1

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

2

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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

Journ

al Pre-

proof

“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

Journ

al Pre-

proof

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

7. References 615

Alcantara-Rodriguez, M., Françozo, M., van Andel, T., 2019. Plant knowledge in the Historia Naturalis 616 Brasiliae (1648): Retentions of seventeenth-century plant use in Brazil. Econ. Bot. 73, 390–617 404. https://doi.org/10.1007/s12231-019-09469-w 618

Anagnostou, S., 2005. Jesuits in Spanish America: contributions to the exploration of the American 619 materia medica. Pharm. in Hist. 47, 3–17. 620

Antczak, A., Urbani, B., Antczak, M.M., 2017. Re-thinking the migration of Cariban-speakers from the 621 Middle Orinoco River to North-Central Venezuela (AD 800). J. World Prehist. 30, 131–175. 622 https://doi.org/10.1007/s10963-017-9102-y 623

Aublet, F., 1775. Histoire des plantes de la Guiane françoise, rangées suivant la méthode sexuelle. 624 Didot, Paris. 625

Bajin ba Ndob, I., Mengome, L.E., Bourobou Bourobou, H.-P., Lossangoye Banfora, Y., Bivigou, F., 626 2016. Ethnobotanical survey of medicinal plants used as anthelmintic remedies in Gabon. J. 627 Ethnopharm. 191, 360–371. https://doi.org/10.1016/j.jep.2016.06.026 628

Balée, W., 2003. Historical-ecological influences on the word for Cacao in Ka’apor. Anthropol. Ling. 45 629 (3), 259–280. 630

Balée, W.L., Moore, D., 1991. Similarity and variation in plant names in five Tupi-Guarani languages 631 (Eastern Amazonia). Bulletin of the Florida Museum of Natural History 35 (4), University of 632 Florida. 633

Barbier, J.B.G., 1824. Traité élémentaire de matière médicale. Méquignon-Marvis, Paris. 634 Barrère, P., 1749. Essai sur l’histoire naturelle de la France équinoxiale. Piget, Paris. 635

Journ

al Pre-

proof

Barrère, P., 1743. Nouvelle relation de la France Equinoxiale. Piget, Paris. 636 Bermúdez, A., Velázquez, D., 2002. Etnobotánica médica de una comunidad campesina del estado 637

Trujillo, Venezuela: un estudio preliminar usando técnicas cuantitativas. Rev. Fac. Farm. 44, 638 2–6. 639

Berry, P.E. (Ed.), 2005. Flora of the Venezuelan Guayana; Rutaceae - Zygophyllaceae, Flora of the 640 Venezuelan Guayana. Missouri Botanical Garden Press, St. Louis. 641

Biblioteca Nacional de Colombia [WWW Document], 2020. https://bibliotecanacional.gov.co/es-co 642 (accessed 6.13.20). 643

Bilby, K., 2000. Reevaluating the African lexical component of the Surinamese Maroon creoles: The 644 Aluku case, in: Thirty-First Annual Conference on African Linguistics, Boston University, 645 March. 646

Blom, C.M., 1763. Dissertatio botanico-medica sistens lignum quassiæ. Upsala. 647 Bodin, S.C., Morin-Rivat, J., Bremond, L., Scheel-Ybert, R., Tardy, C., Vaschalde, C., 2020. Anthracology 648

in the tropics: how wood charcoals help us to better understand today ecosystems, in: 649 Methods in historical ecology: Insights from Amazonia (Odonne, G., Molino, J.-F., Eds.). 650 Routledge, London & New York, pp 87–93. 651

Botanic Garden Meise [WWW Document], 2020. 652 https://www.botanicalcollections.be/#/en/search/specimen (accessed 5.8.20). 653

Botanicus [WWW Document], 2020. http://www.botanicus.org/ (accessed 6.13.20). 654 Boulangeat, L., 2014. Le recensement des Herbiers de France : un nouvel enjeu pour la connaissance. 655

La lettre de l’OCIM 152, 12–16. https://doi.org/10.4000/ocim.1457 656 Boumediene, S., 2020. Jesuit recipes, Jesuit receipts: the Society of Jesus and the introduction of 657

exotic materia medica into Europe, in: Newson, L.A. (Ed.), Cultural worlds of the Jesuits in 658 colonial Latin America. London, pp. 229–254. 659

