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Please note that this is an author-produced PDF of an article accepted for publication following peer review. The definitive publisher-authenticated version is available on the publisher Web site.
Journal of Ethnopharmacology March 2021, Volume 267, Pages 113546 (12p.) https://doi.org/10.1016/j.jep.2020.113546 https://archimer.ifremer.fr/doc/00658/77018/
Archimer https://archimer.ifremer.fr
Geopolitics of bitterness: deciphering the history and cultural biogeography of Quassia amara L
Odonne Guillaume 1, *, Tareau Marc-Alexandre 1, Van Andel Tinde 2
1 LEEISA (Laboratoire Ecologie, Evolution, Interactions des SystèmLEEISA (Laboratoire Ecologie, Evolution, Interactions des Systèmes Amazoniens), CNRS, Université de Guyane, IFREMER, 97300, Cayenne, French Guiana 2 Naturalis Biodiversity Center, Leiden, the Netherlands
* Corresponding author : Guillaume Odonne, email address : guillaume.odonne@cnrs.fr
Abstract : Ethnopharmacological relevance Quassia amara L. recently came into the spotlight in French Guiana, when it became the object of a biopiracy claim. Due to the numerous use records throughout the Guiana shield, at least since the 18th century, a thorough investigation of its origin seemed relevant and timely. In the light of the Convention on Biological Diversity (CBD) and the Nagoya protocol, questions about the origin of local knowledge are important to debate. Aim of the study: Defining cultural biogeography as the dynamics through space and time of biocultural complexes, we used this theoretical framework to shed light on the complex biogeographical and cultural history of Q. amara. We explored in particular the possible transfer of medicinal knowledge on an Old World species to a botanically related New World one by enslaved Africans in Suriname. Materials and methods Historical and contemporary literature research was performed by means of digitized manuscripts, archives and databases from the 17th to the 21st century. We retrieved data from digitized herbarium vouchers in herbaria of the Botanic Garden Meise (Belgium); Naturalis Biodiversity Center (the Netherlands); Missouri Botanical Garden, the Smithsonian National Museum of Natural History, the Field Museum (USA); Royal Botanic Gardens Kew (UK); the IRD Herbarium, French Guiana and the Museum National d’Histoire Naturelle (France). Vernacular names were retrieved from literature and herbarium specimens and compared to verify the origin of Quassia amara and its uses. Results Our exploration of digitized herbarium vouchers resulted in 1287 records, of which 661 were Q. amara and 636 were Q. africana. We observed that the destiny of this species, over at least 300 years, interweaves politics, economy, culture and medicine in a very complex way. Quassia amara’s uses are difficult to attribute to specific cultural groups: the species is widely distributed in Central and South America, where it is popular among many ethnic groups. The species spread from Central to South
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Please note that this is an author-produced PDF of an article accepted for publication following peer review. The definitive publisher-authenticated version is available on the publisher Web site.
America during the early 18th century due to political and economic reasons. This migration possibly resulted from simultaneous migration by religious orders (Jesuits) from Central America to northern South America and by Carib-speaking Amerindians (from northern South America to Suriname). Subsequently, through colonial trade networks, Q. amara spread to the rest of the world. The absence of African-derived local names in the Guiana shield suggests that Q. africana was not sufficiently familiar to enslaved Africans in the region that they preserved its names and transferred the associated medicinal knowledge to Q. amara. Conclusions Cultural biogeography has proven an interesting concept to reconstruct the dynamics of biocultural interactions through space and time, while herbarium databases have shown to be useful to decipher evolution of local plant knowledge. Tracing the origin of a knowledge is nevertheless a complex adventure that deserves time and interdisciplinary studies. Graphical abstract
Keywords : cultural biogeography, herbarium specimens, colonial history, Kwasi, Suriname, French Guiana, Quassia africana
through colonial trade networks, Q. amara spread to the rest of the world. The absence of 41
African-derived local names in the Guiana shield suggests that Q. africana was not 42
sufficiently familiar to enslaved Africans in the region that they preserved its names and 43
transferred the associated medicinal knowledge to Q. amara. 44
Conclusions 45
Cultural biogeography has proven an interesting concept to reconstruct the dynamics of 46
biocultural interactions through space and time, while herbarium databases have shown to 47
be useful to decipher evolution of local plant knowledge. Tracing the origin of a knowledge is 48
nevertheless a complex adventure that deserves time and interdisciplinary studies. 49
50
51
Keywords: cultural biogeography, herbarium specimens, colonial history, Kwasi, Suriname, 52
French Guiana, Quassia africana 53
54
1. Introduction 55
56
Historical ethnobotany seeks to decipher the complex coevolution of people and plants 57
through time, and is a truly interdisciplinary field. Although the convergence between 58
historical and biological sciences is facing epistemological difficulties (Heinrich et al., 2006), 59
the study of historical texts and archives is highly valuable to understand the dynamics of the 60
relations between humans and plants from a diachronic perspective (Medeiros, 2016), and 61
particularly regarding medicinal uses (Lardos, 2015). This has been successfully applied to 62
the description of the evolution of phytotherapeutical practices and of medicinal floras, and 63
numerous examples from Europe (Lardos and Heinrich, 2013; Leonti et al., 2010), Latin-64
America (Brandão et al., 2012, 2008), the Caribbean (Soelberg et al., 2016), Asia 65
(Sathasivampillai et al., 2017) or Africa (El-Gharbaoui et al., 2017; Soelberg et al., 2015) help 66
to understand the dynamics underpinning medicinal plant knowledge and transmission. 67
Beside these historical studies, considering cultural exchanges of plants at a geographical 68
level in the contemporary era is another important field of ethnobiology, particularly 69
developed with regards to the South American and Caribbean diaspora in urban areas in 70
Europe and the US (Ceuterick et al., 2011, 2008; Pieroni and Vandebroek, 2007). Plant 71
transfers have had particular biocultural repercussions in the case of historic migrations, 72
such as the transatlantic slave trade, a subject that has been the focus of recent 73
anthropological and ethnobotanical research (van Andel et al., 2014; 2016; Voeks and 74
Rashford, 2012). Nevertheless, disentangling such processes at both a historical and a 75
geographical level is an ambitious goal, as some started before the Colombian encounter, 76
others were enhanced by the transatlantic trade, and some accelerated through colonial and 77
preindustrial times. 78
A few years ago, one particular plant species (Quassia amara L.) entered the spotlight in 79
French Guiana, when it became the object of a biopiracy claim. In 2016, the France Libertés 80
Foundation filed a patent opposition to the European Office on the legitimacy of two patents 81
deposited by the Institut pour la Recherche et le Développement (IRD) on Simalikalactone E, 82
a pharmacologically active compound against malaria and cancer, present in the leaves of Q. 83
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amara. This event became known afterwards in France as the “kwasi affair” (l’affaire 84
kwachi), after the local name of the plant in the region (Bourdy et al., 2017; Burelli, 2019; 85
Collomb, 2018; Thomas, 2018). Steering clear from the political aspects of this debate, in 86
substance as well as in form, we were interested to shed light on the complex 87
biogeographical and cultural history of this species. We observed that the destiny of this 88
plant, since at least 300 years, interweaves politics, economy, culture and medicine in a very 89
complex way, shaking up thoughts on the definition of traditional knowledge and the 90
dynamics behind it. In the framework of the Convention on Biological Diversity (CBD) and the 91
Nagoya protocol, such controversial points related to origin and ownership of knowledge are 92
to be debated carefully, being the center of political claims (Eimer, 2020; Pedrollo and 93
Kinupp, 2015). 94
Q. amara is a shrubby tree (2-8 m high) in the Simaroubaceae family, with imparipinnate 95
leaves (5-7 leaflets) and a conspicuously winged rachis. Inflorescences are racemose, flowers 96
are bisexual and bright red. The fruits are apocarpous, and consist of multiple drupes 97
(Woodson et al., 1973). Although the species is said to be native to Central America 98
(Woodson et al., 1973), the first mention of Q. amara, as well as the type specimen, come 99
from Suriname (Pulle et al., 1979). Both its local and its scientific name refer to Kwasi, an 100
enslaved African who was born in Guinea (Fermin, 1769; Dragtenstein, 2004), who possibly 101
discovered its antimalarial properties, and certainly made this species famous in Paramaribo 102
(Price, 1979). 103
The ‘scientific history’ of Q. amara reflects complex historical events of trade and exchange 104
in plants and knowledge, and may also testify of the transfer of medicinal knowledge from 105
an Old World species to a botanically related New World one by enslaved Africans in 106
Suriname. Indeed, the cultural connection with Africa is even more important, as shown by 107
recent ethnobotanical studies. Enslaved Africans contributed to the addition of 108
contemporary American medicinal flora by recognizing in the Neotropical flora a number of 109
taxa that they knew as useful in their countries of origin in western Africa. They often 110
transposed vernacular names and uses of Old World plants to the related New World species 111
(Bilby, 2000; Carney and Rosomoff, 2011; van Andel, 2016; van Andel et al., 2014). 112
Our objective is to elucidate and trace back the complex “cultural biogeography” of Q. 113
amara in colonial times by using historical written sources and herbarium vouchers. Despite 114
an increasing use of data from herbarium collections by ecologists to reveal phenological 115
