Journal of Ethnopharmacology

From forest to pharmacy: should we be depressed about a sustainable Griffonia

simplicifolia (Fabaceae) seed supply chain?
--Manuscript Draft--

Manuscript Number: JETHNO-D-21-01041

Article Type: Review Article

Keywords: Griffonia; harvesting impacts; 5-Hydroxytryptophan; interactive impacts; population

dynamics

Abstract: Ethnopharmacological relevance

Griffonia simplicifolia D.C (Baill.) (Fabaceae) seeds are unusually high (6-20% wet
weight) in 5-HTP (5-Hydroxytryptophan), a serotonin precursor widely used to treat
depression. Consequently, this species is regarded as a herbal “Prozac®”.
Contemprary use as an anti-depressant contrasts with traditional uses for insecticides,
arachnicides, fodder, dyes, mordants and chewing-sticks. G. simplicifolia seeds are
wild-harvested for the export trade. Over the past 15 years, use of 5-HTP extracted
from G. simplicifolia in cosmetics has added to global demand. Wild populations in
West Africa are the sole commercial source of G. simplicifolia seed. Aims of the study:
Were to (i) assess the scale of the global trade in G. simplicifolia seeds and (ii) produce
a synthesis of the challenges facing sustainable harvest of G. simplicifolia.

Materials and approach

Firstly, we analysed global trade data for G. simplicifolia, taking into account historical
trends over the past 40 years. Secondly, we reviewed published studies on the
distribution, population biology and harvest impacts of wild G. simplicifolia populations.

Results and Conclusions

Wild G. simplicifolia populations have been the focus of commercial harvest of their
pods (for seeds) for international trade from West Africa for almost 50 years. In the late
1980's, when Ghana exported 75-80 metric tonnes (MT) of G. simplicifolia seed to
Europe, this species was already Ghana's main medicinal plant export. Currently, 5
West African countries export G. simplicifolia seeds (Cote d'Ivoire, Ghana, Liberia,
Nigeria and Togo). Although in the 1980’s, most seed exports were to Europe, today
China is the main importer of G. simplicifolia seed. These seeds are value-added for
production of 5-HTP extracts, and then re-exported, particularly to North America
(c.48% of exports). The low habitat specificity and vigorous re-sprouting of G.
simplicifolia after cutting, plus its occurrence in forest reserves and national parks
confer some resilience on wild populations. Sustaining future supply chains faces six
future challenges, however: (1) Rapid loss of forest habitats; (2) Declining populations
of understorey birds and disruption of G. simplicifolia pollination in this bird pollinated
species; (3) Negative effects of introduced invasive plant species (Broussonetia
papyrifera, Chromolaena odorata) on G. simplicifolia regeneration; (4) Grazing by
livestock and use of G. simplicifolia leaves as forage; (5) The long-term impact of
industrial scale seed “predation”: Over a 9-year period (2005 – 2013), G. simplicifolia
exports from Ghana totalled at least 5550 metric tonnes (or between 9.1 billion to 13.5
billion seeds). This could affect the long-term population dynamics of this species,
which produces a low number of seeds per pod (1-4 seeds) and has short distance
(ballistic) seed dispersal; and (6) Destructive harvest methods, when plants are cut to
harvest get the seed pods. Improved resource management, monitoring, quality control
and careful pricing are important if supply chains from wild stocks are to be maintained.
If wild populations decline, then 5-HTP biosynthesis may compete with low G.
simplicifolia seed yields, leading to loss of income to West African harvesters and
traders.

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Manuscript File

From forest to pharmacy: should we be depressed about a sustainable Griffonia

simplicifolia (Fabaceae) seed supply chain?

A.B. Cunninghama, b*, J.A. Brinckmannc and D.E.V. Harter d

aSchool of Life Sciences, University of KwaZulu-Natal, King Edward Avenue, Pietermaritzburg, 3209, South Africa; and b

School of Veterinary and Life Sciences, Murdoch University, 90 South St., Murdoch WA 6150, Australia.

c Traditional Medicinals, 4515 Ross Road, Sebastopol, California 95472, USA

d Bundesamt für Naturschutz (BfN), Konstantinstr. 110, Bonn 53179, Germany

*Corresponding author. E-mail address: tonyc05@bigpond.net.au

Keywords: Griffonia, harvesting impacts, 5-Hydroxytryptophan, interactive impacts, population dynamics.

Abbreviations: 5-HTP, 5-Hydroxytryptophan; ASNAPP: Agribusiness in Sustainable Natural African Plant

Products; B2B, Business to Business; BOTPAL, Botanical Products Association of Liberia; BTI, Binding Tariff
Information rulings of the European Commission Taxation and Customs Union; CITES, Convention on
International Trade of Endangered Species; COSING, Cosmetic Ingredients & Substances Database (European
Commission); EU, European Union; FOB, Free-on-Board; HS, harmonized system (tariff codes); FAO, Food and
Agriculture Organization of the United Nations; IPM, Integral Projection Models; NACGRAB, National Centre
for Genetic Resources and Biotechnology; NNHPD, Natural and Non-prescription Health Products Directorate
(Health Canada); PROSPER, People, Rules and Organizations Supporting the Protection of Ecosystem
Resources; TMR, Transparency Market Research; ZAUBA, Zauba Technologies Pvt Ltd.