Boumediene, S., 2016. La colonisation du savoir. Des mondes à faire, Vaulx-en-Velin. 660 Bourdy, G., Aubertin, C., Jullian, V., Deharo, E., 2017. Quassia “biopiracy” case and the Nagoya 661

Protocol: A researcher’s perspective. J. Ethnopharm. 206, 290–297. 662 https://doi.org/10.1016/j.jep.2017.05.030 663

Brack Egg, A., 1999. Diccionario enciclopédico de plantas útiles del Perú. Centro de Estudios 664 Regionales Andinos Bartolomé de las Casas. 665

Brand, E., Leon, C., Nesbitt, M., Guo, P., Huang, R., Chen, H., Liang, L., Zhao, Z., 2017. Economic 666 botany collections: A source of material evidence for exploring historical changes in Chinese 667 medicinal materials. J. Ethnopharm. 200, 209–227. 668 https://doi.org/10.1016/j.jep.2017.02.028 669

Brandão, M.G.L., Cosenza, G.P., Pereira, F.L., Vasconcelos, A.S., Fagg, C.W., 2013. Changes in the 670 trade in native medicinal plants in Brazilian public markets. Environ. Monit. Assess. 185, 671 7013–7023. https://doi.org/10.1007/s10661-013-3081-y 672

Brandão, M.G.L., Pignal, M., Romaniuc, S., Grael, C.F.F., Fagg, C.W., 2012. Useful Brazilian plants 673 listed in the field books of the French naturalist Auguste de Saint-Hilaire (1779–1853). J. 674 Ethnopharm. 143, 488–500. https://doi.org/10.1016/j.jep.2012.06.052 675

Brandão, M.G.L., Zanetti, N.N.S., Oliveira, P., Grael, C.F.F., Santos, A.C.P., Monte-Mór, R.L.M., 2008. 676 Brazilian medicinal plants described by 19th century European naturalists and in the Official 677 Pharmacopoeia. J. Ethnopharm. 120, 141–148. https://doi.org/10.1016/j.jep.2008.08.004 678

Breitbach, U.B., Niehues, M., Lopes, N.P., Faria, J.E.Q., Brandão, M.G.L., 2013. Amazonian Brazilian 679 medicinal plants described by C.F.P. von Martius in the 19th century. J. Ethnopharm. 147, 680 180–189. https://doi.org/10.1016/j.jep.2013.02.030 681

Brousseau, L., Garnier-Géré, P., Clement, C.R., 2020. Historical Genomics, in: Methods in Historical 682 Ecology: Insights from Amazonia (Odonne, G., Molino, J.-F., Eds.). Routledge, London & New 683 York, pp 104–111. 684

Brown, C.H., Clement, C.R., Epps, P., Luedeling, E., Wichmann, S., 2013a. The Paleobiolinguistics of 685 Domesticated Chili Pepper (Capsicum spp.). Ethnobiol. Let. 4, 1–11. 686

Journ

al Pre-

proof

Brown, C.H., Clement, C.R., Epps, P., Luedeling, E., Wichmann, S., 2013b. The Paleobiolinguistics of 687 Domesticated Manioc (Manihot esculenta). Ethnobiol. Let. 4, 61–70. 688

Brown, T.A., Cappellini, E., Kistler, L., Lister, D.L., Oliveira, H.R., Wales, N., Schlumbaum, A., 2015. 689 Recent advances in ancient DNA research and their implications for archaeobotany. Veget. 690 Hist. Archaeobot. 24, 207–214. https://doi.org/10.1007/s00334-014-0489-4 691

Burelli, T., 2019. L’affaire Quassia amara: un cas emblématique de biopiraterie catalyseur de 692 nombreuses évolutions sociales et juridiques. Rev. Jur. Environ. 44, 677–703. 693

Bye, R., Linares, E., Estrada, E., 1995. Biological diversity of medicinal plants in México, in: 694 Phytochemistry of medicinal plants (Arnason, J.T., Mata, R., Romeo, J.T., Eds.). Springer US, 695 Boston, pp. 65–82. https://doi.org/10.1007/978-1-4899-1778-2_4 696