trends related to climate change (Calinger et al., 2013; Hart et al., 2014; Willis et al., 2017), 116
or to investigate phytogeography by means of DNA (Roullier et al., 2013b), such material has 117
been little explored regarding geographical similarities and differences in medicinal uses of 118
plants. Exceptions include a recent study on the ‘evolution’ of Traditional Chinese Medicine 119
by using the Economic Botany collections in Kew (Brand et al., 2017) and a comparison of 120
Surinamese phytotherapy over 250 years (van Andel et al., 2012a). 121
Cultural biogeography is defined here as the dynamics through space and time of biocultural 122
complexes. In the present paper, we explain how Q. amara spread from Central to South 123
America and the rest of the world, and verify if (and why) its use as a febrifuge was 124
discovered far from its origin, and whether traditional uses of the botanically related African 125
species Quassia africana (Baill.) Baill. may help explain this key event. 126
2. Material and methods 127
2.1 Historical and contemporary literature research 128
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Following recent research in historical ethnobotany (Lardos, 2015), we traced back the 129
history of Q. amara’s uses by consulting compilations and original historic sources, in the 130
form of books (hardcopy or eBooks) retrieved on Biblioteca Nacional de Colombia (2020), 131
Google books (2020), Gallica (2020), Botanicus (2020) and journal articles. We reviewed the 132
(ethno-) botanical literature of northern South America and Central America, such as floras 133
and ethnobotanical compilations, for Q. amara and related species, and the Central and 134
West African literature for Q. africana and related species to record past and contemporary 135
uses. 136
2.2 Data mining from digitized herbarium vouchers 137
Herbarium voucher labels are an important source of ecological and (ethno-) botanical 138
information, as they often include data on geographic localities, surrounding ecosystems, 139
vernacular names, ethnic groups, medicinal or other uses that help to reconstruct the 140
(ancient) habitats, biogeography or cultural knowledge of a species. Since herbarium 141
collections are increasingly digitalized and published online, access to label information is 142
easier and allows for a greater diffusion of knowledge. Some vouchers date back to the 17th 143
century, and contain information that was once confined to museums (Boulangeat, 2014; 144
Chupin, 2018; van Andel et al., 2012c). From March to June 2020, we explored herbarium 145
databases and digitized vouchers to gather information related to contemporary or ancient 146
distribution, ecology or uses of Q. amara and Q. africana, mostly from online resources: the 147
Botanic Garden Meise (2020), Belgium (BR); Naturalis Biodiversity Center (2020), Leiden, the 148
Netherlands (L); Missouri Botanical Garden (2020), (MO); the Smithsonian National Museum 149
of Natural History (2020), (US); and the Field Museum (2020), (F) in the US; Royal Botanic 150
Gardens Kew (2020), UK (K); the IRD Herbarium (2020) in Cayenne, French Guiana (CAY); and 151
the Museum National d’Histoire Naturelle (2020), in Paris, France (P). Brazilian vouchers 152
from other institutions were also checked through the website Reflora (2020). When data 153
originated from a herbarium specimen, we cited it with the herbarium abbreviation, the 154
name of the first collector and the specimen number (e.g., MO-Smith 15674). 155
2.3 Linguistic comparison of folk names 156
The study of local plant names is a useful way to decipher biocultural history, as local names 157
often travel with the plants and are exchanged among different cultural groups (Balée, 2003; 158
Balée and Moore, 1991; Grenand, 1995; van Andel et al., 2014). In order to have a clear 159
insight into Q. amara’s travel through the Americas, and to verify the transatlantic 160
hypothesis of African plant knowledge applied to the flora of the Americas, local names from 161
both Q. amara and Q. africana were collected from literature, herbarium labels and 162
databases and compared. According to methods previously published (van Andel et al., 163
2014) our comparison was based on phonological, morphosyntactic, and semantic 164
similarities between all American names for Q. amara (from Amerindian, European or Afro-165
American languages) as well as for names for Q. africana in African languages. A strong 166
correspondence in sound, structure, and semantics between vernacular names in different 167
languages was considered as evidence for a shared origin. We paid attention not to cite 168
redundant information and refer to primary sources as much as possible. 169
3. Results and discussion 170
3.1 Herbarium data mining 171
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Exploration of digitized herbarium vouchers and labels resulted in 1287 records, of which 172
661 were Q. amara and 636 were Q. africana, or labelled as such, because a few of them 173
were misidentified. For our analysis, we used virtual specimens from L (102), P (157), US 174
(135), BR (30), F (26), CAY (23), K (1), and labels from MO (187). For Q. africana, virtual 175
specimens were analyzed from BR (436), L (184), US (13) and K (3). All specimen information 176
directly cited in our analysis is listed in Appendix A. 177
3.2 A rapid history of Quassia’s name and fame 178
The febrifugal and stomachic properties are said to have been discovered by an enslaved 179
African in Suriname, known as Kwasi or Quassie van Nieuw Timotibo (Dragtenstein, 2004). 180
Born between 1692 and 1697 in ‘Guinea’, Kwasi was enslaved before 1712 and put to work 181
around 1727 on the plantation Nieuw Timotibo in the former Dutch colony (Dragtenstein, 182
2004; Stedman, 1796). As Kwasi is an Akan ‘day’ name for a male person born on Sunday, he 183
may have been born in Ghana, but it is also possible that he received this name in Suriname. 184
Around 1727, he started working as a scout for the colonial authorities and headed 185
expeditions against Maroon villages, descendants of escaped slaves who settled in the 186
interior forests. For his services, Kwasi was freed in 1755, though he was already honored as 187
“faithful to the Whites” in 1730, which gave him the nickname ‘Kwasimukamba’ (Kwasi the 188
white man) among the Maroons (Dragtenstein, 2004; Price, 1979). As an outspoken and 189
controversial person, he was both feared and renowned as a traditional healer, illegal trader 190
in enslaved Amerindians, and as a sorcerer, renowned for his protective amulets. Kwasi 191
served as the official healer of the governor of the Dutch colony and eventually became a 192
plantation owner himself (Dragtenstein, 2004; Price, 1979). Kwasi became famous after he 193
(supposedly) discovered the healing properties of Q. amara around 1730, when he had 194
frequent interactions with Amerindians, and his successful treatments of both enslaved 195
Africans and Europeans suffering from fever (Dragtenstein, 2004). 196
On 20 December 1755, the Swedish biologist Daniel Rolander, sent to Suriname to collect 197
specimens for the famous botanist Carolus Linnaeus, wrote in his diary: ‘‘A man skilled in 198
plants and their uses is viewed in this region as both enviable and dangerous. I myself 199
conversed with Quassi on a couple of occasions. He was quite guarded with his wisdom; he 200
said he would reveal nothing until receiving a considerable sum of money” (Rolander, 2008). 201
Although the wood of Q. amara, known as lignum quassiae, was already exported in large 202
quantities to Europe in 1755, Rolander was the first to provide a (Latin) description of the 203
flowering and fruiting parts of this plant, but Linnaeus never got hold of his specimens (van 204
Andel et al., 2012a). Kwasi is said to have sold his secret knowledge regarding Q. amara to 205
Carl Gustav Dahlberg, the Swedish owner of the plantation Nieuw Timotibo and Rolander’s 206
mentor (Davis, 2016). In 1761, Dahlberg brought a specimen of the plant to Linnaeus, who 207
dedicated its scientific name to Kwasi and the bitter taste of the wood (Blom, 1763; Linné, 208
1762). 209
The Belgian physician Phillipe Fermin challenged Kwasi’s discovery of Q. amara: “This does 210
not seem quite probable to me, since it had already been known, for nearly forty years, to 211
almost all the inhabitants of Surinam, who made use of the flowers brought by this tree, and 212
who regarded them as very stomachic; and this when I arrived in 1754" (Fermin, 1769). 213
Fermin’s remarks suggest that around 1714 this remedy for stomach pain was already well-214
known. However, Q. amara is absent from the first herbarium collections from Suriname, 215
dating from 1687 and representing 48 useful species (van Andel et al., 2012c). The species 216
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neither appears in the plant drawings of Maria Sybilla Merian, who portrayed 60, mostly 217
cultivated plant species in Suriname around 1699 (Merian, 1705). Kwasi was invited to travel 218
to the Dutch Republic in 1776 and fêted by Willem V, Prince of Orange (Price, 1979; Voeks 219
and Greene, 2018), a rather uncommon treatment for a former enslaved African at that 220
time. During this visit, he complained that Dahlberg had earned a fortune with the export of 221
the bitter wood, for which he had received very little (Dragtenstein, 2004). 222
3.3 Quassia amara’s long journey southeast 223
According to Woodson et al. (1973), Grandtner and Chevrette (2013) and herbarium 224
specimens in GBIF (2020a), the present distribution of Quassia amara ranges from 225
Michoacan (Mexico) to the northeast Brazil, with the highest density between northern 226
Panama and southern Nicaragua (Figure 1). This distribution may reflect a collection bias, 227
and GBIF data include herbarium specimens from cultivated individuals. However, despite 228
the assumption of Woodson et al. (1973) that its exact origin is impossible to determine 229
because of its frequent cultivation in the last centuries, Q. amara is unambiguously 230
considered as native to tropical America (Figure 1). It is observed wild in Panama in 231
‘monsoon’ forest and in evergreen seasonal forests up to 1000 m (D’Arcy, 1987; Woodson et 232
al., 1973), and in evergreen moist forest in Costa Rica (Díaz et al., 2006). According to Pulle 233
et al. (1979) and the herbarium labels consulted, the species has an affinity for wetlands, 234
flooded forests or riverbanks. It is nowadays commonly found naturalized along rivers in 235