TABLE OF CONTENTS
Abstract
1.0 Introduction
2.0 Methods and approach
3.0 Results and Discussion
3.1 Scientific names, synonyms, trade names and vernacular names
3.2 Geographic distribution
3.3 Utilization and trade
3.3.1 Uses of G. simplicifolia in West Africa
3.3.2 International export trade
3.4 Seed and leaf chemistry
3.5 Challenges of complex interactive effects on G. simplicifolia populations
3.6 A future challenge: the economics of 5-HTP from wild harvest vs. metabolic engineering

4.0 Conclusions

5.0 Acknowledgements
6.0 References
Authors contributions
Conflict of interest statement

Abstract

Ethnopharmacological relevance: Griffonia simplicifolia D.C (Baill.) (Fabaceae) seeds are unusually
high (6-20% wet weight) in 5-HTP (5-Hydroxytryptophan), a serotonin precursor widely used to treat
depression. Consequently, this species is regarded as a herbal “Prozac®”. Contemprary use as an
anti-depressant contrasts with traditional uses for insecticides, arachnicides, fodder, dyes, mordants
and chewing-sticks. G. simplicifolia seeds are wild-harvested for the export trade. Over the past 15
years, use of 5-HTP extracted from G. simplicifolia in cosmetics has added to global demand. Wild
populations in West Africa are the sole commercial source of G. simplicifolia seed. Aims of the
study: Were to (i) assess the scale of the global trade in G. simplicifolia seeds and (ii) produce a
synthesis of the challenges facing sustainable harvest of G. simplicifolia. Materials and approach:
Firstly, we analysed global trade data for G. simplicifolia, taking into account historical trends over
the past 40 years. Secondly, we reviewed published studies on the distribution, population biology
and harvest impacts of wild G. simplicifolia populations. Results and Conclusions: Wild G.
simplicifolia populations have been the focus of commercial harvest of their pods (for seeds) for
international trade from West Africa for almost 50 years. In the late 1980's, when Ghana exported
75-80 metric tonnes (MT) of G. simplicifolia seed to Europe, this species was already Ghana's
main medicinal plant export. Currently, 5 West African countries export G. simplicifolia seeds (Cote
d'Ivoire, Ghana, Liberia, Nigeria and Togo). Although in the 1980’s, most seed exports were to
Europe, today China is the main importer of G. simplicifolia seed. These seeds are value-added for
production of 5-HTP extracts, and then re-exported, particularly to North America (c.48% of
exports). The low habitat specificity and vigorous re-sprouting of G. simplicifolia after cutting, plus
its occurrence in forest reserves and national parks confer some resilience on wild populations.
Sustaining future supply chains faces six future challenges, however: (1) Rapid loss of forest
habitats; (2) Declining populations of understorey birds and disruption of G. simplicifolia pollination
in this bird pollinated species; (3) Negative effects of introduced invasive plant species (Broussonetia
papyrifera, Chromolaena odorata) on G. simplicifolia regeneration; (4) Grazing by livestock and use
of G. simplicifolia leaves as forage; (5) The long-term impact of industrial scale seed “predation”:
Over a 9-year period (2005 – 2013), G. simplicifolia exports from Ghana totalled at least 5550 metric
tonnes (or between 9.1 billion to 13.5 billion seeds). This could affect the long-term population
dynamics of this species, which produces a low number of seeds per pod (1-4 seeds) and has short
distance (ballistic) seed dispersal; and (6) Destructive harvest methods, when plants are cut to
harvest get the seed pods. Improved resource management, monitoring, quality control and careful
pricing are important if supply chains from wild stocks are to be maintained. If wild populations
decline, then 5-HTP biosynthesis may compete with low G. simplicifolia seed yields, leading to loss
of income to West African harvesters and traders.

1.0 Introduction

Depression is a widely diagnosed mental disorder affecting the lives of over 300 million people
worldwide (or 4.4 % of the global population), posing a huge demand for pharmaceutical
antidepressants and therapies (Gaynes et al., 2005; Kessler et al., 1994, World Health Organization
2017). Serotonin is well known as one of three important hormones produced by the pineal gland in
mammals (including humans) and is a neurotransmitter regulating mood, sleep, and anxiety in
mammals, as well as alcoholism in humans (Veenstra-VanderWeele, et al., 2000). Interest in use of
herbal alternatives to pharmaceutical antidepressants is increasing. Hypericum perforatum L., for
example, is used to treat mild to moderate depression, despite concerns about toxicity and drug
interactions (Hammerness et al., 2003). Kava (Piper methysticum G. Forst.) is also sometimes used
to treat depression and anxiety, with research into aqueous kava extracts after withdrawal of some
other forms of P. methysticum from sale in Europe, the UK and Canada due to worries about
hepatotoxic reactions (Sarris et. al., 2009). Since the late 1950’s, serotonin deficiency in the brain
has been considered one of the causes of depression, with medication known to increase serotonin
levels (Coppen, 1963; 1967; Pare, 1959). And since the 1970’s, studies have been done on the
serotonin precursor, 5-HTP (5-Hydroxytryptophan) (Berdonces, 2015).

