Additional notes on Ipomoea (Convolvulaceae) in Bolivia
109 species of Ipomoea L. are recorded from Bolivia. This total includes six new country records and two species, I. prolifera J. R. I. Wood & Scotland and I. inaccessa J. R. I. Wood & Scotland, which are described as new, while I. jujuyensis O’Donell is excluded having been included previously in error. The little-known I. subalata Hassl. is described in full and compared with I. chondrosepala Hallier f. and other species with which it has been confused. The paper is illustrated with line drawings, photographs and distribution maps of the main species discussed.
Key Wordsnew records new species Paraguay taxonomy
More species of Ipomoea L. are recorded from Bolivia than from any other country in the Americas apart from Mexico and Brazil. 102 species were listed and described by Wood et al. (2015) and the number has now risen to 109 as a result of further field work and herbarium studies. Although there is no easy explanation, the most important factor for this large number appears to be the latitude where Bolivia lies. Both in the northern and the southern hemispheres of the American continent, the greatest diversity in Ipomoea is found between 15 and 25 degrees latitude with diminishing numbers recorded from equatorial regions as well as from the more temperate zones. The diversity of habitats and the presence of several distinct biomes are clearly other factors but are unlikely to be the sole reasons, as some equatorial countries, such as Colombia, have many fewer species of Ipomoea despite their greater size and even more varied habitats.
Since the publication of our account of Ipomoea in Bolivia (Wood et al. 2015), we have had the opportunity to carry out fieldwork in both Bolivia and Paraguay and examine herbarium collections in both these countries and elsewhere. This has resulted in many new records of recognised species and a greater understanding of the characteristics, distribution and conservation status of a number of recently described species, especially I. appendiculata J. R. I. Wood & Scotland, I. juliagutierreziae J. R. I. Wood & Scotland, I. lactifera J. R. I. Wood & Scotland, I. longibarbis J. R. I. Wood & Scotland, I. mucronatoproducta J. R. I. Wood & Scotland and I. odontophylla J. R. I. Wood & Scotland. Also, of particular interest is a new record of I. acanthocarpa (Choisy) Asch. & Schweinf. from Ibiato in Cercado, Beni (Martinez 38), only the second record from Bolivia and this 62 years after the only previous record. We have also seen specimens from other countries of species previously thought to be Bolivian endemics including I. chiquitensis J. R. I. Wood & Scotland in Brazil (Wood et al. 2017a) and I. mucronifolia J. R. I. Wood & Scotland in Paraguay (Wood et al. 2017b).
It is now clear that we cited Ipomoea jujuyensis O’Donell in error for I. squamosa Choisy and so I. jujuyensis should be excluded from the list of accepted species for Bolivia. Molecular sequencing of the sole Bolivian collection, Mendoza et al. 2622 showed clearly that it belonged to I. squamosa and this was confirmed by re-examination of the specimen. The absence of I. jujuyensis from Bolivia, however, is odd given its presence in the Andes of Argentina, Peru and Ecuador.
The following two recently discovered species from Bolivia are new to science:
Ipomoea prolifera J. R. I. Wood & Scotland, sp. nov. Type: Bolivia. Dept. Santa Cruz, Prov. Vallegrande, on descent to Pampa Negra, 15 March 2018, J. R. I. Wood, M. T. Martinez & G. Aramayo 28441 (holotype USZ; isotypes LPB, OXF).
recognition. The pubescent sepals and corolla clearly place this species in a large radiation of mainly South American species of Ipomoea that share these characteristics and this is confirmed by unpublished molecular data. However, the dimorphic leaves are unusual and serve to separate it immediately from species with somewhat similar palmatilobed leaves such as I. stuckertii O’Donell, I. padillae O’Donell and I. pampeana P. P. A. Ferreira & Miotto. Ipomoea cardenasiana O’Donell does present dimorphic leaves but is readily distinguished by its much larger sepals (15 – 22 mm, not 8 – 9 mm long). The subterminal inflorescence is unusual in decumbent species but is somewhat similar to that of I. mendozae J. R. I. Wood & Scotland but the cuneate-based, ovate to subrhomboid, entire leaves of that species are very different from the dimorphic, cordate-based leaves of I. prolifera. The proliferating inflorescence is also unusual. Unpublished molecular data suggest this species is closely related to I. cuneifolia Meisn and I. bombycina (Choisy) Benth. & Hook. f., but there is no obvious close morphological resemblance between these three species.