Suriname (F-Maguire 23851, van Andel personal observation) and on former plantations in 236
French Guiana (CAY-Feuillet 2859 and CAY-Houël 1). It is also frequently cultivated in 237
gardens in the coastal areas of Guyana, Suriname and French Guiana. The map of Q. amara 238
distribution shows a relatively standard pattern, with a high density of specimens collected 239
at a central position, indicating the putative center of origin, and a decreasing occurrence 240
towards the periphery of its range. 241
242
243
Figure 1. Approximate range of Q. amara in Central and South America from herbarium collection 244 localities recorded in GBIF. 245
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246
Q. amara supposedly arrived in French Guiana in 1772 (Barbier, 1824), where it gained its 247
local name “quinquina de Cayenne”, referring to the Andean quinine bark (Cinchona spp., 248
Rubiaceae), subsequently leading to several confusions. The first herbarium specimen from 249
Cayenne was collected in 1788 (P-Stoupy s.n.), and the second in 1792 (P-Leblond s.n.). The 250
fact that Q. amara does not appear in the inventories of Barrère (1743, 1749) or Aublet 251
(1775) supports this time of introduction, despite the difficulty to correspond the 18th 252
century names and identifications to modern ones if no specimens exist. In 1896, the species 253
was collected growing spontaneously close to Cayenne (P-Soubirou s.n.). Circulation of 254
knowledge in French Guiana mainly occurs (today and in the past) within the coastal area 255
and along the main rivers, such as the Maroni and the Oyapock (Tareau, 2019; Tareau et al., 256
2019). The fact that the species is unknown among the Wayãpi, Wayana and Teko 257
indigenous groups currently living in the remote South of French Guiana, whose only links to 258
the coast are these rivers, also support this coastal way of introduction (Grenand et al., 259
2004; Odonne & Davy, unpublished results). 260
In Colombia, Q. amara was first collected along the Rio Viejo (close to the Magdalena River) 261
in 1801 (P-Bonpland 1536), the voucher being dated after Sprague (1926). Although we were 262
unable to determine the context of this collection, it’s local name (cruzette) is similar to what 263
it is called two centuries later. The Spanish physician and botanist José Celestino Mutis 264
imported the plant to Colombia around 1775 (Mutis, 1778), in spite of its probable natural 265
occurrence in the northeastern parts of the country, which was probably unknown to Mutis 266
as one of the first botanists exploring the country. At the Missouri herbarium (MO), there 267
are eight Colombian specimens (mostly from the northwest of the country), of which three 268
were collected from the wild in disturbed forest. 269
Although Berry (2005) ambiguously considers Q. amara as “native to the Venezuelan 270
Guayana, the Guianas and Northern Brazil”, based on only one collection from a wild 271
individual in secondary forest, the absence of specimens from Eastern Colombia and 272
Venezuela is striking (Figure 1). As far as we know, the plant is very little used in Venezuela 273
today, as it is absent from medicinal market surveys held in Caracas (Giraldo et al., 2009) and 274
not cited among the medicinal species recorded in the Trujillo state (Bermúdez and 275
Velázquez, 2002; Carrillo-Rosario and Moreno, 2006), nor in the Aragua state (Jaramillo et 276
al., 2014). The only Venezuelan specimen with a local name (MO-Trujillo 21360) was 277
collected in 1988 in a fairly isolated community of Carib-speaking Akawaio Amerindians, who 278
live close to the border with Guyana. 279
The plant is notably absent from the earliest extensive natural history description from 280
Brazil, the Historia Naturalis Brasiliae (Alcantara-Rodriguez et al., 2019). It is also lacking in 281
the early 19th century floristic inventories by C.F.P. von Martius and A. de Saint-Hilaire, and 282
neither listed by European botanists in Brazil in the 19th century (Brandão et al., 2012, 2008; 283
Breitbach et al., 2013). Its first collection was made in 1841 and is labelled “woods 284
maranham” (K-Gardner 5982), suggesting a possible collection in a forest from the 285
Maranhão (called Maranham in the 19th century). The harbor of Saõ Luis (Maranhão’s 286
capital) was an important colonial trade hub, and might have been the entry point of this 287
species. According to the previous references, we consider its native status in Brazil very 288
unlikely and this specimen might have been collected in a secondary forest, as observed in 289
Suriname or French Guiana. The second most ancient specimen was collected in 1873, 290
cultivated in Barreiras (lower Amazon) (K-Traill 102), then in 1918 in the Rio de Janeiro 291
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botanical garden (F-Whitford 21), while the few other early 20th century specimens (e.g. F-292
Dahlgreen 49) are from the Para state, which is close to French Guiana. 293
Q. amara does not appear in the comprehensive list by Rios et al. (2007) of 1191 useful 294
species in Ecuador, based on specimens in the Herbarium da Pontificia Universidad Católica 295
del Ecuador (QCA), and likewise in Peru, as it is not cited in the Peruvian Checklist from the 296
Missouri Botanical Garden website (2020). The species was collected in the wild in Nicaragua 297
in 1853 (P-Wright s.n.), and several wild specimens were found in the southeastern part of 298
Mexico, with the first record from the state of Colima in 1891 (US-Palmer 1338). 299
Regarding the Caribbean, the first collections come from the French West Indies, probably 300
introduced there from French Guiana. In 1820, a sample was collected from a cultivated 301
plant in Martinique known as “quinquina du pays” and used against fevers, and said to come 302
from French Guiana (P-Plée s.n.), while the first specimen collected in Guadeloupe dates 303
from 1808 and was cultivated as well (P-L’Herminier s.n.). In 1895, the plant was present in 304
Trinidad (L-Hart 4492), and maybe even earlier, as a specimen from the A.L. de Jussieu 305
herbarium (P-Riedlé s.n.) was thought to be collected on this island around 1798, as well as a 306
specimen from 1828 (P-Brongniart s.n.). In the US Virgin Islands, one specimen supposedly 307
comes “ad habitat naturalis” from Saint-Thomas in 1887 (P-Eggers 155), but the specimen 308
collected in Saint-Croix in 1896 was cultivated (P-Ricksecker s.n.). 309
Just like the fate of many other useful plants after the Columbian encounter (Boumediene, 310
2016; Voeks and Greene, 2018), Q. amara experienced a rapid fame in colonial territories in 311
the Old World, and spread as far as Saigon (sent from Suriname) in 1874 (P-Pierre 3864; P-312
Poli s.n.), Bangkok in 1899 (US-Zimmermann 67), Manila and Batavia in 1903 (US-Merril 313
3434; L-Backer 82), Guinea in 1905 (P-Chevalier 13043), Ambon (Indonesia) in 1913 (US-314
Robinson 1765), and the Congo in 1922 (P-Goossens 3076). It arrived in Hawaii in 1931 from 315
Singapore (L-Shearard et al. 27), and in Micronesia in 1949 (US-Glassman 2562). 316
3.4 Geopolitics of bitterness… 317
The typical distribution of Q. amara and the fact it has been rarely found outside gardens 318
and former plantation areas in the Guiana shield, suggest that it is probably not native in the 319
southeastern part of its range. The species probably originates from the area between 320
Nicaragua, Costa Rica and Panama (Woodson et al., 1973), and has been introduced 321
northwest to Mexico and southeast by travelers towards northern Brazil between the early 322
18th and the 19th century. 323
The absence from Southern Colombia and Ecuador is striking: a species that can thrive in 324
such a wide area and survives in secondary forests would have been able to spread further 325
southeast, at least following human introduction. Ecuador was part of the Virreinato del Perú 326
until 1720, whereupon it became part of the Virreinato de Nueva Granada (currently 327
Colombia). This area (southern Colombia, Ecuador and northern Peru) is the natural home of 328
the quina trees (Cinchona spp., Rubiaceae), which were at that time the main source of 329
bitter antimalarials and the most sought-after medicinal plant species. Mutis, who organized 330
in 1783 the “Real Expedición Botánica del Nuevo Reino de Granada” (Jaramillo-Arango, 331
1952), wrote an influential text on the study of quina bark (Mutis and de Gregorio, 1828). 332
However, an unpublished manuscript by Mutis, dating from 1778 (Figure 2), is of particular 333
interest for our story: the “informe presentado al virrey sobre muestras de quina de la 334
Guayana”. In this text, Mutis, after having analyzed some samples from (most probably) Q. 335
amara “affirms [...] that the so-called quina de la Guyana not only lacks all the characters of 336
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the real quina or Cinchona officinal, but not even [...] enters as a subordinate species of this 337
genus, in the concept of the botanists” (Mutis, 1778) 338
339
340
Figure 2. Excerpt of Mutis (1778) stating that “in repeated trials as I have practiced, will be able to 341 conclude that whatever the bark of Guiana is, it is not Quina”. 342
343
This short text, indirectly speaking of the increasing fame of Q. amara, helps us to grasp one 344
of the great stakes of that time. Cinchona bark was at the very core of intense politico-345
economical battles and finding alternative sources (or other species with similar properties) 346
was essential for the colonial nations that competed with Spain (Crawford, 2016). This 347
aspect is confirmed by Leblond (1789), when he states “I set out for the Oyapock River with 348
much food, trade goods, and twenty-four men with the intention of reaching the sources of 349
that river […] to search for [Cinchona], […] but there was none of that.” 350
The spread of Q. amara far from its natural range may be related to religious discords, 351
notably between orders. Around the mid-18th century, the Jesuits were famous for their 352
interest in South-American materia medica (Boumediene, 2020), natural substances with 353
potential value for European pharmacies (Anagnostou, 2005). While seeking these medicinal 354
plants, they published some interesting descriptions of natural history. The “Orinoco 355
ilustrado y defendido” (Gumilla, 1745) is one of the important works on uses and customs in 356
the area covered by Jesuit’s missions, notably along the Orinoco River. The only very bitter 357