Currently, the most cost-effective source of 5-HTP is from seeds of Griffonia simplicifolia D.C (Baill.),
a scandent climber or liana, that is one of four species in a small genus in the tribe Cercideae, which
forms the most ancient living clade in the Fabaceae. While the other three Griffonia species are
restricted to rainforest habitat, G. simplicifolia is more widespread, occurring in a variety of habitats
in eight West African countries: Benin, Cameroon, Cote d'Ivoire, Gabon, Ghana, Liberia, Nigeria and
Togo. All four Griffonia species are characterised by an unusual metabolic pathway that produces
very high concentrations of an unusual amino acid, 5-hydroxytryptophan (5-HTP) (also commercially
known as oxitriptan) in the seeds (Bell, 1970; Fellows & Bell, 1971). 5-HTP is an intermediate
metabolite of the amino acid L-tryptophan (LT) in the biosynthetic pathway for the
neurotransmitter serotonin in animals (Birdsall, 1998). In addition, G. simplicifolia seeds (and leaves
in small quantities) also contain serotonin (5-hydroxytryptamine (5-HT)). Experimental studies
support the theory that the production of 5-HTP helps defend plants against seed predators such as
Bruchid beetles (Janzen, et al., 1977) and that 5-HT reduces fungal infection of plant leaves, such as
in rice (Ishihara, et al., 2008; Kang et al., 2007). Ironically, while the high concentration of 5-HTP in
G. simplicifolia seeds (up to ~20% (w/w), Lemaire and Adosraku, 2002) defend plants against
predatory insects, it has had the opposite effect in terms of human use: attracting commercial
harvest for trade and consumption. Consequently, G. simplicifolia has become known as an herbal
alternative to the anti-depressant, selective serotonin reuptake inhibitor (SSRI) drug “Fluoxetine
hydrochloride USP” (brand name Prozac®, Eli Lilly and Company). And for almost 50 years since
clinical use of 5-HTP started, there has been international demand for G. simplicifolia seeds
precisely because they contain 5-HTP. Use of 5-HTP from G. simplicifolia seeds in cosmetics (to
reduce the effects of ultra-violet radiation on the skin) also appears to be increasing, with patents
taken out in France (Meybeck, 2005) and an application the USA (Meybeck, 2009).

By the late 1980’s, there already was unease about destructive cutting of G. simplicifolia stems to
get the pods for commercial trade (Ake-Assi, L., pers.comm., 1989; Cunningham, 1993; Cunningham
& Schippmann, 2000; Bosch, 2008). In 1989, international trade in G. simplicifolia for 5-HTP
increased when the United States Food and Drug Administration (FDA) banned its precursor, L-
tryptophan, because of links to the eosinophilia-myalgia syndrome (Birdsall, 1998). Since then, 5-
HTP has been marketed as a safe alternative to L-tryptophan (Bowers, et al., 2003). The rise in G.
simplicifolia seed exports from Ghana, the main exporting country, reflects the rising demand for 5-
HTP. In the late 1980's, when Abbiw (1990) reported exports of 75-80 metric tonnes (MT) of G.
simplicifolia seed, this species had already reached the status of Ghana's main medicinal plant
export. By the early 1990's, G. simplicifolia seed exports were from the two Range States with the
largest G. simplicifolia populations (Cote d'Ivoire and Ghana), with Germany the main importing
country. By 2003, a single company in Ghana (Tek-Spec. Co. Ltd.) reported being able to supply up
to 100 MT of G. simplicifolia seeds annually (Brinckmann, 2003). Estimates of the size of the global
market for bulk 5-HTP vary. One report valued the market at US$ 31.7 million, based on demand for
136.4 tonnes of extract (Transparency Market Research (TMR), 2014), while another source
estimated the market at about 120 tonnes (bulk value US$ 50 to 100 million) with an annual growth
rate of about 7% (Yan & Lin, 2015). According to the latter source, in terms of global markets, the
North American region forms about 48% of the global market for bulk 5-HTP produced from G.
simplicifolia seeds, followed by Europe and Asia Pacific.

Although G. simplicifolia is one of the most studied commercially exploited African medicinal plants,
with 1010 publications on this species at the time van Wyk (2015) did his review, studies on trade
and sustainable harvest in G. simplicifolia products remained a gap. An exception are the studies by
Hwang (2017) and Hwang et al. (2018), whose participatory work with G. simplicifolia harvesters in
Liberia showed that non-destructive harvest of pods could benefit harvesters themselves.
Underscoring the need for resource management plans and monitoring across the range of a
valuable West Africa export for which wild populations are the sole source of supply.

Stimulated by increased G. simplicifolia seed exports from West Africa that reflect the growing
global demand for 5-HTP and anti-depressants, this study had two aims: (i) to assess the scale of
the global trade in G. simplicifolia seeds and (ii) due to the industrial scale of wild harvest, to review
studies on the population biology of G. simplicifolia that assessed the challenges facing sustainable
harvest.

2.0 Methods and approach

Firstly, to get clarity on the trade assessment, we accessed all available published information on
the taxonomy and trade names for G. simplicifolia.