Cabrera, L., 1993. El monte [The forest]. Ediciones Universal. Miami. 697 Calinger, K.M., Queenborough, S., Curtis, P.S., 2013. Herbarium specimens reveal the footprint of 698

climate change on flowering trends across north-central North America. Ecol. Let. 16, 1037–699 1044. https://doi.org/10.1111/ele.12135 700

Carney, J., Rosomoff, R.N., 2011. In the shadow of slavery: Africa’s botanical legacy in the Atlantic 701 world. University of California Press. 702

Carrillo-Rosario, T., Moreno, G., 2006. Importancia de las plantas medicinales en el autocuidado de la 703 salud en tres caseríos de Santa Ana Trujillo, Venezuela. Rev. Fac. Farm. 48, 21–28. 704

Castner, J.L., Timme, S.L., Duke, J.A., 1998. A field guide to medicinal and useful plants of the upper 705 Amazon. Feline Press, Gainesville. 706

Ceuterick, M., Vandebroek, I., Pieroni, A., 2011. Resilience of Andean urban ethnobotanies: A 707 comparison of medicinal plant use among Bolivian and Peruvian migrants in the United 708 Kingdom and in their countries of origin. J. Ethnopharm. 136, 27–54. 709 https://doi.org/10.1016/j.jep.2011.03.038 710

Ceuterick, M., Vandebroek, I., Torry, B., Pieroni, A., 2008. Cross-cultural adaptation in urban 711 ethnobotany: The Colombian folk pharmacopoeia in London. J. Ethnopharm. 120, 342–359. 712 https://doi.org/10.1016/j.jep.2008.09.004 713

Chevalier, A., Russell, W., 1936. Sur un Quassia africain utilisé par les Noirs comme plante médicinale. 714 J. Agr. Trad. Bot. App. 16, 364–367. https://doi.org/10.3406/jatba.1936.5621 715

Chupin, L., 2018. L’informatisation participative d’herbiers comme dispositif de médiation du 716 patrimoine et de la culture scientifique. Quelles convergences avec les dispositifs de 717 formation à la médiation ? Trema 119–131. https://doi.org/10.4000/trema.3954 718

Clement, C.R., Cristo-Araújo, M. de, Coppens d’Eeckenbrugge, G., Reis, V.M. dos, Lehnebach, R., 719 Picanço-Rodrigues, D., 2017. Origin and dispersal of domesticated peach palm. Front. Ecol. 720 Evol. 5, 148. 721

Coe, F.G., 2008. Ethnobotany of the Rama of southeastern Nicaragua and comparisons with Miskitu 722 plant lore. Econ. Bot. 62, 40–59. https://doi.org/10.1007/s12231-008-9006-y 723

Coe, F.G., Anderson, G.J., 1999. Ethnobotany of the Sumu (Ulwa) of southeastern Nicaragua and 724 comparisons with Miskitu plant lore. Econ. Bot. 53, 363–386. 725 https://doi.org/10.1007/BF02866715 726

Coe, F.G., Anderson, G.J., 1996. Ethnobotany of the garífuna of Eastern Nicaragua. Econ. Bot. 50, 71–727 107. https://doi.org/10.1007/BF02862114 728

Collomb, G., 2018. The Quassia amara affair: Political games and issues in French Guyana, from 729 Global to Local. Rech. Amérind. Québec 48 (1/2), 254–255. 730

Cosenza, G.P., Somavilla, N.S., Fagg, C.W., Brandão, M.G.L., 2013. Bitter plants used as substitute of 731 Cinchona spp. (quina) in Brazilian traditional medicine. J. Ethnopharm. 149, 790–796. 732 https://doi.org/10.1016/j.jep.2013.08.004 733

Crawford, M.J., 2016. The Andean wonder drug: Cinchona bark and imperial science in the Spanish 734 Atlantic, 1630-1800. University of Pittsburgh Press, Pittsburgh. 735

Crookes, S.W., 1860. Chemical news and journal of industrial science. Chemical news office. 736 D’Arcy, W.G., 1987. Flora of Panama: checklist and index, Monographs in systematic botany from the 737

Missouri Botanical Garden. Missouri Botanical Garden, Saint Louis. 738

Journ

al Pre-

proof

Davis, D.D., Goodwin, R.C., 1990. Island Carib origins: evidence and nonevidence. Am. Antiq. 55, 37–739 48. 740