plant described in this book was identified as Crotalaria stipularia Desv. by del Rey Fajardo 358
(2017), from which we may interpret that they were unfamiliar with Q. amara. However, the 359
eventuality that the Jesuits would have spread this yet undescribed plant eastward is not 360
impossible, although it is not the most likely hypothesis. The Orinoco was roughly the 361
eastern limit of the Jesuit settlements (Tarble and Scaramelli, 2004) and can thus be 362
considered as a cultural biogeographical limit for the use of Q. amara, which is more 363
frequent east of this river in the Guiana shield. Considering that the Jesuits were ruling the 364
trade in Cinchona, widely known by the name of Jesuit’s bark (Boumediene, 2020), we 365
speculate that the appearance of Q. amara in the few territories where the Jesuits were 366
absent could be related to some kind of disinterest from this order over the concurrent 367
species of Cinchona. Even if Q. amara was already famous in Suriname at this time, the 368
Jesuit’s expulsion from Spanish territories in 1767 may have contributed to free its way to 369
Europe and other South American colonies. Jesuit’s networks and the commercial 370
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exploitation of Cinchona bark likely explains the absence of Q. amara from southern 371
Colombia to Peru, while the influence of other religious and political organizations could be a 372
possible explanation of its ‘sudden appearance’ in a Dutch colony. 373
3.5 Quassia amara’s linguistic legacy 374
The greater the variety of local names for a plant species in a specific region, the higher the 375
probability that plant is native from that area (Brown et al., 2013a; Shepard Jr and Ramirez, 376
2011; Westengen et al., 2014). Our review of the vernacular names of Q. amara (Table 1) 377
confirms that the species is native to Central America, given its higher number of 378
(indigenous) names compared to the Guiana Shield and Brazil, where most names are 379
derived from post-Colombian names, such as kwasi or quinine/quina or Cayenne. Moreover, 380
in Brazil nearly 25 species of febrifugal plants share the name of quina (Pio Corrêa, 1926; 381
1969; 1978). It is interesting that the Kali'na and Palikur indigenous peoples in French Guiana 382
name this species kuwasi or kwasβan, names also derivated from kwasi. 383
384
[Table 1]: Local names of Q. amara in the Neotropics mention in literature and on herbarium labels. 385
Country
(North to South)
Cultural
group
Language Name References
Mexico ? Spanish quina US-Hinton 15865 Mexico ? Spanish quina roja MO-Hinton 16221
Mexico ? Spanish cuasia MO-Breen 1077
Nicaragua ? Spanish hombre grande F-Proctor 26929
Nicaragua ? ? wama baka MO-Grijalva et al 6044
Nicaragua Garifuna Arawak wéwe gífi Coe and Anderson, 1996
Nicaragua Rama Chibchan (name in English Creole)
bitta wood Coe, 2008
Nicaragua Ulwa (Sumu) Misumalpan batakka dî basta Coe and Anderson, 1999
Costa Rica ? ? bakonki Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? Spanish cuasia Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? Spanish hombre grande MO-Mertinez 280; Pittier, 1908
Costa Rica ? ? guabo Pittier, 1908
Costa Rica ? Spanish hombrón Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? ? kläklö Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? ? pito kicha Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? ? quini Fournier & García 1998 in Díaz et al. (2006)
Costa Rica ? ? webblakló Fournier & García 1998 in Díaz et al. (2006)
Panama ? Spanish crucete MO-Sharp 1962; Mo-Standley 27417
Panama ? Spanish guabo amargo MO-Howell 8
Panama ? Spanish guavita amarga MO-Sharp 1962
Panama ? Spanish guavito MO-Kluge 36, Sexton 114
Panama Chocó Chocó (name in Spanish)
hombre grande MO-Duke & Kirkbride 14074; Duke, 1970
Panama ? Spanish juavita amarga MO-Smith et al. 3280
Panama Kuna (Bayano)
Chibchan udut pulu MO-Duke 14471
Colombia ? Spanish contra-cruceta US-Romero Castaneda 604
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386
Amerindians in Suriname, and particularly Carib-speaking groups, are the only people in the 387
Guiana shield to have their own specific vernacular names that are not derived from kwasi 388
(Table 1). The single Venezuelan unique vernacular name is also from Carib-speaking 389
Amerindians. The diversity of names in Carib languages for Q. amara may be a sign of a long-390
standing associated knowledge of this group, which originally comes from northern South 391
Colombia ? Spanish cruceto morado F-Dugand et al. 367
Colombia ? French (name derived from Spanish)
cruzette P-Bonpland 1536
Colombia ? Spanish cuásia US-Dugand et al. 4105
Venezuela ? Spanish bejuco barbasco Berry, 2005
Venezuela Akawaio Carib maipa MO-Trujillo 21360
Venezuela ? Spanish salsa hueca F-Steyermark 60962
Venezuela ? Spanish palo isidoro F-Aristeguieta 2110, US-Pittier 10970
Venezuela ? Spanish mamoncillo F-Steyermark 86595
Guyana All Guyanese Creole quashi bitter; quassy bitters
F-Dahlgreen et al. sn., van Andel, 2000
Guyana All Guyanese Creole bitter wood Lindeman et al., 1963
Suriname All Surinamese
Dutch Creole kwasibita van Andel and Ruysschaert, 2014
Suriname All Surinamese
Dutch bitterhout van Andel and Ruysschaert, 2014
Suriname Wayana Carib peunpe van Andel and Ruysschaert, 2014
Suriname Arawaks Arawak kareudan van Andel and Ruysschaert, 2014
Suriname Kali’na Carib apekyi/apekïi, (a) pekeï
van’t Klooster et al., 2003
Suriname Kali’na Carib këripu/k’eripu van’t Klooster et al., 2003
French Guiana
Kali’na Carib kuwasi Grenand pers. com.
French Guiana
Palikur Amerindian kwasβan Grenand et al., 2004
French Guiana
Créoles French Creole kwachi/couachi Grenand et al., 2004 and P-Benoist s.n.
Brazil ? Portuguese amargo Lorenzi and Matos, 2008
Brazil ? Portuguese quássia (amarga)
Lorenzi and Matos, 2008
Brazil ? Portuguese quássia-de-caiena
Lorenzi and Matos, 2008
Brazil ? Portuguese quina Lorenzi and Matos, 2008 & F-Archer 7543
Brazil ? Portuguese quinarana Lorenzi and Matos, 2008
Brazil ? Portuguese pau-quássia Lorenzi and Matos, 2008
Brazil ? Portuguese murubá, murupá
Pio Corrêa, 1978
Martinique ? French Creole quinquina de Cayenne
US-Stehlé 5872
Martinique ? French Creole quina pays P-Bélanger s.n.
Guadeloupe ? French Creole quinine caraïbe P-Stehlé 8038
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America (Antczak et al., 2017; Davis and Goodwin, 1990). However, the limited number of 392
specimens, indigenous names and uses recorded in Venezuela (from where the Carib groups 393
originate) makes the hypothesis of a strictly Carib-based discovery and diffusion unlikely. 394
Some names, such as peunpe, from the Carib-speaking Wayana, also refer to other species, 395
notably the other bitter Simaroubaceae tree Simarouba amara Aubl. 396
A possible explanatory hypothesis would be that the plant was spread by European colonists 397
during the early 18th century, without leaving written traces, from Central America to 398
Northern South America (current Venezuela), from where Carib-speaking groups have 399
transported it through the Guiana shield down to Suriname, where Kwasi made it famous. 400
Then, from Suriname, the species was subsequently brought to French Guiana and Brazil by 401
European, Afro-American and/or and Amerindian groups. The unfamiliarity with this species 402
among the southernmost indigenous groups of French Guiana (Grenand et al., 2004; Odonne 403
and Davy, unpublished results), combined with the lack of herbarium specimens outside the 404
coastal area, support the latter part of this hypothesis. Still, the possibility of a transatlantic 405
knowledge transfer remains to be analyzed. 406
3.6 Medicinal uses of Q. amara in the Neotropics 407
To complete the diffusion and subsequent biogeography of Q. amara, we reviewed its uses 408
in South and Central America. Throughout the Neotropics, it is used for a wide range of 409
affections such as fever, snakebites (US-de Bruijn 1042; US-Romero Castaneda 604), 410
diarrhea, urinary infections, diabetes and stomach pain (Coe and Anderson, 1996; Girón et 411
al., 1991; House et al., 1995). According to Pittier (1908), the plant was one of the most 412
commonly used remedies by the native communities in Costa Rica, where it was employed 413
to heal cold sores and as an aperitive, and Duke (2008) also reported its use against 414
hangovers. It was found as a remedy against fevers in Panama in 1924 (US-Kluge 36), and 415
against malaria in Nicaragua in 1946 by natives of the Isla Zapatero and in 1990 in the Rio 416
San Juan area (MO-Grijalva 1946, MO-Salick 7832). In Mexico, it was found in a herbalist 417
store in the Monterrey area and sold to stimulate drainage of the gall bladder (MO-Breen 418
1077). Few uses were registered in Venezuela, but it was used in 1944 in Tumeremo (Bolivar) 419
to treat fevers and hemorrhages (F-Steyermark 60962) and in 1954 in Perija (Zulia) against 420
“calenturas”, probably also referring to fever (F-Aristeguieta et al. 2110). 421
In Suriname, it is widely found on the markets of Paramaribo and Albina (van Andel et al., 422
2007), where carved wooden cups called kwasi bita bekers are sold to be filled with water or 423
rum (Figure 3). The content is drunk as a bitter tonic, stomachic or remedy against fevers 424
(Odonne et al., 2007). The wood is also a frequent ingredient of bitter aphrodisiac bottles 425
(van Andel et al., 2012b) (Figure 3). These cups allow the Saamaka Maroons to perpetuate 426
their great ability in wood carving, an undeniable African heritage (Price et al., 2005). 427
428
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429 Figure 3. Carved medicinal cup made from Q. amara wood and an aphrodisiac bottle containing 430 pieces of wood from the same species. Picture: G. Odonne. 431
432
This practice of shaping goblets was also current in Guyana in 1934, as retrieved from a 433
specimen label (US-Archer 2615), stating that bitter cups shaped like wine glasses were filled 434
with cane juice distillate and the extract used as a substitute for quinine, which was 435
confirmed by another specimen label from 1955 (US-Little 16700). 436
In French Guiana, the wood of Q. amara was once thought to be mainly a Creole remedy 437
(Grenand et al., 2004), but it recently emerged as the fifth most cited medicinal species, 438
among 12 of the 16 cultural groups interviewed by Tareau (2019). The bitter wood was even 439
cited by recent Haitian migrants in Cayenne, showing the adaptation of these cultures to the 440