Secondly, trade data for G. simplicifolia was analysed using a range of sources. Trade from West
Africa countries was derived from internet based sources, including B2B trade portals such as Africa
Export Forum (http://www.export-forum.com/africa/), Alibaba (https://www.alibaba.com/),
Exporters India (https://www.exportersindia.com/), and TradeBoss
(http://www.tradeboss.com/default.cgi/), the websites of Griffonia seed exporting companies, and
through personal communication (by JB) with Ghanaian exporters. The Alibaba Group on-line
platform for global wholesale trade was also used to help quantify locations of companies listed at
“supplying” Griffonia raw materials, processed ingredients and finished products by using three
search terms: “Griffonia seed”, “Griffonia seed extract” and “Griffonia seed extract 5 htp”. Recent
data on US import shipments of Griffonia ingredients (Jan-Sept. 2019) were purchased from the
India-based Zauba Technologies & Data Services Pvt Ltd (https://www.zauba.com/), which provided
data from actual shipment documents including date of shipment, harmonized system tariff codes
(HS Codes) used per shipment, ports of loading and discharge, names of the shippers and buyers,
quantity of shipping units and net weights. Analysis of trade by HS Code was complicated as we
realized that some exporters had assigned incorrect codes in their shipment documentation. For
example, incorrect HS Codes used for shipments of Griffonia seed extract included HS 1209.24
(Kentucky bluegrass (Poa pratensis L.) seed), HS 2933.99 (other heterocyclic compounds with
nitrogen hetero-atom(s) only), HS 3104.30 (potassium sulphate), HS 3201.10 (quebracho bark
extract), HS 3301.29 (other essential oils other than those of citrus fruit), and HS 3912.90 (other
cellulose and its chemical derivatives). The correct (albeit general and not species-specific) HS Code
for Griffonia seed would be HS 1211.90 while HS 1302.19 or HS 1302.39 would be appropriate for
different types of extracts of the seeds. The available data also suggested that exporters of purified
5-HTP are applying the general HS Code for extracts (HS 1302.19) rather than using an HS Code
suitable for purified chemical substances. Incorrect or inconsistent tariff classification by exporters
confounds analysis.

Thirdly, we reviewed published information on G. simplicifolia ecology and population biology and
on the impacts of seed harvest on G. simplicifolia populations from across the geographic range of
this species.

3.0 Results and Discussion

3.1 Scientific names, synonyms, trade names and vernacular names

Correct species classification and naming is not only important for assessing trade and traditional
use, but is also important for correct labeling of herbal ingredients and dietary supplements. With
G. simplicifolia (formerly Bandeiraea simplicifolia (DC.) Benth., Schotia simplicifolia M. Vahl ex DC.
(The Plant List, Version 1.1, 2013), this is relatively easy, as Griffonia is used as a trade name in
English, French and German (which also uses the name Afrikanische Schwarzbohne). Local names
reflect the linguistic diversity of West Africa, however. Cote d'Ivoire: Ga: tôtô, Akan-brong: kadia;
Anyin (or Anyi): borokotoa; Baule: eggo; ‘Kru’: balio, Kru-Bete: kokula, kpe koléï; Wè (or
Guere): boogra; buogla; gera; ko kwè; kpakwalè; Ghana: Akan: Kagya; Adangme: gbogbloto, with
totolimo the name given to the fruits; Abron (or Brong): kagya; Fante: kagya; Wasa: kagya or
aboto; Twi: kagya or kanya, with toto the name for the fruits; Ga: kanya; tooto (and toto for the
fruits); Éwé (or Vhe): gbato gbato; gogo and also in Éwé, but listed in Burkill (1995) as Gbe names
(although Gbe is an alternative name for Éwé): akpee, gbogbotri; woto-woto; Liberia: Maan (or
Mano): po pio lo (Burkill, 1985) and glabler, guoblo or wawatoto (ASNAPP (www.asnapp.org.gh));
Nigeria: Igbo: kpokirikpo, with kporikpo (referring to the plant) and edekpurakpu (referring to the
fruit and seed), Yoruba: alukoko. With the exception of the names recorded for Liberia by ASNAPP
the rest of these local names were recorded in Burkill (1985). Some of which are onomatopoeic, due
to the sound made by the ripe pods when they explosively disperse the seeds. However we have
updated the names for local languages in Burkill (1985) based on currently accepted spellings in
www.ethnologue.com).

3.2 Geographic distribution

Griffonia simplicifolia is the most widespread species in this small genus (of 4 species) restricted to
west tropical Africa. Wild populations of G. simplicifolia occur in eight African countries: Benin,
Cameroon, Cote d'Ivoire, Gabon, Ghana, Liberia, Nigeria and the Togolese Republic (Aubréville,
1968; Lock, 1989; Figure 1A). Interestingly, neither Aubréville (1970) in the Flora of Cameroon nor
Lock (1989) record G. simplicifolia in Benin or Cameroon. Griffonia simplicifolia is however recorded
in the Campo-Ma'an forest area of southern Cameroon by Tchouto (2004). Both the “Plants For A
Future” (PFAF) and the “Useful Tropical Plants” databases give the Congo as a Range State but this is
doubtful.