Davis, N.Z., 2016. Physicians, healers, and their remedies in colonial Suriname. Can. Bull. Med. Hist. 741 33, 3–34. https://doi.org/10.3138/cbmh.33.1.3 742

del Rey Fajardo, J., 2017. La botica y los boticarios de los Jesuitas en Santafé de Bogotá (1616-1767). 743 Proc. Hist. 32, 103–119. 744

Díaz, R., Hernández, L., Ocampo, R., Cicció, J., 2006. Domesticación y fitoquímica de Quassia amara 745 (Simaroubaceae) en el trópico húmedo de Costa Rica. Lankesteriana 6(2), 49–64. 746 https://doi.org/10.15517/lank.v6i2.19702 747

Dragtenstein, F., 2004. “Trouw aan de blanken”: Quassie van Nieuw Timotibo, twist en strijd in de 748 18de eeuw in Suriname. KIT Publishers, Amsterdam. 749

Duke, J.A., 2008. Duke’s Handbook of Medicinal Plants of Latin America. CRC Press, Boca Raton. 750 Duke, J.A., 1970. Ethnobotanical observations on the Chocó Indians. Econ. Bot. 24, 344–366. 751

https://doi.org/10.1007/BF02860669 752 Eimer, T.R., 2020. What if the subaltern speaks? Traditional knowledge policies in Brazil and India. 753

Third W. Quart. 41, 96–112. 754 El-Gharbaoui, A., Benítez, G., González-Tejero, M.R., Molero-Mesa, J., Merzouki, A., 2017. 755

Comparison of Lamiaceae medicinal uses in eastern Morocco and eastern Andalusia and in 756 Ibn al-Baytar’s Compendium of Simple Medicaments (13th century CE). J. Ethnopharm. 202, 757 208–224. https://doi.org/10.1016/j.jep.2017.03.014 758

Fermin, P., 1769. Description générale, historique, géographique et physique de la colonie de 759 Surinam. Van Harrevelt, Amsterdam. 760

Field Museum [WWW Document], 2020. https://collections-botany.fieldmuseum.org/ (accessed 761 5.8.20). 762

Fournet, J., 1978. Flore illustrée des phanérogames de Guadeloupe et de Martinique. Institut 763 National de la Recherche Agronomique, Paris. 764

French Guiana IRD Herbarium [WWW Document], 2020. http://publish.plantnet-765 project.org/project/caypub (accessed 5.8.20). 766

Gallica [WWW Document], 2020. https://gallica.bnf.fr/html/und/livres/livres?mode=desktop 767 (accessed 6.13.20). 768

Gänger, S., 2015. World trade in medicinal plants from Spanish America, 1717–1815. Med. Hist. 59, 769 44–62. 770

GBIF, Q. africana records [WWW Document], 2020. https://www.gbif.org/fr/species/3709178 771 (accessed 6.12.20). 772

GBIF, Q. amara records [WWW Document], 2020. https://www.gbif.org/species/3190647 (accessed 773 6.11.20). 774

Gentry, A.H., 1996. A field guide to the families and genera of woody plants of north west South 775 America: New edition. University of Chicago Press, Chicago. 776

Giraldo, D., Baquero, E., Bermúdez, A., 2009. Caracterización del comercio de plantas medicinales en 777 los mercados populares de Caracas, Venezuela. Act. Bot. Venez. 32, 267–301. 778

Girón, L.M., Freire, V., Alonzo, A., Cáceres, A., 1991. Ethnobotanical survey of the medicinal flora 779 used by the Caribs of Guatemala. J. Ethnopharm. 34, 173–187. 780 https://doi.org/10.1016/0378-8741(91)90035-C 781

Google Books [WWW Document], 2020. https://books.google.com/?hl=en (accessed 6.13.20). 782 Grandtner, M.M., Chevrette, J., 2013. Dictionary of trees, Volume 2: South America: Nomenclature, 783

taxonomy and ecology. Academic Press/Elsevier, Amsterdam. 784 Grenand, 1995. Le voyage des mots. Logique de la nomination des plantes: exemples dans des 785

langues tupi du Brésil, in: Les mécanismes du changement culturel et linguistique, numéro 786 thématique. Cahiers du Lacito. Paris, pp. 23–42. 787