medicinal flora of their new host country. Q. amara was also the most cited species in Saint-441
Georges de l’Oyapock, located at the border between French Guiana and Brazil, as a general 442
medicinal plant (Tareau et al., 2019). 443
In the upper Amazonia, the plant is infused or macerated as a febrifuge (Castner et al., 444
1998), used to treat hepatitis in Peru (Brack, 1999) and to combat malaria, vesicular colic, 445
intestinal gases and dyspepsia in Brazil (Gupta, 1995; Lorenzi and Matos, 2008). In brief, Q. 446
amara has two main domains of uses: gastro-intestinal complaints and fevers. 447
3.7 Colonial and industrial uses (and the rise of taxonomical confusion) 448
Soon acclaimed in the 19th century European medical treaties (Gänger, 2015; Heckel, 1897; 449
Mérat & de Lens, 1837; Reclu, 1889; Roques, 1837), Q. amara became popular in Europe for 450
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diverse medicinal applications. It also was employed as a bitter ingredient in beer (Heckel, 451
1897), which might explain its confusing names bois d’absynthe [absynth wood] and bois de 452
frêne [ash wood] on a specimen’s label from 1894 collected on Guadeloupe (US-Duss 2977). 453
In 1869, Suriname exported 245 tons of Q. amara wood to Europe (Price, 1983). Quassia is 454
cited in the 1820 version of the US Pharmacopeia (Hershenson, 1964), but with the species 455
name Q. excelsa, which may refer to another Simaroubaceae (Picrasma excelsa Planch.) with 456
which Q. amara was often mistaken or adulterated in the 19th century (Ocampo and Mora, 457
2011). This confusion is also observed in the Brazilian Pharmacopeia from 1926 (Brandão et 458
al., 2013), as well as in products sold in other places of the world (Figure 4) and 459
acknowledged by Duke (2008). 460
461
462
Figure 4. 19th century box of “Quassia” chips with two different scientific names (© The Field 463 Museum of Natural History - Botany Department - Catalog Number: 1970701. CC BY-NC). 464
465
Around the middle of the 19th century (Crookes, 1860), “tonic cups”, “tonic goblets” or 466
“Quassia cups” where sold against both gastro-intestinal complaints and fevers in drugstores 467
in the US, Canada and UK (Figure 5). However, defining the species used for each of these 468
wooden cups is difficult. They are unlikely to be all made from Q. amara wood, as they differ 469
in size, color and wood structure (Odonne et al. 2007). One of the most challenging issues in 470
historical (ethno-)pharmacology is to ensure the correct identification of plants in plant-471
based remedies. The world history of plant trade and pharmacy is a story of drug 472
adulteration and frauds, associated to the carelessness of some plant providers. Many 473
mismatches between names also originate from the use of different languages. Possibly, the 474
current local uses of Q. amara have been influenced by these 19th/20th century uses, in a 475
kind of retroactive knowledge hybridization, although this is difficult to prove. 476
477
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478
479
Figure 5. Early 20th century tonic cup, probably from Picrasma excelsa (© The Field Museum of 480 Natural History - Botany Department - Catalog Number: 271505. CC BY-NC). 481
482
3.8 Related species in the Neotropics 483
Several other species in the Simaroubaceae family are used as medicinal plants in tropical 484
America, occasionally sharing folk names, uses, morphological or chemical properties, in 485
what Bye et al. (1995) call a “medicinal plant complex”. The most salient species, due to its 486
frequent confusion with Q. amara for centuries, is Picrasma excelsa (Jamaican quassia, 487
Figures 4 and 5), which is still a popular medicinal plant in Jamaica today (Ocampo and Mora, 488
2011; Picking et al., 2015). In the Guiana shield, Simarouba amara Aubl. is a tree up to 40 m, 489
with similar white and bitter wood that is used against malaria in French Guiana (Gentry, 490
1996; Grenand et al., 2004). Picrolemma sprucei Hook.f. is a shrub or small tree (1-10 m) 491
with orange fruits (Berry, 2005), of which the wood is used against diabetes and fever in 492
Amazonia (Grenand et al., 2004), and sometimes replaces Q. amara in French Guiana. 493
Picramnia pentandra Sw. is a shrub or small tree (1.5-8 m) named bwa mondong ( “wood of 494
the African Mandingo cultural group”) in Martinique, where it grows in xerophytic forests on 495
basaltic substrate (Fournet, 1978). This name explicitly refers to its use by enslaved Africans, 496
and both in Martinique (Nossin, 2019) and Cuba (Cabrera, 1993), this species is a popular 497
ritual plant among people of African descent. In Cuba, it is also used against fevers (Cabrera, 498
1993). 499
3.9 Out of Africa? 500
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Several Neotropical plant species have been named by enslaved Africans after botanically 501
related species in Africa (van Andel et al., 2014). This recognition of New World medicinal 502
species by analogy with related Old World species is a perfect illustration of a cultural 503
hybridization process (Stockhammer, 2012), and it could be a plausible scenario that 504
enslaved Africans recognized Q. amara in Suriname because they knew the properties of a 505
related African species: Quassia africana. This would be concurrent with what Chevalier and 506
Russell (1936) wrote on this topic with an intuition whose accuracy was perhaps 507
underestimated at the time: “Isn't it a noteworthy fact that the use of the bitter wood 508
provided by the Simaroubaceae among the primitive peoples [sic] of both Tropical America 509
and Black Africa?” The American Q. amara was only recently introduced in Africa, and nearly 510
all the specimens are cultivated (e.g. from Ivory Coast: L-Leeuwenberg 3829) or clearly 511
stated as imported, such as the Ghanaian specimen from 1972 that was introduced from 512
Brazil (L-Cudjoe 588). 513
Q. africana is mainly distributed in Central Africa: the Democratic Republic of Congo, the 514
Republic of Congo, Gabon, Cameroon and Equatorial Guinea (GBIF, 2020b). There, it is used 515
as an antimalarial (Mbatchi et al., 2006), febrifuge (Musuyu Muganza et al., 2012), and 516
against diarrhea, worms, stomach pains and as a tonic (Longanga Otshudi et al., 2000; Bajin 517
ba Ndob et al., 2016). This corresponds to current uses of Q. amara in the Guiana shield 518
(Grenand et al., 2004; Odonne et al., 2007; Tareau, 2019), which makes the hypothesis that 519
enslaved Africans (instead of Amerindians) discovered its medicinal uses more credible. 520
However, the most salient uses of Q. africana that emerges from herbarium voucher labels 521
are to treat lice (quoted 10 times), digestive disorders (7), worms (4), fever (3), poisoning (3), 522
sexually transmitted diseases (2), and other ailments (3). Given the diverse uses of Q. 523
africana and its occurrence in a region where many slave traders were purchasing their 524
captives (Warner-Lewis, 2003), it is surprising that none of the Central African names for Q. 525
africana were transferred to Q. amara in the Americas. The transfer from Africa to America 526
is eventuality possible explanation for gastrointestinal-related disorders only, which are in 527
both places the most frequently treated disorders with these species. Our hypothesis, 528
however, is nevertheless not supported by the absence of any local American names for Q. 529
amara (Table 1) to the hundred names reported for Q. africana (Appendix B). However, 530
most local names for Q. africana are from Central Africa and the Bantu linguistic family. As 531
Kwasi probably had an Akan-speaking (West African) origin, he or fellow Akan-speakers may 532
not have recognized this Central African species. 533
The discrepancy between the few early 20th century specimens mentioning African uses of 534
Quassia against fevers, and the many contemporary references that claim it, suggest a 535
possible cultural transfer from Q. amara to Q. africana, i.e. from America to Africa. Did the 536
colonial trade of Q. amara bitters stimulate the use of the botanically related African 537
species? The massive worldwide use of bitter Cinchona barks from the 17th to the 19th 538
century, and the subsequent use of its isolated bitter component quinine from the mid-19th 539
onwards as a treatment against malaria, may have shaped a post-colonial cognitive 540
relationship between malaria treatment and bitterness. Such a relation is observed every 541
day in field interviews when speaking about malaria and its herbal remedies, for which 542
bitterness is often sought after. The connection between bitterness and malaria may have a 543
historical base (Cosenza et al. 2013). 544
3.10 Perspectives for Cultural Biogeography 545
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“Cultural biogeography” is an interesting concept to reconstruct the dynamics of biocultural 546
interactions through space and time. In this particular case, relying on herbarium databases 547
helps to decipher the “evolution” of medicinal plant use, particularly for the last few 548
centuries, and to understand the ecological niches (primary or secondary forests) and 549
domestication status of plants (cultivated, escaped from gardens, etc.). Some major 550
limitations of herbarium labels, however, is the scarce ethnobotanical information provided, 551
the bias of the botanical collector and the fact that the recorded plant use is often based on 552
the knowledge of a single person. Historic specimens often come without detailed locality 553
data, local names or uses, but valuable information emerges when huge quantities of 554
specimens are reviewed. 555
A next step in tracing the details of the origin and spread of Q. amara would be to perform 556
DNA analyses on herbarium specimens, as genetic tools now allow to precise the ancient 557
geographical origin of useful species and trace pre- and postcolonial exchange networks and 558
other long-time movements of plants and people (Clement et al., 2017; Moreira et al., 2017; 559
Przelomska et al., 2020; Rossetto et al., 2017; Roullier et al., 2013b). The field of Historical 560
genomics (Brousseau et al., 2020) can decipher long-term migrations of plants, and indicate 561
domestication events or strong biocultural interactions (van Andel et al., 2016). Molecular 562
studies on herbarium samples would shed more light on the genetic distance between Q. 563
amara samples from the Guiana Shield and Central America, which can reveal historic 564