[insert Figure 1. A. The geographic distribution of G. simplicifolia (based on distribution data from the GBIF
Secretariat (2017), excluding specimen localities that are most likely from planted specimens (for example, in
Dakar, Senegal). B. Map summarising cross-border trade and exports, showing estimated quantities of seed
exports from Cote d’Ivoire, Ghana and Liberia, with export quantities from Cameroon and Nigeria unknown.
Juliani (2013) suggested that Ghana exported 2000 tonnes of G. simplicifolia seed, 30-40% of which was from
neighbouring countries such as Cote d’Ivoire and Liberia. Our export data for Ghana is based on ASNAPP data
(2005-2013, Table 2), which gave an average of 616.67 tonnes of seeds (air dry mass) exported per year
(rounded to 617 tonnes here). We suggest that 616.67 tonnes/year represents an underestimate of total
exports, as the ASNAPP data for 2005-2013 was based on interviews with key exporters, so may have missed
smaller quantities exported.

Griffonia simplicifolia has a fragmented distribution across a variety of habitats across the
phytogeographic zones of Guineo-Congolian forest, the Guineo-Congolian Regional transition zone
forest and Guineo-Congolian bushland and thicket, sensu White (1983). Unlike the other three
species of Griffonia, which are restricted to forests, G. simplicifolia is less habitat specific, occurring
in savanna, on coastal plains (particularly termite mounds), scrub thickets, in secondary and
disturbed forest, including roadsides and primary forest margins (Addo-Fordjour, et al., 2009a;
2013, 2015; Hermann, 2011; Figure 2A). In West African dry coastal forest of the Guinea-
Congolia/Sudania regional transition zone, G. simplicifolia is more common in the east, on
inselbergs of the Accra Plains, where G. simplicifolia and Premna quadrifolia Schumach. & Thonn.
are the most common lianas in forests dominated by Diospyros abyssinica (Hiern.) F. White and
Millettia thonningii (Schum. & Thonn.) Baker.
[insert Figure 2. A. Griffonia simplicifolia in Ghana, showing the typical shorter growth form outside of forests,
where natural “tree trellises” are absent. B. Griffonia pods high in tree on the forest margin; C.
Demonstration of the “traditional” method of felling Griffonia stems with a machete to get the pods. D. Detail
showing non-destructive harvest using a hooked stick; E. Tall G. simplicifolia on a forest margin showing the
method advocated in Ghana and Liberia for harvest of G. simplicifolia pods using a hooked stick rather than a
machete.

3.3 Utilization and trade

3.3.1 Uses of G. simplicifolia in West Africa

The small-scale uses of G. simplicifolia across West Africa, include traditional medicinal uses, use of
the leaves for insecticides and arachnicides, fodder (including the fruit/pods) and for dyes, stains,
inks, tattoos and mordants (the stems, roots and fruits) as well as chewing-sticks (Burkill, 1985) are
summarised in Table 1. For the most part, however, G. simplicifolia seeds are wild-harvested
expressly for the export trade, mainly to China and India for extraction of 5-HTP (Figures 1B and 2,
Section 3.2.2).
Table 1 lists the countries where wild G. simplicifolia occurs along with information on whether it is
commercially harvested in each country from cultivated or wild population sources. The harvest
areas and methods are provided in cases where data is available.

Table 1. A summary of Griffonia simplicifolia sources and uses across the geographic range of
this species, noting that wild populations are the sole source of supply for all countries.
Range
Source Trends in use
State
Benin Wild Oral hygiene (Hounnankpon, et al., 2017).
Cameroon Wild No Cameroon-specific information found.
In Côte d’Ivoire (and Nigeria) “The pulped bark is applied to
syphilitic sores. A leaf decoction is used as an emetic, cough
medicine and aphrodisiac. A decoction of stems and leaves is
taken as a purgative to treat constipation and is used externally
Côte
Wild as an antiseptic wash to treat suppurating wounds. Leaf sap is
d’Ivoire
used as eye drops to cure inflamed eyes and is drunk or applied
as an enema to cure kidney problems. Stems and stem bark are
made into a paste that is applied to decaying teeth, and a paste
made from the leaves is applied to burns” (Bosch, 2008).
Gabon Wild No Gabon-specific information found.
Raw “Kagya” seeds are taken orally for treating stomach ache
(Appiah, et al., 2019). “Kagya” leaves are traditionally used in
the treatment of boils, fresh wounds, sores in the head and
insect stings (CSIR-Forestry Research Institute of Ghana, 2020).
The leaves are also used to make a “substantive dye” (a direct
Ghana Wild
dye that does not require mordants) for cotton fabrics yielding
a brown colour (Opoku-Asare, et al., 2013). The stem is
traditionally used as chewing stick, fuelwood and for rope
making, and the leaves for fodder for sheep and goats (UNDP,
2010).
Liberia Wild No Liberia-specific information found.
In Nigeria, stems are chewed by men like chewing stick as
Nigeria Wild aphrodisiac; powdered leaves mixed with pap (Nigerian corn
meal) to treat male infertility (Borokini, et al., 2013).
In Togo, decoction of the leafy stems of “Egbodouti” is taken
for treatment of malaria anaemia (Klotoé, et al., 2018). The
Togo Wild leaves are eaten like vegetables, while various other plant parts
are used by traditional healers for the manufacture of
traditional medicines (Novidzro, et al., 2019).