Grenand, P., Moretti, C., Jacquemin, H., Prévost, M.-F., 2004. Pharmacopées traditionnelles en 788 Guyane: Créoles, Palikur, Wayãpi. IRD Editions, Paris. 789

Journ

al Pre-

proof

Gumilla, J., 1745. El Orinoco ilustrado y defendido. Manuel Fernandez, Supremo Consejo de la 790 Inquisition, Madrid. 791

Gupta, M.P. (Ed.), 1995. 270 plantas medicinales iberamericanas. CYTED-SECAB, Santa Fé de Bogotá. 792 Hart, R., Salick, J., Ranjitkar, S., Xu, J., 2014. Herbarium specimens show contrasting phenological 793

responses to Himalayan climate. Proc. Nat. Acad. Sci. 111, 10615–10619. 794 https://doi.org/10.1073/pnas.1403376111 795

Hather, J.G., 2010. Tropical archaeobotany: applications and new developments. Routledge, London. 796 Heckel, É., 1897. Les plantes médicinales et toxiques de la guyane française: catalogue raisonné et 797

alphabétique. Protat frères, Macon. 798 Heinrich, M., Kufer, J., Leonti, M., Pardo-de-Santayana, M., 2006. Ethnobotany and 799

ethnopharmacology—Interdisciplinary links with the historical sciences. J. Ethnopharm. 107, 800 157–160. https://doi.org/10.1016/j.jep.2006.05.035 801

Hershenson, B.R., 1964. A botanical comparison of the United States pharmacopoeias of 1820 and 802 1960. Econ. Bot. 18, 342–356. 803

House, P.R., Lagos-Witte, S., Ochoa, L., Torres, C., Mejía, T., Rivas, M., 1995. Plantas medicinales 804 comunes de Honduras. Universidad Nacional Autónoma de Honduras, Tegucigalpa. 805

Jaramillo, M.A., Castro, M., Ruiz-Zapata, T., Lastres, M., Torrecilla, P., Lapp, M., Hernández-Chong, L., 806 Muñoz, D., 2014. Estudio etnobotánico de plantas medicinales en la comunidad campesina 807 de Pelelojo, municipio Urdaneta, estado Aragua, Venezuela. Ernstia 24, 85–110. 808

Jaramillo-Arango, J., 1952. Don José Celestino Mutis y las expediciones botánicas españolas del siglo 809 XVIII al Nuevo Mundo. Prensas del Ministerio de Educación, Bogota. 810

Lardos, A., 2015. Historical approaches in ethnopharmacology, in: Ethnopharmacology (Heinrich, M., 811 Jäger, A.K., Eds.). J. Wiley & sons, Chichester. pp. 333–341. 812

Lardos, A., Heinrich, M., 2013. Continuity and change in medicinal plant use: The example of 813 monasteries on Cyprus and historical iatrosophia texts. J. Ethnopharm. 150, 202–214. 814 https://doi.org/10.1016/j.jep.2013.08.026 815

Leblond, J.-B., 1789. Lettre de Jean-Baptiste Leblond au ministre de la Marine. Cayenne, 28 816 novembre 1789. Archives Nationales de l’Outre-Mer (ref ANOM/EE 1232). 817

Leonti, M., Cabras, S., Weckerle, C.S., Solinas, M.N., Casu, L., 2010. The causal dependence of present 818 plant knowledge on herbals—Contemporary medicinal plant use in Campania (Italy) 819 compared to Matthioli (1568). J. Ethnopharm. 130, 379–391. 820 https://doi.org/10.1016/j.jep.2010.05.021 821

Lindeman, J.C., Mennega, A.M.W., Hekking, W.H.A., 1963. Bomenboek voor Suriname. Dienst's Lands 822 Bosbeheer Suriname, Paramaribo. 823

Linné, C. von, 1762. Species plantarum. Editio Secunda, Tomus 1. Laurentius Salvius, Stockholm. 824 Longanga Otshudi, A., Vercruysse, A., Foriers, A., 2000. Contribution to the ethnobotanical, 825

phytochemical and pharmacological studies of traditionally used medicinal plants in the 826 treatment of dysentery and diarrhoea in Lomela area, Democratic Republic of Congo (DRC). J. 827 Ethnopharm. 71, 411–423. https://doi.org/10.1016/S0378-8741(00)00167-7 828