migration routes. In our case, DNA barcoding could also help to identify the wooden objects 565
in museums labelled as “Quassia”. 566
Linguistics, and particularly paleobiolinguistics, is a useful tool to study cultural 567
biogeography, but so far is only applied to trace back the origins of food crops (Brown et al., 568
2013a, 2013b; Westengen et al., 2014). Interdisciplinary studies are needed more than ever 569
to unravel complex itineraries of plants and people (Perrier et al., 2011), as represented in 570
the archaeobotany of both physical remains and pictorial representations (Hather, 2010), 571
and in the study of ancient DNA (Brown et al., 2015; Przelomska et al., 2020) or in 572
anthracology (Bodin et al., 2020). 573
4. Conclusion 574
Our research points out that Quassia amara’s uses are difficult to attribute to a particular 575
cultural group: the species is widely distributed in Central and South America, where it is 576
popular among many indigenous and non-indigenous groups. Based on our results, we 577
conclude that Q. amara was spread during the early 18th century due to political and 578
economic reasons. This spread is possibly the result of simultaneous actions by European 579
colonists from Central America to northern South America and by Carib-speaking 580
Amerindians from northern South America to Suriname, where Kwasi increased its fame as 581
an antimalarial around 1730. From there, it spread southeast towards French Guiana (in 582
1772) and Brazil in the 19th century (maybe via São Luis or Belem, the main harbors on the 583
North), and in the beginning of the 19th century, to the rest of the world. The presence of 584
Cinchona spp. in Southern Colombia, Ecuador and Peru may have caused little interest in Q. 585
amara, and prevented a successful introduction in that region. Today, the species is 586
cultivated throughout the tropics, both for domestic medicine and for pharmaceutical 587
applications. 588
The absence of African-derived local names in the Guiana shield suggests that enslaved 589
Africans in the Guiana shield were not familiar with Q. africana to have preserved its name 590
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and transferred this to Q. amara. The “discovery” of the medicinal properties of Q. amara in 591
Suriname seems, thus, to be a knowledge transfer to a West African by Carib-speaking 592
Amerindians, who may have imported the species for medicinal reasons. However, no 593
documented traces remain of this event. 594
Cultural biogeography had proven an interesting concept to reconstruct the dynamics of 595
biocultural interactions through space and time. Herbarium databases have proved useful to 596
decipher evolution of local knowledge in our research. Nevertheless, given the current 597
threats to ethnobotanical (and ethnoecological) knowledge, it is time to react with an 598
inclusive approach fostering the preservation of biocultural diversity more broadly. 599
Explaining the species, their uses, and their associated knowledge as global biocultural 600
heritage, and engaging efforts toward a better comprehension of their history, may help 601
avoid counterproductive political standstill. 602
5. Acknowledgements 603
We acknowledge Sophie Gonzalez, curator of the CAY herbarium, for her advices and for 604
providing us data and pictures of Cayenne’s samples, Pierre Grenand for his comments on 605
this paper, Maria Franco Trindade Medeiros for her advices on historical ethnobiology. Extra 606
thanks for those herbarium artists who transformed the painful examination of hundreds of 607
specimens into sometimes surprising laughs (Appendix A). 608
6. Funding 609
The authors were funded by an Investissement d’Avenir grant from the Agence Nationale de 610
la Recherche (CEBA: ANR-10-LABX-25-01), by the PEPS-ECOMOB and the Prime80-DiaspoREs 611
projects from the CNRS’ Environment and Ecology Institute (CNRS/INEE) and the Mission 612
pour l’Interdisciplinarité et les Initiatives Transverses (CNRS/MITI), and Naturalis Biodiversity 613
Center. 614
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Soelberg, J., Asase, A., Akwetey, G., Jäger, A.K., 2015. Historical versus contemporary medicinal plant 937 uses in Ghana. J. Ethnopharm. 160, 109–132. https://doi.org/10.1016/j.jep.2014.11.036 938
Soelberg, J., Davis, O., Jäger, A.K., 2016. Historical versus contemporary medicinal plant uses in the 939 US Virgin Islands. J. Ethnopharm. 192, 74–89. https://doi.org/10.1016/j.jep.2016.07.005 940
Sprague, T.A., 1926. Humboldt and Bonpland’s itinerary in Colombia. Bull. Misc. Inf. (Royal Botanic 941 Gardens, Kew) 1926, 23–30. https://doi.org/10.2307/4114319 942
Stedman, J.G., 1796. Narrative of a five years’ expedition, against the revolted negroes of Surinam, in 943 Guiana, on the wild coast of South America: From the year 1772, to 1777. Johnson & 944 Edwards, London. 945
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Tarble, K., Scaramelli, F., 2004. A brief but critical presence: The archaeology of a Jesuit mission in the 946 middle Orinoco (1730-47). Missionalia: South. Afr. J. Miss. Stud. 32, 419–444. 947
Tareau, M.-A., 2019. Les pharmacopées métissées de Guyane: ethnobotanique d’une phytothérapie 948 en mouvement. PhD thesis, Université de Guyane, Cayenne. 949
Tareau, M.-A., Dejouhanet, L., Palisse, M., Odonne, G., 2019. Circulations et échanges de plantes et 950 de savoirs phytomédicinaux sur la frontière Franco-Brésilienne. Rev. Franc. Santé Terr. 19. 951
Thomas, F., 2018. Patents and benefit sharing: What can we learn from the Quassia amara lawsuit? 952 What is the problem? in: The commons, plant breeding and agricultural research (Girard, F., 953 Frison, C.). Routledge, London, pp. 61–73. 954
van Andel, T., 2000. Non-timber forest products of the North-West District of Guyana, part I & II. 955 Utrecht University, Utrecht. 956
van Andel, T., 2016. The reinvention of household medicine by enslaved Africans in Suriname. Soc. 957 Hist. Med. 29, 676–694. https://doi.org/10.1093/shm/hkv014 958
van Andel, T., Behari-Ramdas, J., Havinga, R., Groenendijk, S., 2007. The medicinal plant trade in 959 Suriname. Ethnobot. Res. App. 5, 351. https://doi.org/10.17348/era.5.0.351-372 960
van Andel, T., Maas, P., Dobreff, J., 2012a. Ethnobotanical notes from Daniel Rolander’s diarium 961 Surinamicum (1754–1756): Are these plants still used in Suriname today? Taxon 61, 852–863. 962 https://doi.org/10.1002/tax.614010 963
van Andel, T., Meyer, R.S., Aflitos, S.A., Carney, J.A., Veltman, M.A., Copetti, D., Flowers, J.M., 964 Havinga, R.M., Maat, H., Purugganan, M.D., 2016. Tracing ancestor rice of Suriname Maroons 965 back to its African origin. Nat. Pl. 2, 16149. 966
van Andel, T., Mitchell, S., Volpato, G., Vandebroek, I., Swier, J., Ruysschaert, S., Rentería Jiménez, 967 C.A., Raes, N., 2012b. In search of the perfect aphrodisiac: Parallel use of bitter tonics in West 968 Africa and the Caribbean. J. Ethnopharm. 143, 840–850. 969 https://doi.org/10.1016/j.jep.2012.08.008 970
van Andel, T., Ruysschaert, S., 2014. Medicinale en rituele planten van Suriname. LM Publishers, 971 Paramaribo. 972
van Andel, T., van‘t Klooster, C.I., Quiroz, D., Towns, A.M., Ruysschaert, S., van den Berg, M., 2014. 973 Local plant names reveal that enslaved Africans recognized substantial parts of the New 974 World flora. Proc. Nat. Acad. Sci. 111, e5346–e5353. 975
van Andel, T., Veldman, S., Maas, P., Thijsse, G., Eurlings, M., 2012c. The forgotten Hermann 976 Herbarium: a 17th century collection of useful plants from Suriname. Taxon 61, 1296–1304. 977
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Country Date Voucher or reference Local name Ethnolinguistic group Use URL
DRC (Mobwasa) 1913 BR-de Giorgi 884 balebame https://www.botanicalcollections.be/#/en/specimen/BR0000019460130DRC (Mobwasa) 1913 BR-de Giorgi 660 balibamu against belly aches https://www.botanicalcollections.be/#/en/specimen/BR0000019460093DRC (Efomi) 1982 BR-Gorbatoff 136 balombe https://www.botanicalcollections.be/#/en/specimen/BR0000019460802DRC 1925 BR-Robyns 1037 betoma https://www.botanicalcollections.be/#/en/specimen/BR0000021300608DRC (Bonzefo) 1952 BR-Gorbatoff 232 bokota gombe https://www.botanicalcollections.be/#/en/specimen/BR0000019460819DRC 1958 BR-Robin 19 bokota https://www.botanicalcollections.be/#/en/specimen/BR0000021300592
DRC (Mobwasa) 1913 BR-de Giorgi 838 bolé fish poison https://www.botanicalcollections.be/#/en/specimen/BR0000019460123DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 bolome lokundu http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137
Zaïre (Botsima) 1991 L-Dhetchuvi 716 bomengea lokundu https://medialib.naturalis.nl/file/id/WAG.1525266/format/largeDRC (Botsima) 1991 BR-Dhetchuvi 716 bomengea anthelminthic https://www.botanicalcollections.be/#/en/specimen/BR0000019460222DRC (Bokondji) 1959 BR-de Wanckel 13 bomengea against lices and anthelminthic https://www.botanicalcollections.be/#/en/specimen/BR0000019460178
DRC 1955 BR-Évrard 1223 bondjelete ngwaka https://www.botanicalcollections.be/#/en/specimen/BR0000019460314DRC (Bikoro) 1920 BR-Goossens 2331 bosole bangala against fever https://www.botanicalcollections.be/#/en/specimen/BR0000019460604DRC (Monsole lac Tumba)1920 BR-Goossens 2364 bosolemote bangala against fever https://www.botanicalcollections.be/#/en/specimen/BR0000019460628DRC (Luki) 1947 BR-Donis 1459 bundu https://www.botanicalcollections.be/#/en/specimen/BR0000018127515DRC (Luki) 1946 BR-Toussaint 2052 bundu tsi kiombe against fevers https://www.botanicalcollections.be/#/en/specimen/BR0000018127584DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 bundu tsi mayumbe http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC (N'kakala) 1947 BR-Toussaint 2177 bundu tsi https://www.botanicalcollections.be/#/en/specimen/BR0000006875435DRC (Luki) 1953 BR-Wagemans 739 dinianza https://www.botanicalcollections.be/#/en/specimen/BR0000018127607DRC 1956 BR-Hombert 210 dinianza https://www.botanicalcollections.be/#/en/specimen/BR0000013623395s.l. s.d. BR-Nsimundele 116 dinianza https://www.botanicalcollections.be/#/en/specimen/BR0000006262006DRC (Luki) 1946 BR-Toussaint 2052 dinianza against fevers https://www.botanicalcollections.be/#/en/specimen/BR0000018127584DRC 1913 BR-Lemaire 164 dumbo dumbo against ticks https://www.botanicalcollections.be/#/en/specimen/BR0000019461069DRC (Dundusana) 1913 BR-de Giorgi 1035 ebamu against constipation https://www.botanicalcollections.be/#/en/specimen/BR0000019460147