3.3.2 International export trade

Although wild collection of the leaves and other plant parts occurs for use in local traditional
medicines, the wild collection of G. simplicifolia seed in West Africa is primarily for export (Figure 3).
Value-addition in exporting countries (such as Côte d’Ivoire, Ghana and Liberia) is limited to
harvesters and buying and packing agents cleaning, drying, sorting, grading or packaging the seeds
prior to export.
Figure 3. From forest to pharmacy: A. Shelled seeds and empty pods (plus part of a cut stem) on top of a
banana leaf during sorting in a local village. B. C. A Griffonia seed trader in Zorgowee (Nimba county, Liberia),
who was member of BOTPAL, a trader network started by the PROSPER project, which aimed at improving
drying, packing, seed quality and prices along Griffonia value chains. D. G. simplicifolia capsules produced in
Austria.

The dried raw material (seed or powdered seed) is exported to high-technology extraction houses
situated mainly in China, followed by India and some European countries (Figure 4). The seeds are
harvested within a short window annually from December to February and shipped via containers to
China/India/Europe. The shipment could take up to 2-4 months to reach extractors in Asia, which can
affect 5-HTP levels. After processing it may take until the summer months before the value-added
extract is re-exported to the 5-HTP market in the USA. Consequently, the demand vs. supply time-
lines are misaligned, causing fluctuations in prices of both seeds and 5-HTP. When inventories are
low, demand results in increases in the price of 5-HTP extracts. Multiple extractors then hurry to buy
seeds in the next harvest season, exerting greater pressures on wild stocks of G. simplicifolia seed
(Kodzo Gbewonyo, pers. comm., 15.2.2020 and 22.11.2020).

Figure 4. A basic long distance supply chain for wild harvested G. simplicifolia seeds, including the cross-
border trade from Cote d’Ivoire, Liberia and Togo to Ghana. In the early 1980’s, most seed was exported from
West Africa (Cote d’Ivoire and Ghana) to Europe (particularly Germany). By the early 2000’s, most seed was
exported to China for extraction of 5-HTP and re-export of the extract or of capsules and other products, most
(c.48%) to the USA. Juliani (2013) records additional details for Liberia, including transport times and costs
(US$2.10 – 5.70 per “rice bag” of G. simplicifolia seed) depending on distance, with the highest costs from
collection areas over 70km away. The excellent poster by Acquaye et al. (2014) illustrates the detailed “do’s
and don’ts” of good collection practices very well and was useful in informing this supply chain diagram.

Precisely quantifying the international export trade of Griffonia seed raw material, and extracts
made from it, was not possible due to the absence of species-specific HS Codes. We therefore
evaluated data provided by private companies, NGOs involved in sustainable Griffonia production,
and online B2B trade databases. Because the US is a major importer of Griffonia extracts from
extraction houses situated in China and India, we purchased USA import trade data from an Indian
private trade analysis firm Zauba Technologies Pvt. Ltd. The shipments data obtained from Zauba
could serve as a proxy, representative of the overall export trade from Asia with regard to types of
ingredients produced, information on the main processors and exporters, and the main US
importers. These data filled in the gaps for the African export trade of Griffonia raw materials. We
could weave together, through the various data sets, that the main Chinese exporters of extracts to
the USA were the main importers of raw material from western African countries. This section on
international trade firstly provides country-specific details on exports of Griffonia seed from Africa
and secondly, summarizes the international trade scenario based, in part, on information from
Alibaba, ASNAPP, and Zauba, as well as information providers in the Griffonia trade.

We did not find evidence of export trade of G. simplicifolia seeds from Benin or Gabon. And
although the report to FAO by Nigeria’s National Centre for Genetic Resources and Biotechnology
(2008) lists G. simplicifolia as an “endangered” species collected by the National Centre for Genetic
Resources and Biotechnology (NACGRAB), no mention is made of exports. Researchers at the
Department of Botany, University of Lagos refer to G. simplicifolia as a rare endemic medicinal plant
species of Nigeria, and have begun cultivation trials (Balogun, et al., 2020). A legal export trade is
recorded from:

Cameroon: The Exporters India searchable B2B online database (https://www.exportersindia.com/)

listed one Cameroonian company, Loum Agric Exports, exporting seeds of G. simplicifolia to India (at
US$800 – 900 per tonne in minimum quantities of 2 tonnes). However, more information is not
available on exports from Cameroon.