specimens examined. bolivia. Santa Cruz: Prov. Vallegrande, on descent to Pampa Negra, 18°27.746'S 64°16.322'W, 1802 m, 15 March 2018, J. R. I. Wood et al. 28441 (holotype USZ; isotypes LPB, OXF); ibid., 18°28.202'S 64°16.127'W, 1671 m, J. R. I. Wood et al. 28443 (LPB, OXF, USZ).
conservation status. This is a narrowly endemic species, known only from a single location, but similar dry bushland covers the hill slopes in the area and it may prove to be more common than is apparent, particularly, if as seems to be the case, it only flowers in years of good rainfall. There is no obvious threat to its habitat, so it should be classified as data deficient (DD) within IUCN guidelines until accurate population studies can be carried out.
phenology. Found in flower in March but probably continuing to flower into April. This plant was not seen in 2013, a relatively dry year, at the same location and in the same season, so flowering may be very dependent on adequate rainfall.
etymology. This species is named Ipomoea prolifera because of its proliferating inflorescences.
notes. This species is yet another endemic to the hot, dry inter-Andean valley system of the Rio Grande and Rio Mizque. Although we did not find any other endemic species in the immediate vicinity of the two collections, both Ipomoea juliagutierreziae and Varronia lantanifolia J. S. Miller & J. R. I. Wood were found nearby and other Bolivian endemic species are found at the bottom of the valley.
Ipomoea inaccessa J. R. I. Wood & Scotland, sp. nov. Type: Bolivia. Dept. La Paz, Prov. Caranavi, on W side of Serrania de Bellavista, above Carrasco, J. R. I. Wood & S. G. Beck 28539 (holotype LPB; isotypes K, OXF, USZ).
recognition. A very vigorous liana reaching heights unattained by most species of Ipomoea. Herbarium specimens are most likely to be confused with I. philomega (Vell.) House but that species has smaller, deep pink corollas, 5 – 6 cm in length and, usually, glabrous leaves and distinctive reddish sepals. I. inaccessa has a larger corolla about 9 cm long, which is white or white with a pale pink throat. Its leaves are uniformly densely puberulent on both surfaces and the sepals are pale green. The flower colour and sepal shape suggest it is related to I. reticulata O’Donell and I. saopaulista O’Donell and this is confirmed by unpublished molecular sequence data. However, the much larger dimensions of the sepals (13 – 20 mm long, not 7 – 10 mm) and corolla (9 – 9.5 cm long, not 2.5 – 5 cm) rule out both these species.
distribution & habitat. Endemic to moist hill forest with frequent cloud at 1400 – 1500 m on the west side of the Serrania de Bellavista in Caranavi Province of the La Paz Department. Map 1.
specimens examined. bolivia. La Paz: Prov. Caranavi, on road from Caranavi to Puerto Linares, 1500 m, 5 July 1980, T. Feuerer & N. Höhne 4662 (LPB); Caranavi hacia Alto Beni, subiendo 9 km hacia Buenavista, 16°18'S 67°50'W, 1400 m, 5 May 1989, S. G. Beck 17205 (LPB); on W side of Serrania de Bellavista, above Carrasco, 15°42'21"S 67°29'454"W, 1420 m, J. R. I. Wood & S. G. Beck 28539 (holotype LPB; isotypes K, OXF, USZ); ibid.,15°40'49"S 67°29'524"W, 1525 m, J. R. I. Wood & S. G. Beck 28542 (LPB, K, OXF, USZ); ibid, J. R. I. Wood & S. G. Beck 28543 (LPB, K, OXF, USZ).