Lorenzi, H., Matos, F.J. de A., 2008. Plantas medicinais no Brasil: nativas e exóticas. Instituto 829 Plantarum de Estudos da Flora, Nova Odessa/São Paulo. 830

Mbatchi, S.F., Mbatchi, B., Banzouzi, J.T., Bansimba, T., Nsonde Ntandou, G.F., Ouamba, J.-M., Berry, 831 A., Benoit-Vical, F., 2006. In vitro antiplasmodial activity of 18 plants used in Congo 832 Brazzaville traditional medicine. J. Ethnopharm. 104, 168–174. 833 https://doi.org/10.1016/j.jep.2005.08.068 834

Medeiros, M.F.T., 2016. Historical ethnobiology, in: Introduction to Ethnobiology (Albuquerque, U.P., 835 Nobrega Alves, R.R., Eds.). Springer, Heidelberg, pp. 19–24. 836

Mérat, J.-F., de Lens, A.-J., 1837. Dictionnaire universel de matière médicale et de thérapeutique 837 générale, contenant l’indication, la description et l’emploi de tous les médicaments connus 838 dans les diverses parties du globe. Société belge de librairie, Bruxelles. 839

Merian, M.S., 1705. Metamorphosis insectorum Surinamensium: Of te verandering der Surinaamsche 840 insecten. G. Valck, Amsterdam. 841

Journ

al Pre-

proof

Missouri Botanical Garden [WWW Document], 2020. 842 http://legacy.tropicos.org/Name/29400114?projectid=3 (accessed 5.8.20). 843

Moreira, P.A., Aguirre-Dugua, X., Mariac, C., Zekraoui, L., Couderc, M., Rodrigues, D.P., Casas, A., 844 Clement, C.R., Vigouroux, Y., 2017. Diversity of treegourd (Crescentia cujete) suggests 845 introduction and prehistoric dispersal routes into Amazonia. Front. Ecol. Evol. 5, 150. 846

Museum National d’Histoire Naturelle [WWW Document], 2020. 847 https://science.mnhn.fr/institution/mnhn/collection/p/item/search/form (accessed 5.8.20). 848

Musuyu Muganza, D., Fruth, B.I., Nzunzu Lami, J., Mesia, G.K., Kambu, O.K., Tona, G.L., Cimanga 849 Kanyanga, R., Cos, P., Maes, L., Apers, S., Pieters, L., 2012. In vitro antiprotozoal and cytotoxic 850 activity of 33 ethnopharmacologically selected medicinal plants from Democratic Republic of 851 Congo. J. Ethnopharm. 141, 301–308. https://doi.org/10.1016/j.jep.2012.02.035 852

Mutis, J.C., 1778. Copia fragmentada del informe presentado por José Celestino Mutis al virrey sobre 853 muestras de quina de la Guayana. 854

Mutis, J.C., de Gregorio, M.H., 1828. El arcano de la quina: discurso que contiene la parte médica de 855 las cuatro especies de quinas oficinales, sus virtudes eminentes y su legítima preparación: 856 obra póstuma. Ibarra, Madrid. 857

Naturalis Biodiversity Center [WWW Document], 2020. https://bioportal.naturalis.nl/ (accessed 858 5.8.20). 859

Nossin, E., 2019. Plantes magiques de Martinique, Guadeloupe et des Petites Antilles. Exbrayat 860 Editions, Fort de France. 861

Ocampo, R., Mora, G., 2011. Ethnomedicine of Quassia and related plants in Tropical America. in: 862 Ethnomedicinal Plants: Revitalizing of Traditional Knowledge of Herbs (Rai, M., Acharya, D., 863 Rios, J.L., Eds.). CRC Press, Boca Raton, pp. 301-325. 864

Odonne, G., Bourdy, G., Beauchêne, J., Houël, E., Stien, D., Chevolot, L., Deharo, E., 2007. From Tonic-865 cups to Bitter-cups: Kwasi bita beker from Suriname: Determination, past and present use of 866 an ancient galenic artefact. J. Ethnopharm. 110, 318–322. 867 https://doi.org/10.1016/j.jep.2006.09.045 868

Pedrollo, C.T., Kinupp, V.F., 2015. Sustainability or colonialism? Legislative obstacles to research and 869 development of natural products and patents on traditional knowledge in Brazil. Act. Bot. 870 Bras. 29, 452–456. 871