Côte d’Ivoire: For over 30 years, seeds of G. simplicifolia have been exported from Côte d’Ivoire. In
1990, Professor Ake-Assi (pers. Comm. to A.B.C) spoke about the destructive harvest caused by
felling of G. simplicifolia stems when pods were collected for seed exports to Germany
(Cunningham, 1993). Today, Côte d’Ivoire, along with Ghana, is one of the two main exporters of G.
simplicifolia seed, which, according to the Africa Export Forum B2B online database
(http://www.export-forum.com/africa.html) is collected in central Côte d’Ivoire, including in the
Daoukro area. Dried seeds are packed in 60 kg jute bags. Minimum batches exported from the
Abidjan port commonly are 500kg at a time. However, the export trade destination has shifted from
Europe in the 1980’s (mainly Germany) to China and India today. For example, as listed in the
Exporters India B2B database, the Olakila Import and Export Company Ltd., based in Hong Kong has
a branch office in Côte d’Ivoire to facilitate trade in seeds of G. simplicifolia and other medicinals
such as Voacanga africana Stapf ex Scott-Elliott (Apocynaceae) seed, Picralima nitida Stapf (T.
Durand & H. Durand) (Apocynaceae) seed and Rauvolfia vomitoria Afzel. (Apocynaceae) root and
rhizome, and spices such as Guinea pepper (Xylopia aethiopica (Dunal.) A. Rich, Annonaceae),
alligator pepper (Aframomum daniellii (Hook.f) K. Schum., A. citratum (C. Pereira) K. Schum. or A.
exscapum (Sims) Hepper, Zingiberaceae) seeds. In addition to the aforementioned Olakila company,
the Alibaba online B2B portal lists other Griffonia seed exporters including Sarepta Enterprises
(minimum order quantity of 12 MT), Ivoire Negoce (min. order 100 MT), and Cooperative
Agricultural Toudougou (min. order 500 MT).

Ghana: Seeds of G. simplicifolia and Voacanga africana are the two main Ghanaian medicinal
products with export value. Based on internet-based surveys of e-commerce and local information,
at least seventeen companies are involved in G. simplicifolia exports from Ghana (Afodak Company
Ltd.. Africa First LLC, Africa Naturals Co. Ltd., Akowilson Trading and Farms Ltd., Bethel Incorporated
Agencies, Bioresources International Inc., Bride Tidings Enterprise Ltd., Chalbedico Co. Ltd.,
Gbagfarms Ltd., Green Extract Ghana, Herrnhut Commodities, IRECO, Koafric Ventures, Norwek
Trading and Farms Ltd., Ofosael Ghana Limited, Spearson Group, Ultimate Imports and Exports). In
2003, a representative of the (former) Tek-Spec. Co. Ltd., located in Ghana, informed one of us (JB)
that his company could supply up to 100 MT of G. simplicifolia seeds annually, with a harvest
window of December through March (Brinckmann, 2003). In a 2008 study on wild collected
medicinal plants carried out by one of us (JB) for WWF Deutschland, one of the surveyed Ghanaian
companies (Spearson Ltd., Ashanti Region) reported trading about 150 MT of wild collected
Griffonia annually. These estimates represent individual companies and not the total wild-collected
quantity from all of Ghana (Brinckmann & Goldberg, 2008). By 2007, based on a survey of Ghanaian
traders carried out by ASNAPP (Agribusiness in Sustainable Natural African Plant Products), Ghana
exported 700 MT of G. simplicifolia seeds, with exports dropping to 450 MT the following year,
peaking at 900 MT exported in 2010, and 600 MT of seeds exported in 2013 (Kodzo Gbewonyo,
BioResources International Inc., Accra, 15.2.2020, pers. comm.). Interestingly, Juliani (2013) stated
that Ghana exports 2000 MT of G. simplicifolia seeds per year, 30-40% of which is derived from
cross-border trade from Côte d’Ivoire and Liberia. The United Nations Development Programme
(UNDP) in Ghana had estimated Ghanaian annual production of G. simplicifolia seeds in 2010 at
about 300 tonnes, one-third of the ASNAPP export estimate (UNDP, 2010). Another source reported
that about 120 tonnes of the 2010 Ghanaian export total went to Western Europe (Chris Okunji,
Global Phytotherapeutics Consulting LLC, Accra, 17.2.2020, pers. comm.).

Table 2 shows estimated annual quantities of Griffonia seeds exported from Ghana, 2005 through
2013 in comparison to total medicinal plant exports. More recent export data are not currently
available.

Table 2. The total quantity and value of medicinal plants exported from Ghana (2005-2019)
compared to estimated quantities of G. simplicifolia seed harvested and shipped or exported from
Ghana. The total exports data from was kindly obtained from the Ghana Export Authority in 2020 by
ASNAPP (Daniel Acquaye, pers. comm., 2020). The Griffonia export data is based on ASNAPP data
compiled from 2005-2013 by ASNAPP (Ghana) from interviews with key exporters of Griffonia in
Ghana, so probably represents an underestimate of total Griffonia exports, as Juliani (2013) reported
that Ghana exported 2000 tonnes per year, 30-40% of which came from Liberia or Cote d’Ivoire. No
disaggregated data for Griffonia was available for the period 2014-2019.

YEAR TOTAL MEDICINAL TOTAL VALUE OF GRIFFONIA

PLANT EXPORTS MEDICINAL PLANT EXPORTS FROM
(metric tonnes) EXPORTS (US$) KEY EXPORTERS
2005 1111.192 2 461 984 700
2006 2041.133 2 080 704 450
2007 4916.142 4 066 879 600
2008 1421.900 2 408 905 700
2009 4854.123 8 147 137 400
2010 10904.265 21 126 397 900
2011 21261.098 16 580 440 700
2012 22391.413 14 198 430 500
2013 54048.906 43 116 561 600
2014 68337.679 51 991 308
2015 35527.540 28 995 008
2016 32303.243 26 512 063
2017 8989.405 9 605 712
2018 7986.951 9 432 808
2019 3353.212 5 726 419
Most G. simplicifolia seed exports from Ghana are shipped to China, with lower quantities exported
to India and Europe for value-addition. Griffonia simplicifolia seed extracts are not yet produced in
Ghana at a commercial scale. They are, for the most part, produced at extraction houses situated in
China, India and in some European countries (Richard, E., Africa Naturals, Accra, Ghana, 11.2.2020,
pers. comm.). The extraction houses then export the bulk ingredients (extract or purified substance)
to ingredient distributors or finished product brands mainly in the Americas and Europe.