conservation status. This is a narrowly endemic species, known only from a single area of the Serrania de Bellavista. This is a large ridge, reaching about 2000 m in height, covered in moist hill forest. The habitat is vulnerable but the very steep slopes, the very moist, cloudy climate and its unsuitability for agriculture provide a degree of protection. Careful population studies are needed to assess the extent of populations of Ipomoea inaccessa in the area as the Serrania is extensive and mostly botanically unknown. It can only be treated as Data Deficient (DD) within IUCN guidelines until its populations are carefully studied.
phenology. Flowering presumably begins in March and continues into May with mature fruit present in July.
etymology. This species is named Ipomoea inaccessa because of the difficulties of collecting flowering specimens from high forest on precipitous wooded Andean slopes.
The following species are additional to those recorded for Bolivia by Wood et al. (2015):
Ipomoea deminuta J. R. I. Wood & Scotland (2017a: 9).
This was described as a new species based on a single collection (E. Gutiérrez et al. 1152) from Flor de Oro on the western side of the Rio Iténez in the Noel Kempff National Park on the Bolivian frontier with Brazil. No other collection has been seen but it might be expected to occur elsewhere in the region both in Bolivia and in Rondonia and Mato Grosso States in Brazil as the recorded habitat of flooded pampa is common in the region.
Ipomoea longeramosa Choisy (1845: 384).
This was published as a new record for Bolivia by Wood & Scotland (2017b: 6) based on Wood et al. 24770 (K, LPB, USZ) collected in cerrado on fine sandy soil near San Rafael in Velasco Province in the Chiquitania region of Eastern Bolivia.
Ipomoea lindenii M. Martens & Galeotti (1845: 264).
This species is common in Central America from Mexico southwards but is much less common in South America where it occurs sporadically along the Andes south to Peru and, now, to Bolivia. The only Bolivian collection (N. Salinas 3134 (LPB)) was made in Quime Province in the Department of La Paz between Comunidad Khora and Mikilpirhua at 1900 m. This collection is in fruit but shows the characteristic leaf shape, shortly pedunculate cymes and rostrate capsules typical of this species.
Ipomoea peruviana O’Donell (1948: 4).
Ipomoea volcanensis O’Donell (1953: 398).
This was regarded as an Argentinian endemic before it was collected in Bolivia in 2016. In Bolivia it is known from a single collection (Wood, Hind & Gutiérrez 28059 [K, LPB, OXF, USZ]) from Bosque Tucumano Boliviano at 1446 m on the ridge between Palos Blancos and Entre Rios in Tarija Department. Its occurrence in Bolivia is not unexpected but its discovery makes the absence of Ipomoea jujuyensis more surprising.
Ipomoea subalata Hassl. (Hassler 1911: 157).
This is described in full below as it is a very poorly known species and is recorded for Bolivia for the first time:
specimens examined. bolivia. Chuquisaca: Prov. Luis Calvo, Mun. Villa Vaca Guzman, pie de monte de la Serrania de Incahuasi, 19°52'47"S 63°44'16"W, 1020 m, 17 Feb. 2006, A. Lliully & Portal 725 (HSB, MO, OXF). Santa Cruz: Prov. Cordillera, Tatarenda, R. E. Fries 1451 (S); Camiri, 800 m, Feb. 1951, M. Cárdenas 4734 (LIL); Abapó, camino a Com. Morocco, 18°52'01"S 63°22'52"W, 487 m, 26 April 2012, J. R. I. Wood & F. Mamani 27477 (K, LPB, UZ); Abapó–Laguna Tatarenda, 19°03'13"S 63°28'01"W, 786 m, 23 March 2013, J. R. I. Wood et al. 27590 (K, LPB, USZ); entre Ipatí y Lagunillas, 19°47'22S 63°39'38"W, 1070 m, 25 March 2013, J. R. I. Wood et al. 