantsoa), with flowers in August through November and with fruits in April and October. This species is unusual for Psychotria in being represented by more flowering than fruiting specimens.
Conservation status. – Psychotria razafimandimbisonii is known from 13 specimen collections representing 12 unique occurrences in humid evergreen forest at 400–1100 m elevation. The EOO of the species is 322 km ², within the limits for “Endangered” under IUCN Red List Criterion B1; and the AOO is 32 km ², also within the limits for “Endangered” under Criterion B2 ( IUCN, 2012). Eleven of the collection sites are within two protected areas: Fandrina Vondrozo Forest Corridor Reserve, whose interior is well protected but parts close to villages seriously impacted by exploitation of timber and shifting cultivation (C. Birkinshaw, pers. comm.); and the well-protected Ranomafana PA. One collection is on the eastern boundary of Ranomafana PA and label information indicates the habitat is infested with invasive Psidium L.; this constitutes a single location (sensu IUCN, 2012) based on the unique threats known to occur. It is likely that the border edges of the protected area are subject to degradation by small-scale slash and burn agriculture and resource exploitation including logging, hunting and mining ( GOODMAN et al., 2018) and this location may have loss of suitable habitat. The northernmost collection is within Fandrina Vondrozo Forest Corridor Reserve, and constitutes a second location. The southernmost collection is also within Fandrina Vondrozo, along the Tatamaly River, a possible sensitive habitat due to the water resources and in a separate parcel of the Reserve, and is considered a third location. The remaining collection sites are all in the interior of Ranomafana PA and thus not subject to the threats at the first location. These constitute another location for a total of four locations with respect to known threats. Three locations are in generally well-protected areas but the fourth can be considered vulnerable for loss of habitat quality, thus the Red List status of P. razafimandimbisonii is assessed as “Endangered” [EN B1ab(iii)+2ab(iii)].
Notes. – Psychotria razafimandimbisonii is characterized by the combination of its glabrous vegetative structures; medium-sized leaves that are markedly retuse at the top and lack domatia; relatively small, broadly rounded to truncate stipules; pedunculate corymbiform inflorescences with the flowers pedicellate in dichasial groups and the peduncle borne on a reduced internode topped by a leafless node; truncate to denticulate, medium-sized calyx limbs; slender, yellow, medium-sized corollas; medium-sized ellipsoid fruits; pyrenes that are weakly ridged abaxially; and endosperm that is ruminate on both surfaces. The specimens characteristically dry brown or yellowish brown. The markedly retuse leaves are distinctive, and unusual in Rubiaceae . These are easily mistaken at first glance for leaves that are damaged by insect or physical tearing, but the complete leaves consistently have this form. The sinus is usually edged for part or all of its length by the distalmost pair of secondary veins, with no blade tissue extending out from them on the distal side. The secondary leaf veins are not markedly more closely set near the apex, so the retuse shape appears to be due to additional growth of the lamina and marginal tissue rather than to a foreshortened central portion and costa. The margins of most of the leaf blades are thinly revolute, but this may be an artifact of drying rather than a character of the living plants. The adaxial trichomes of the stipules are a little longer than the sheath, so these extend past it and can appear to be marginal cilia; whether these trichomes are covered by the stipule in life then the stipule shrinks in drying is not known. The inflorescences are distinctive in the reduced stem segment at the base of the peduncle, which is leafless but has two truncate stipules. The inflorescences are described on some labels as whitened. This new species agrees with BREMEKAMP (1963) ’s Mapouria Group VI.
Psychotria razaf imandimbisonii is similar to P. retusa (Bremek.) A.P. Davis & Govaerts , which can be separated by its leaves that are only shallowly retuse at the apex and have regularly developed domatia in the abaxial vein axils, larger stipules 9–25 mm long, abaxially smooth pyrenes, and endosperm that is ruminate only on the abaxial side. Psychotria razafimandimbisonii is also similar to P. manampamihensis (Bremek.) A.P. Davis & Govaerts , which can be separated by its generally smaller, obovate leaf blades, 5–12 × 3–6 cm, that have regularly developed domatia and are obtuse to truncate at the apex with acuminate tips 1–8 mm long, stipules that are slenderly acuminate at the top, and flowers that are subsessile or borne on shorter pedicels, up to 0.5 mm long.
Paratypi. – MADAGASCAR. Reg. Vatovavy-Fitovinanay [ Prov. Fianarantsoa]: distr. Ifadenia, Ranomafana PN , 21°15'25"S 47°25'30"E, 918 m, 24.IV.2005, Acevedo-Rodríguez & Razafindraibe 14457 ( MO, US) GoogleMaps ; 21°02'00"S 47°18'00"E – 21°25'00"S 47°37'00"E, 400–1375 m, 14.VIII.1988, Kremen 88-1 ( MO) GoogleMaps ; 21°15'36"S 47°25'12"E, 854–915 m, 1.XI.1998, Almeda 7932 ( CAS, MO, TAN) GoogleMaps ; 21°16'S 47°25'E, 900–1100 m, 31.X.1998, Daniel 9011 ( CAS, MO, TAN) GoogleMaps ; 21°16'S 47°26'E, 1000–1100 m, 4.XI.1998, Daniel 9098 ( CAS, MO, TAN) GoogleMaps ; 21°15'S 47°25'E, 1100 m, 3.XI.1997, Davis et al. 1045 ( K, MO, TAN) GoogleMaps ; 21°15'S 47°27'E, 800–1000 m, 3–8.IX.1993, Kotozafy et al. 212 ( MO) GoogleMaps ; ibid. loco GoogleMaps , 900 m, 19–21.X.1993, Kotozafy & Randriamanantena 335 ( MO) ; 21°15'S 47°28'E, 950 m, 3.X.1986, Nicoll 116 ( MO) GoogleMaps ; 7 km W of Ranomafana, Duke University Primate Center , 21°16'S 47°25'E, 1000 m, 27.X.1987, Overdorff 39 ( MO) GoogleMaps ; 21°15'50"S 47°25'17"E, 998 m, 14.IV.2016, Razafimandimbison et al. 1381 ( MO, S, TAN) GoogleMaps ; Vatoharanana, 40 km S Ranomafana (ville) , 21°17'04"S 47°26'00"E, 1025 m, 2.X.2000, Rakotovao & Randriantafika 957 ( MO, P, TAN) GoogleMaps .
Royal British Columbia Museum - Herbarium
Missouri Botanical Garden
Museum National d' Histoire Naturelle, Paris (MNHN) - Vascular Plants
Centre National de la Recherche Appliquée au Developement Rural
Mykotektet, National Veterinary Institute
Royal Botanic Garden Edinburgh
Royal Botanic Gardens
Parc de Tsimbazaza
Embrapa Agrobiology Diazothrophic Microbial Culture Collection
Naturhistorisches Museum Wien
California Academy of Sciences
Department of Botany, Swedish Museum of Natural History
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