Perrier, X., Langhe, E.D., Donohue, M., Lentfer, C., Vrydaghs, L., Bakry, F., Carreel, F., Hippolyte, I., 872 Horry, J.-P., Jenny, C., Lebot, V., Risterucci, A.-M., Tomekpe, K., Doutrelepont, H., Ball, T., 873 Manwaring, J., Maret, P. de, Denham, T., 2011. Multidisciplinary perspectives on banana 874 (Musa spp.) domestication. Proc. Nat. Acad. Sci. 108, 11311–11318. 875 https://doi.org/10.1073/pnas.1102001108 876

Peruvian Checklist, Missouri Botanical Garden [WWW Document], 2020. 877 http://legacy.tropicos.org/Name/29400114?projectid=5 (accessed 5.8.20). 878

Picking, D., Delgoda, R., Younger, N., Germosén-Robineau, L., Boulogne, I., Mitchell, S., 2015. TRAMIL 879 ethnomedicinal survey in Jamaica. J. Ethnopharm. 169, 314–327. 880 https://doi.org/10.1016/j.jep.2015.04.027 881

Pieroni, A., Vandebroek, I., 2007. Traveling cultures and plants: The ethnobiology and 882 ethnopharmacy of Human migrations, 1st ed. Berghahn Books, New York. 883

Pio Corrêa, M., 1926. Dicionário das plantas úteis do Brasil e das exóticas cultivadas, vol. I, II. 884 Imprensa Nacional, Rio de Janeiro. 885

Pio Corrêa, M., 1969. Dicionário das plantas úteis do Brasil e das exóticas cultivadas, vol. IV. Imprensa 886 Nacional, Rio de Janeiro. 887

Pio Corrêa, M., 1978. Dicionário das plantas úteis do Brasil e das exóticas cultivadas, vol. III, V, VI. 888 Imprensa Nacional, Rio de Janeiro. 889

Pittier, H., 1908. Ensayo sobre las plantas usuales de Costa Rica. H.L. & J.B. McQueen Inc., 890 Washington. 891

Price, R., 1983. First-Time: The historical vision of an Afro-American people. John Hopkins Studies in 892 Atlantic History and Culture. The John Hopkins University Press, Baltimore. 893

Journ

al Pre-

proof

Price, R., 1979. Kwasimukamba’s gambit. Bijdragen tot de Taal-, Land- en Volkenkunde 135 (1), 151–894 169. 895

Price, S., Price, R., Strobel, M.-B., Dutrait, L., Robert, D., Colleu, G., 2005. Les arts des Marrons. Vents 896 d’ailleurs, La Roque-d'Anthéron. 897

Przelomska, N.A.S., Armstrong, C.G., Kistler, L., 2020. Ancient plant DNA as a window into the cultural 898 heritage and biodiversity of our food system. Front. Ecol. Evol. 8. 899 https://doi.org/10.3389/fevo.2020.00074 900

Pulle, A.A., Stoffers, A.L., Lindeman, J.C., Foundation van Eedenfonds (Eds.), 1979. Flora of Suriname. 901 Vol. 5 Pt. 1: Simaroubaceae, Papaveraceae, Vitaceae, Icacinaceae, Theaceae, 902 Theophrastaceae, Nymphaeaceae, Cabombaceae, Nelumbonaceae, Musaceae, 903 Zingiberaceae, Liliaceae. Brill, Leiden. 904

Reclu, M., 1889. Manuel de l’herboriste: comprenant la culture, la récolte, la conservation les 905 propriétés médicinales des plantes du commerce et un dictionnaire des maladies et des 906 remèdes. JB Baillière, Paris. 907

Reflora [WWW Document], 2020. URL 908 http://reflora.jbrj.gov.br/reflora/herbarioVirtual/ConsultaPublicoHVUC/ConsultaPublicoHVU909 C.do (accessed 9.25.20). 910

Rios, M., Koziol, M.J., Pedersen, H.B., Granda (Eds.), 2007. Plantas útiles del Ecuador: aplicaciones, 911 retos y perspectivas. Ediciones Abya-Yala, Quito. 912

Rolander, D., 2008. The Suriname journal: Composed during an exotic journey (trans. J. Dobreff, C. 913 Dahlman, D. Morgan & J. Tipton). The Linnaeus apostles: Global science & adventure 3, 914 1215–1576. 915