27637 (K, LPB, OXF, USZ); between Ipata and Tatarenda, 19°18'45"S 63°28'54"W, 917 m, 10 March 2018, J. R. I. Wood et al. 28398 (LPB, USZ); between Abapó and Laguna Tatarenda, 19°03'15"S 63°22'51"W, 769 m, 10 March 2018, J. R. I. Wood et al. 28399 (LPB, USZ); 5 km S of Abapó on toad to Camiri, 18°57'18"S 63°23'42"W, 492 m, 26 March 2018, J. R. I. Wood et al. 28477 (LPB, USZ); 10 km S of Camiri on road to Boyuibe, 20°06'14"S 63°29'13"W, 750 m, 27 March 2018, J. R. I. Wood et al. 28398 (LPB, USZ); 5 – 10 km from Ipata along road to Vado Yeso, 19°16' 52"S 63°32'07", 861 m, 28 March 2018, J. R. I. Wood et al. 28515 (LPB, USZ). Prov. Ichilo, Buenavista, c. 2 km along road to Yapacani, 17°21'11"S 63°40'24"W, 398 m, 4 April 2013, J. R. I. Wood & B. Williams 27735 (K, LPB, USZ). Prov. Vallegrande, c. 20 km E of Vallegrande along road to Postrervalle, 18°33'29"S 63°59'09"W, 1880 m, J. R. I. Wood et al. 28431 (LPB, USZ). Tarija: Prov. Gran Chaco, 2 km from Palos Blancos towards Villamontes, M. Mendoza et al. 2662 (K, USZ); Yacunda, carretera hacia Campo Largo, 22°01'16"S 63°57'27"W, 650 m, 6 March 2006, F. Zenteno et al. 4454 (CTES, LPB).
conservation status. Using Geocat (https//www.kew.org/science/projects/geocat) the extent of occupancy of this species within Bolivia is 19,744 km2, so it might be treated as Vulnerable (VU) according to IUCN guidelines, but based on its area of occupancy of 56,000 km2, it would be categorised as Endangered (EN). The categorisation as Vulnerable (VU) would seem to be more appropriate as the species thrives in disturbed scrub by roads and around “potreros” in areas of dry scrub, where there is little likelihood of significant habitat change, even though its area of occupancy is limited. Globally, this species is difficult to categorise, as the single collection from Paraguay is over a hundred years old. As no search has been made for this species in the area of the type locality, any judgement of its global status is likely to be premature. It might be extinct in Paraguay or, on the contrary, locally frequent and unthreatened given its predilection for disturbed scrubby habitats.
phenology. In Bolivia flowering in March and April.
notes. This species was described from San Luis in northern Paraguay but has never been recollected there. It has been the source of much misunderstanding by us, as well as by other botanists. In Brazil it has been equated with the plant represented by G. & L. T. Eiten 4077A (K, NY, SP) from Maranhão which we believe is the equally poorly known Ipomoea cearensis O’Donell. More recently in Flora do Brasil 2020 (under construction) the name has been applied to Harley 21983, which is I. pterocaulis J. R. I. Wood & Scotland. In Wood et al. (2015: 59) we equated it with I. megapotamica Choisy, but this was also wrong. I. subalata differs in its larger corolla, longer glabrous sepals and different inflorescence. In our account of Ipomoea in Bolivia (Wood et al. 2015) we mistakenly widened our concept of I. chondrosepala Hallier f. (Wood et al. 2015: 52) to include plants whose corolla was pubescent in bud. We noted that specimens of this species from Paraguay had glabrous corollas and were uneasy about this anomaly at the time, as corolla indumentum is nearly always species-specific in Ipomoea. Our unease increased after observing that some Bolivian collections also had glabrous corollas but it was only after fieldwork in 2017 and 2018 in Paraguay and Bolivia, that we accepted that our concept of I. chondrosepala Hallier f., in fact consisted of two elements, I. chondrosepala itself and the little-known I. subalata.