DRC 1938 BR-Léontovitch 111 ebamwa https://www.botanicalcollections.be/#/en/specimen/BR0000021300028DRC (Yambata) 1913 BR-de Giorgi 1372 ebedema against lice https://www.botanicalcollections.be/#/en/specimen/BR0000019460154DRC 1913 BR-Montchal 123 ebetoma https://www.botanicalcollections.be/#/en/specimen/BR0000021300370
DRC 1938 BR-Léontovitch 111 egongo https://www.botanicalcollections.be/#/en/specimen/BR0000021300028DRC (Yambata) 1914 BR-Montchal 166 eliekema (?) against belly aches and to heal wounds https://www.botanicalcollections.be/#/en/specimen/BR0000021300387DRC (Bonzefo) 1952 BR-Gorbatoff 232 etio gombe https://www.botanicalcollections.be/#/en/specimen/BR0000019460819DRC (Menkete) 1933 BR-Dubois 69 ialayetemba poison https://www.botanicalcollections.be/#/en/specimen/BR0000019460253DRC (Amadilu) 1906 BR-Sapin s.n. iaraiotomba against lice https://www.botanicalcollections.be/#/en/specimen/BR0000019459578DRC 1906 BR-Sapin s.n. iasaiotumba against lice https://www.botanicalcollections.be/#/en/specimen/BR0000005051854DRC (Ilenge) 1921 BR-Goossens 2685 ielalotombe bangala https://www.botanicalcollections.be/#/en/specimen/BR0000019460666DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 ielatomba http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 ieloiotomba http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC 1931 BR-Dubois 39B ilele poison, antisyphilitic https://www.botanicalcollections.be/#/en/specimen/BR0000019460239Gabon 1936 Chevalier and Russell, 1936 isindu igala punuDRC (Patumbalu) 1952 BR-Tailfer 11 kangololo https://www.botanicalcollections.be/#/en/specimen/BR0000021300684DRC (Tumba) 1957 BR-Liben 3402 katsha tshiluba https://www.botanicalcollections.be/#/en/specimen/BR0000019459509DRC (N'kakala) 1947 BR-Toussaint 2177 kikema https://www.botanicalcollections.be/#/en/specimen/BR0000006875435DRC (Kivangu) 1948 BR-Donis 1752 kikema https://www.botanicalcollections.be/#/en/specimen/BR0000018127522DRC (Luki) 1946 BR-Toussaint 2052 kikema against fevers https://www.botanicalcollections.be/#/en/specimen/BR0000018127584DRC (Luki) 1947 BR-Donis 1615 kikema https://www.botanicalcollections.be/#/en/specimen/BR0000006261993DRC (Luki) 1948 BR-Donis 2218 kikembe https://www.botanicalcollections.be/#/en/specimen/BR0000013246662DRC (Luki) 1949 BR-Donis 2301 kinkema https://www.botanicalcollections.be/#/en/specimen/BR0000006262327DRC (Luki) 1949 BR-anonymous 2301 kinkema https://www.botanicalcollections.be/#/en/specimen/BR0000021300790DRC (Luki) 1948 BR-Donis 2218 kodia kodia https://www.botanicalcollections.be/#/en/specimen/BR0000013246662DRC (Yonga) 1960 BR-Jespersen 24 kongololo against cholic, purgative, anthelminthic https://www.botanicalcollections.be/#/en/specimen/BR0000019460888DRC (Mayombe) 1921 BR-Nannan 46 kumbi https://www.botanicalcollections.be/#/en/specimen/BR0000018127553DRC 1921 BR-Goossens 2601 lealotomba bangala https://www.botanicalcollections.be/#/en/specimen/BR0000019460659DRC 1921 BR-Goossens 2937 lelaiotomba bangula anthelminthic https://www.botanicalcollections.be/#/en/specimen/BR0000019460673DRC (Jambuya) 1921 BR-Goossens 2541 leloiotomba https://www.botanicalcollections.be/#/en/specimen/BR0000019460642Gabon 2012 WAG-Queiroz-Villareal 1398 lindughala punu antidiabetic https://medialib.naturalis.nl/file/id/WAG.1952061/format/largeDRC (Mobwasa) 1913 BR-de Giorgi 764 mambabala https://www.botanicalcollections.be/#/en/specimen/BR0000019460116DRC (Lukolela) 1959 BR-Toka 185 mbongeliamba pama https://www.botanicalcollections.be/#/en/specimen/BR0000021300714DRC (Kiobo) 1940 BR-Donis 107 mbundu tsi kiombe https://www.botanicalcollections.be/#/en/specimen/BR0000018127492DRC (Likimi) 1913 BR-de Giorgi 1487 modimbo-dimbo-bali against constipation and lice https://www.botanicalcollections.be/#/en/specimen/BR0000019460161DRC (Kakenge) 1937 BR-Gillardin 333 mufura mukumba lulua (lg. tshiluba kananga) https://www.botanicalcollections.be/#/en/specimen/BR0000020214821DRC (Kakenge) 1937 BR-Gillardin 315 mufwa matumba lulua (lg. tshiluba kananga) against bellyache https://www.botanicalcollections.be/#/en/specimen/BR0000020214814DRC (Jambuya) 1921 BR-Goossens 1656 n'gelango bangala https://www.botanicalcollections.be/#/en/specimen/BR0000019460598Gabon 2008 BR-Ngombe Mamadou 273 ngounie https://www.botanicalcollections.be/#/en/specimen/BR0000018127164DRC (Mukumari) 1939 BR-Gillardin 574 okenzu kiketela https://www.botanicalcollections.be/#/en/specimen/BR0000019460567DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 okenzu kitelela http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC 1913 BR-Elskens s.n. oloko https://www.botanicalcollections.be/#/en/specimen/BR0000019460291Gabon 2012 L-Queiroz-Villareal 1398 oubara massango antidiabetic https://medialib.naturalis.nl/file/id/WAG.1952061/format/largehttps://medialib.naturalis.nl/file/id/WAG.1952061/format/largeCameroon 2002 ovuin miondo (miondo ?) http://specimens.kew.org/herbarium/K000338356DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 samba ? http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137Gabon 1936 Chevalier and Russell, 1936 simigala esbiraDRC (Mayumbe) 1904 BR-anonymous 546 vonda bikadi mayumbe https://www.botanicalcollections.be/#/en/specimen/BR0000021300790DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 vonda kadi mayumbe http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC (Luki) 1984 BR-Mbambi 29 vonda kadi https://www.botanicalcollections.be/#/en/specimen/BR0000018127546DRC (Luki) 1947 BR-Devred 3239 vonda kadi https://www.botanicalcollections.be/#/en/specimen/BR0000018127478DRC (Kivangu) 1948 BR-Donis 1752 vonda kadi https://www.botanicalcollections.be/#/en/specimen/BR0000018127522DRC (Luki) 1947 BR-Donis 1615 vonda kadi https://www.botanicalcollections.be/#/en/specimen/BR0000006261993DRC (Mayombe) 1921 BR-Nannan 305 vonda kati against blennoragy https://www.botanicalcollections.be/#/en/specimen/BR0000018127577DRC (Luki) 1946 BR-Toussaint 2052 vonda-kadi against fevers https://www.botanicalcollections.be/#/en/specimen/BR0000018127584DRC (Luki) 1946 BR-Toussaint 2052 vonda-kadi against fevers https://www.botanicalcollections.be/#/en/specimen/BR0000018127584DRC (Yangambi) 1938 BR-Louis 8340 weko turumbu (lg. lombo) https://www.botanicalcollections.be/#/en/specimen/BR0000013258054DRC (Yangambi) 1936 BR-Louis 1444 weko turumbu (lg. lombo) https://www.botanicalcollections.be/#/en/specimen/BR0000013258085DRC (Yangambi) 1952 BR-Troupin 2066 weko turumbu (lg. lombo) https://www.botanicalcollections.be/#/en/specimen/BR0000021300752DRC (Yangambi) 1935 BR-Louis 467 weko turumbu (lg. lombo) https://www.botanicalcollections.be/#/en/specimen/BR0000021300103DRC (Yangambi) 1952 BR-Troupin 2059 weko turumbu (lg. lombo) https://www.botanicalcollections.be/#/en/specimen/BR0000021300745DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 weko yangambi http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC 1938 BR-Gilbert 1048 weko https://www.botanicalcollections.be/#/en/specimen/BR0000019460444DRC (Yangambi) 1987 BR-Gilbert 22 weko https://www.botanicalcollections.be/#/en/specimen/BR0000019460420DRC 1947 BR-Gilbert 1056 weko https://www.botanicalcollections.be/#/en/specimen/BR0000019460451DRC (Bokolongo) 1958 BR-Evrard 3544 yaayotoma https://www.botanicalcollections.be/#/en/specimen/BR0000019460321
DRC (Bokote) 1943 BR-Hulstaert 1245 yala y'ôtòmba https://www.botanicalcollections.be/#/en/specimen/BR0000019460857DRC (Madibi) 1906 BR-Sapin s.n. yalalotumba against lices https://www.botanicalcollections.be/#/en/specimen/BR0000019459561DRC 1910 BR-Sapin s.n. yalaotumba against lices https://www.botanicalcollections.be/#/en/specimen/BR0000019459585DRC (Atene) 1907 BR-Sapin s.n. yalaotumba against lice https://www.botanicalcollections.be/#/en/specimen/BR0000019459561DRC 1980 BR-Dechamps 8078 yato yotomba mongo (lg. lomongo) https://www.botanicalcollections.be/#/en/specimen/BR0000019460062DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 yayotamda ? http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137DRC (Bolima) BR-Collart 61 yayotomba lomongo https://www.botanicalcollections.be/#/en/specimen/BR0000019460024DRC 1958 G. Gilbert (1958) Flore du Congo Belge et du Ruanda-Urundi: Simaroubaceae. vol. 7 yela y otomba ? http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137
DRC (Mondjo) 1946 BR-Léonard 494 yelayo tomba lokundu https://www.botanicalcollections.be/#/en/specimen/BR0000021300011DRC (Monsole lac Tumba)1920 BR-Goossens 2364 yelayotumba bangala against fever https://www.botanicalcollections.be/#/en/specimen/BR0000019460628DRC 1936 BR-Louis 2751 yeloye otumba ; etumba lokundu against lice https://www.botanicalcollections.be/#/en/specimen/BR0000019460390DRC (Yonga) 1960 BR-Jespersen 24 zlolo against cholic, purgative, anthelminthic https://www.botanicalcollections.be/#/en/specimen/BR0000019460888DRC 1913 BR-Nannan 1 against belly aches https://www.botanicalcollections.be/#/en/specimen/BR0000021300431
DRC (Atènes) 1907 BR-Sapin s.n. against lice https://www.botanicalcollections.be/specimen/BR0000019459592
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https://www.botanicalcollections.be/#/en/specimen/BR0000019460130https://www.botanicalcollections.be/#/en/specimen/BR0000019460093https://www.botanicalcollections.be/#/en/specimen/BR0000019460802https://www.botanicalcollections.be/#/en/specimen/BR0000021300608https://www.botanicalcollections.be/#/en/specimen/BR0000019460819https://www.botanicalcollections.be/#/en/specimen/BR0000021300592
https://www.botanicalcollections.be/#/en/specimen/BR0000019460123http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137
https://medialib.naturalis.nl/file/id/WAG.1525266/format/largehttps://www.botanicalcollections.be/#/en/specimen/BR0000019460222https://www.botanicalcollections.be/#/en/specimen/BR0000019460178
https://www.botanicalcollections.be/#/en/specimen/BR0000019460314https://www.botanicalcollections.be/#/en/specimen/BR0000019460604https://www.botanicalcollections.be/#/en/specimen/BR0000019460628https://www.botanicalcollections.be/#/en/specimen/BR0000018127515https://www.botanicalcollections.be/#/en/specimen/BR0000018127584http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000006875435https://www.botanicalcollections.be/#/en/specimen/BR0000018127607https://www.botanicalcollections.be/#/en/specimen/BR0000013623395https://www.botanicalcollections.be/#/en/specimen/BR0000006262006https://www.botanicalcollections.be/#/en/specimen/BR0000018127584https://www.botanicalcollections.be/#/en/specimen/BR0000019461069https://www.botanicalcollections.be/#/en/specimen/BR0000019460147