Roques, J., 1837. Nouveau traité des plantes usuelles: spécialement appliqué à la médecine 916 domestique, et au régime alimentaire de l’homme sain ou malade. P. Dufart, Paris. 917

Rossetto, M., Ens, E.J., Honings, T., Wilson, P.D., Yap, J.-Y.S., Costello, O., Round, E.R., Bowern, C., 918 2017. From songlines to genomes: Prehistoric assisted migration of a rain forest tree by 919 Australian Aboriginal people. Plos One 12, e0186663. 920 https://doi.org/10.1371/journal.pone.0186663 921

Roullier, C., Benoit, L., McKey, D.B., Lebot, V., 2013a. Historical collections reveal patterns of diffusion 922 of sweet potato in Oceania obscured by modern plant movements and recombination. Proc. 923 Nat. Acad. Sci. 110, 2205–2210. https://doi.org/10.1073/pnas.1211049110 924

Roullier, C., Duputié, A., Wennekes, P., Benoit, L., Fernández Bringas, V.M., Rossel, G., Tay, D., McKey, 925 D., Lebot, V., 2013b. Disentangling the origins of cultivated sweet potato (Ipomoea batatas 926 (L.) Lam.). Plos One 8, e62707. https://doi.org/10.1371/journal.pone.0062707 927

Royal Botanic Gardens Kew [WWW Document], 2020. 928 http://apps.kew.org/herbcat/gotoHomePage.do (accessed 5.8.20). 929

Sathasivampillai, S.V., Rajamanoharan, P.R.S., Munday, M., Heinrich, M., 2017. Plants used to treat 930 diabetes in Sri Lankan Siddha Medicine – An ethnopharmacological review of historical and 931 modern sources. J. Ethnopharm. 198, 531–599. https://doi.org/10.1016/j.jep.2016.07.053 932

Shepard Jr, G.H., Ramirez, H., 2011. “Made in Brazil”: Human dispersal of the Brazil nut (Bertholletia 933 excelsa, Lecythidaceae) in ancient Amazonia. Econ. Bot. 65, 44–65. 934

Smithsonian National Museum of Natural History [WWW Document], 2020. 935 https://collections.nmnh.si.edu/search/botany/ (accessed 5.8.20). 936

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

Journ

al Pre-

proof

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

van’t Klooster, C.I., Lindeman, J.C., Jansen-Jacobs, M.J., 2003. Index of vernacular plant names of 978 Suriname. Nationaal Herbarium Nederland, Universiteit Leiden Branch, Leiden. 979

Voeks, R., Greene, C., 2018. God’s healing leaves: The colonial quest for medicinal plants in the Torrid 980 Zone. Geograph. Rev. 108, 545–565. 981

Voeks, R., Rashford, J. (Eds.), 2012. African ethnobotany in the Americas, 2013 edition. Springer, New 982 York. 983

Warner-Lewis, M., 2003. Central Africa in the Caribbean: Transcending time, transforming cultures. 984 University of the West Indies Press, Kingston. 985

Westengen, O.T., Okongo, M.A., Onek, L., Berg, T., Upadhyaya, H., Birkeland, S., Khalsa, S.D.K., Ring, 986 K.H., Stenseth, N.C., Brysting, A.K., 2014. Ethnolinguistic structuring of sorghum genetic 987 diversity in Africa and the role of local seed systems. Proc. Nat. Acad. Sci. 111, 14100–14105. 988 https://doi.org/10.1073/pnas.1401646111 989

Willis, C.G., Ellwood, E.R., Primack, R.B., Davis, C.C., Pearson, K.D., Gallinat, A.S., Yost, J.M., Nelson, 990 G., Mazer, S.J., Rossington, N.L., Sparks, T.H., Soltis, P.S., 2017. Old plants, new tricks: 991 Phenological research using herbarium specimens. Tr. Ecol. Evol. 32, 531–546. 992 https://doi.org/10.1016/j.tree.2017.03.015 993

Woodson, R.E., Schery, R.W., Porter, D.M., 1973. Flora of Panama. Part VI. Family 90. Simaroubaceae. 994 Ann. Miss. Bot. Gard. 60, 23–39. 995

Journ

al Pre-

proof

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

Journ

al Pre-

proof

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