1. Corolla glabrous on the exterior; leaves and stem glabrous……………………………………………….2
Corolla pubescent on the exterior, at least in bud and towards the tips of the midpetaline bands; leaves usually pubescent, at least on the veins beneath; stem usually thinly hirsute ……………………………………………….3
2. Stems winged; perennial herb; inflorescence of axillary cymes; corolla usually 8 – 9 cm long……………………………………………….I. pterocaulis
Stems unwinged; woody liana, the inflorescence commonly developing on leafy side shoots; corolla 5.5 – 7 cm long……………………………………………….I. chondrosepala
3. Sepals glabrous or nearly so; corolla large, 9 – 12 cm long……………………………………………….4
Sepals tomentellous; corolla 4.5 – 6 cm long……………………………………………….I. megapotamica
4. Stems winged; outer sepals 9 – 12 mm long……………………………………………….I. subalata
Stems unwinged; outer sepals 8 – 9 mm long……………………………………………….I. cearensis
Ipomoea chondrospala Hallier f. (Hallier 1899: 49).
specimens examined. bolivia. Cochabamba: Prov. Carrasco, Bulo Bulo, 17°16.404'S 64°22.957'W, 267 m, 11 July 2018, J. R. I. Wood et al. 28904 (LPB, USZ). Prov. Chapare, Espiritu Santo, 1891, M. Bang 1278 (GH, K, MO, NY, US). Prov. Tiraque, near Hotel El Puente, Villa Tunari, 16°58.955'S 65°24.464'W, 342 m, 12 July 2018, J. R. I. Wood et al. 28904 (LPB, USZ). La Paz: Prov. Sud Yungas, 2 – 3 km S of Alto Beni, Sapecho–Santa Ana, 470 m, R. Seidel & Schulte 2424 (K, LPB). Santa Cruz: Prov. Ibañez, Reserva Arubaí, 8 km de Terebinto, 17°'41'S 63°25'W, 225 m, 25 May 2006, D. Villarroel & I. Linneo 599 (USZ); La Angostura, 18°10'S 63°31'W, 775 m, 20 April 2002, Angostura, M. Nee & M. Sundue 52209 (LPB). Prov. Ichilo, P. N. Amboró, trail along Río Yapacaní towards Camp. Mataracú, 17°32'S 63°52'W, 350 m, 31 May 1998, M. Nee & L. Bohs 49535 (NY, USZ); near Hotel El Cafetal, Candelaria, Buenavista, 363 m, 17°27.79'S 63°41.659'W, 15 July 2017, J. R. I. Wood et al. 28286 (LPB, OXF); ibid., 11 July 2018, J. R. I. Wood et al. 28885 (LPB, USZ); ibid., near Mision Peniel, 17°28.123'S 63°41.358'W, J. R. I. Wood et al. 28890 (LPB, USZ). Prov. Velasco, c. 90 km S of Piso Firme on road to Santa Rosa de la Roca, 14°17'S 61°52.643'W, 25 June 2018, J. R. I. Wood et al. 28613 (LPB, USZ).
We gratefully acknowledge the financial support of the Leverhulme Trust for funding our Ipomoea Foundation Monograph project, an NERC IAA award for the illustrations and the BBSRC GCRF-IAA fund (BB/GCRF-IAA/16 and BB/GCRF-IAA/17/16) for field work in Bolivia and Paraguay in 2017 and 2018. We also benefitted from a grant by the Bentham Moxon Trust to Nicholas Hind which facilitated field work in 2016. Pablo Muñoz was funded by a BBSRC scholarship granted through the Interdisciplinary Bioscience DTP Programme. Thanks are due to Germán González and Rosa Degen de Arrúa in Paraguay and to Gina Aramayo, Roxana Ledezma, Gloria Gutiérrez, Andrea Peñarrieta, Daniel Villarroel, Rosie Clegg and Nicholas Hind in Bolivia for their help in the field; to John Baker and James Ritchie at Oxford for help with the maps, images and photographs. We are especially grateful to Stephan Beck for organising a very memorable trip to the Yungas of La Paz and to Eliana Calzadilla for preparing the illustrations which accompany the text.
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