https://www.botanicalcollections.be/#/en/specimen/BR0000021300028https://www.botanicalcollections.be/#/en/specimen/BR0000019460154https://www.botanicalcollections.be/#/en/specimen/BR0000021300370
https://www.botanicalcollections.be/#/en/specimen/BR0000021300028https://www.botanicalcollections.be/#/en/specimen/BR0000021300387https://www.botanicalcollections.be/#/en/specimen/BR0000019460819https://www.botanicalcollections.be/#/en/specimen/BR0000019460253https://www.botanicalcollections.be/#/en/specimen/BR0000019459578https://www.botanicalcollections.be/#/en/specimen/BR0000005051854https://www.botanicalcollections.be/#/en/specimen/BR0000019460666http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000019460239
https://www.botanicalcollections.be/#/en/specimen/BR0000021300684https://www.botanicalcollections.be/#/en/specimen/BR0000019459509https://www.botanicalcollections.be/#/en/specimen/BR0000006875435https://www.botanicalcollections.be/#/en/specimen/BR0000018127522https://www.botanicalcollections.be/#/en/specimen/BR0000018127584https://www.botanicalcollections.be/#/en/specimen/BR0000006261993https://www.botanicalcollections.be/#/en/specimen/BR0000013246662https://www.botanicalcollections.be/#/en/specimen/BR0000006262327https://www.botanicalcollections.be/#/en/specimen/BR0000021300790https://www.botanicalcollections.be/#/en/specimen/BR0000013246662https://www.botanicalcollections.be/#/en/specimen/BR0000019460888https://www.botanicalcollections.be/#/en/specimen/BR0000018127553https://www.botanicalcollections.be/#/en/specimen/BR0000019460659https://www.botanicalcollections.be/#/en/specimen/BR0000019460673https://www.botanicalcollections.be/#/en/specimen/BR0000019460642https://medialib.naturalis.nl/file/id/WAG.1952061/format/largehttps://www.botanicalcollections.be/#/en/specimen/BR0000019460116https://www.botanicalcollections.be/#/en/specimen/BR0000021300714https://www.botanicalcollections.be/#/en/specimen/BR0000018127492https://www.botanicalcollections.be/#/en/specimen/BR0000019460161https://www.botanicalcollections.be/#/en/specimen/BR0000020214821https://www.botanicalcollections.be/#/en/specimen/BR0000020214814https://www.botanicalcollections.be/#/en/specimen/BR0000019460598https://www.botanicalcollections.be/#/en/specimen/BR0000018127164https://www.botanicalcollections.be/#/en/specimen/BR0000019460567http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000019460291
https://medialib.naturalis.nl/file/id/WAG.1952061/format/largehttp://specimens.kew.org/herbarium/K000338356http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137
https://www.botanicalcollections.be/#/en/specimen/BR0000021300790http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000018127546https://www.botanicalcollections.be/#/en/specimen/BR0000018127478https://www.botanicalcollections.be/#/en/specimen/BR0000018127522https://www.botanicalcollections.be/#/en/specimen/BR0000006261993https://www.botanicalcollections.be/#/en/specimen/BR0000018127577https://www.botanicalcollections.be/#/en/specimen/BR0000018127584https://www.botanicalcollections.be/#/en/specimen/BR0000018127584https://www.botanicalcollections.be/#/en/specimen/BR0000013258054https://www.botanicalcollections.be/#/en/specimen/BR0000013258085https://www.botanicalcollections.be/#/en/specimen/BR0000021300752https://www.botanicalcollections.be/#/en/specimen/BR0000021300103https://www.botanicalcollections.be/#/en/specimen/BR0000021300745http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000019460444https://www.botanicalcollections.be/#/en/specimen/BR0000019460420https://www.botanicalcollections.be/#/en/specimen/BR0000019460451https://www.botanicalcollections.be/#/en/specimen/BR0000019460321
https://www.botanicalcollections.be/#/en/specimen/BR0000019460857https://www.botanicalcollections.be/#/en/specimen/BR0000019459561https://www.botanicalcollections.be/#/en/specimen/BR0000019459585https://www.botanicalcollections.be/#/en/specimen/BR0000019459561https://www.botanicalcollections.be/#/en/specimen/BR0000019460062http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137https://www.botanicalcollections.be/#/en/specimen/BR0000019460024http://db.plantentuinmeise.be/RESEARCH/DATABASES/FOCA/speciesdetails.php?refid=563137
https://www.botanicalcollections.be/#/en/specimen/BR0000021300011https://www.botanicalcollections.be/#/en/specimen/BR0000019460628https://www.botanicalcollections.be/#/en/specimen/BR0000019460390https://www.botanicalcollections.be/#/en/specimen/BR0000019460888https://www.botanicalcollections.be/#/en/specimen/BR0000021300431
https://www.botanicalcollections.be/specimen/BR0000019459592
Journ
al Pre-
proof
Voucher ID URL
F-Archer 7543 https://fm-digital-assets.fieldmuseum.org/668/489/V0203611F.jpg
F-Aristeguieta 2110 https://fm-digital-assets.fieldmuseum.org/668/497/V0203619F.jpg
F-Dahlgreen 49 https://fm-digital-assets.fieldmuseum.org/668/490/V0203612F.jpg
F-Dahlgreen et al. s.n. https://fm-digital-assets.fieldmuseum.org/668/486/V0203608F.jpg
F-Dugand et al. 367 https://fm-digital-assets.fieldmuseum.org/668/492/V0203614F.jpg
F-Proctor 26929 https://plantidtools.fieldmuseum.org/en/rrc/catalogue/399843
F-Steyermark 60962 https://fm-digital-assets.fieldmuseum.org/668/496/V0203618F.jpg
F-Steyermark 86595 https://fm-digital-assets.fieldmuseum.org/668/498/V0203620F.jpg
F-Whitford 21 https://fm-digital-assets.fieldmuseum.org/668/491/V0203613F.jpg
K-Traill 102 http://reflora.jbrj.gov.br/reflora/geral/ExibeFiguraFSIUC/ExibeFiguraFSIUC.do?idFigura=227329818
K-Gardner 5982 http://reflora.jbrj.gov.br/reflora/geral/ExibeFiguraFSIUC/ExibeFiguraFSIUC.do?idFigura=227329828
L-Backer 82 https://data.biodiversitydata.nl/naturalis/specimen/L.2129256
L-Cudjoe 588 https://data.biodiversitydata.nl/naturalis/specimen/WAG.1525309
L-Hart 4492 https://data.biodiversitydata.nl/naturalis/specimen/L.2129265
L-Leeuwenberg 3829 https://data.biodiversitydata.nl/naturalis/specimen/WAG.1525306
L-Shearard et al. 27 https://data.biodiversitydata.nl/naturalis/specimen/L.2129214
MO-Breen 1077 http://legacy.tropicos.org/Specimen/3437069?projectid=83
MO-Breen 1077 http://legacy.tropicos.org/Specimen/3437069?projectid=83
MO-Duke & Kirkbride 14074 http://legacy.tropicos.org/Specimen/1484252
MO-Duke 14471 http://legacy.tropicos.org/Specimen/1509311
MO-Grijalva 1946 http://legacy.tropicos.org/Specimen/119921?projectid=7
MO-Grijalva et al. 6044 http://legacy.tropicos.org/Specimen/1167044?projectid=7
MO-Hinton 16221 http://legacy.tropicos.org/Specimen/3600322?projectid=83
MO-Howell 8 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Kluge 36 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Mertinez 280 http://legacy.tropicos.org/Specimen/906259?projectid=83
MO-Salick 7832 http://legacy.tropicos.org/Specimen/119452?projectid=7
MO-Sexton 114 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Sharp 1962 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Smith et al. 3280 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Standley 27417 http://legacy.tropicos.org/Name/29400114?projectid=56
MO-Trujillo 21360 http://legacy.tropicos.org/Specimen/3233077?projectid=83
P-Benoist s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677857
P-Bélanger s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677815
P-Bonpland 1536 http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677835
P-Brongniart s.n . http://mediaphoto.mnhn.fr/media/1441432208034zCkl0JpVbPYCCrYJ
P-Chevalier 13043 http://mediaphoto.mnhn.fr/media/1441392263356jFHLDhXIkHGFGmRg
P-Eggers 155 http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677894
P-Goossens 3076 http://mediaphoto.mnhn.fr/media/1441392263320idMN4pHedLViPpLS
P-L’Herminier s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p00673894
P-Leblond s.n . http://mediaphoto.mnhn.fr/media/1444518555845GsxSILf84YvzvRhX
P-Pierre 3864 http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677878
P-Plée s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677807
P-Poli s.n. http://mediaphoto.mnhn.fr/media/1441430146200yHNPEAZXFILiS3E2
P-Ricksecker s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677892
P-Riedlé s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p00673892
P-Soubirou s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677852
P-Stehlé 8038 http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677816
P-Stoupy s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p00673893
Journ
al Pre-
proof
P-Wright s.n. http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677897
US-Archer 2615 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1359&width=1024&irn=13625974
US-de Bruijn 1042 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1349&width=1023&irn=13633076
US-Dugand et al. 4105 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1353&width=1023&irn=13613878
US-Duss 2977 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=9003&width=6799&irn=13634824
US-Glassman 2562 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1279&width=1024&irn=10314390
US-Hinton 15865 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1342&width=1023&irn=13631573
US-Kluge 36 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1357&width=1023&irn=13632888
US-Little 16700 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1345&width=1023&irn=13609433
US-Merril 3434 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=9003&width=6845&irn=13618884
US-Palmer 1338 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1307&width=1024&irn=10309737
US-Pittier 10970 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1347&width=1023&irn=13615200
US-Robinson 1765 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1356&width=1023&irn=13618068
US-Romero Castaneda 604 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1341&width=1023&irn=13621309
US-Stehlé 5872 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1351&width=1024&irn=13607126
US-Zimmermann 67 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=640&width=485&irn=13633602
"Artistic" vouchers
P-Hinton 15865 http://coldb.mnhn.fr/catalognumber/mnhn/p/p04677797
P-Leblond s.n. http://mediaphoto.mnhn.fr/media/1444518555845GsxSILf84YvzvRhX
US-Dugand 4105 https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=1342&width=1023&irn=13628137
US-Shannon s.n. https://collections.nmnh.si.edu/search/botany/search.php?action=10&height=639&width=487&irn=13630326
L-Hart s.n. https://data.biodiversitydata.nl/naturalis/specimen/L.2129200
L-Philipson 10414 https://data.biodiversitydata.nl/naturalis/specimen/L.2129260
L-Ibañez & Núñez 816 https://data.biodiversitydata.nl/naturalis/specimen/L.4153935
Journ
al Pre-
proof