In most of the destination countries, the ingredients are handled and regulated as active ingredients
of herbal medicinal products (e.g. Austria, Canada) although some countries treat the ingredients as
dietary supplement components (e.g. the Netherlands, United States). According to the trade
database of Zauba Technologies Pvt Ltd, Griffonia extracts and/or purified 5-HTP produced in India
are exported mainly to customers in the United States, followed by the European countries Latvia,
Czechia, Belgium, Austria, United Kingdom and France (see: https://www.zauba.com/export-
Griffonia-hs-code.html).

Liberia: In 2010, 750 kg of G. simplicifolia seeds were collected and sold. In 2011, the annual total
increased significantly to 10 tonnes collected and sold (Simon, 2012). Three Liberian counties
(Nimba, Bassa, and Lofa) are most important sources of G. simplicifolia (Hwang, 2017). This has
increased significantly, due to cross border trade to Côte d’Ivoire and Ghana, who re-export the
dried, graded seeds. Hwang (2017) for example states that the Chinese company BannerBio
Nutraceuticals, Inc., who purchased G. simplicifolia seeds in Liberia processed 200 MT of seeds per
year.

Togo: Exports of G. simplicifolia seeds from Togo are advertised by HIB & Co., in Lomé, Togo and by
the French company, Latitudes 230 SARL, based in Rouffiac-Tolosan in south-west France. The latter
company has developed G. simplicifolia seed supply chains from seven villages in Togo and claims to
follow Fairtrade principles (see: https://www.latitudes23.com/en/produits/griffonia/).

Based on the Alibaba online B2B database, it is clear that the main exporting countries for the raw
material are Ghana and Cote d'Ivoire. Trading companies in some non-African countries are shown
as re-exporters of the dried seeds, e.g. United Kingdom. Based on the online databases of both
Alibaba and Zauba Technologies, it is also clear that the main exporting country for both Griffonia
seed extract and purified 5-HTP is China. Using the search term “Griffonia” in the Alibaba portal,
there were 1465 results. Each result is not unique however as some companies have multiple
product listings within the 1465 count. Furthermore the results include all processed forms
including natural extracts of the dried seed, semi-purified and purified substances derived from
Griffonia seed (5-HTP), bulk capsules and tablets for filling into bottles, and finished herbal
products, labelled and packed for retail sale. While the vast majority of the 1465 results correspond
to export-oriented Chinese extraction houses, there were five distinct Indian extraction houses
included, as well as some ingredient trading companies from South Africa and the USA.

Concerning US imports of ingredients or products of G. simplicifolia seeds, according to trade data

compiled by Zauba Technologies, 26 shipments arrived at US ports in the calendar year 2019. Of
these, 23 shipments were loaded at a Chinese port. The remaining three shipments were a small re-
export of 100 kg of Griffonia seeds from port of Antwerp, the Netherlands, 125 kg of Griffonia seed
extract from port of Sydney, Australia, and 550 kg of Griffonia seed extract from the port of Pusan,
South Korea. In total, the 26 shipments weighed 123,555 kg. From these data it is, however, not
possible to quantify the portions that represent Griffonia extracts. That is because many of the
shipments included other botanical ingredients, which therefore contribute to the total weights and
quantities shown. The higher weight Griffonia-containing shipments included extracts of other
botanicals and other natural ingredients. To better understand the Griffonia trade and therapeutic
use in destination countries it is illustrative to note what other types of natural ingredients the
Chinese extractions houses are specializing in, along with concentrated extracts of Griffonia seed
and/or purified 5-HTP. In addition to containing Griffonia seed extract, a 21,036 kg container
shipped from AHA International Co. Ltd. (Hefei City, Anhui Province) also contained extracts of
horny goat weed (Epimedium spp., Berberidaceae) aerial parts, L-theanine (>98% pure), maca
(Lepidium meyenii Walp., Brassicaceae) hypocotyle, stinging nettle (Urtica dioica L., Urticaceae)
root, tongkat ali (Eurycoma longifolia Jack, Simaroubaceae) tree root. Another Chinese extraction
company exported extracts of Griffonia seed and yohimbe (Pausinystalia johimbe (K. Schum.) Pierre
ex. Beille, Rubiaceae) tree bark. What all of these extracts have in common is that they are used as
components of dietary supplement products targeted to men, for erectile disfunction, enlarged
prostate and low testosterone levels. These US import data show that a frequent Chinese exporter
of Griffonia seed extract, BannerBio Nutraceuticals Inc. (Shenzhen, Guangdong) exports product
mainly to their own US distributor situated in California, BannerBio USA, Inc.. BannerBio produces a
branded Griffonia seed extract they call GriffsTract® available in three specifications: (1) purified dry
extract powder standardized to contain 99% 5-HTP; (2) customized quantified dry extract
standardized to contain anywhere from 10 to 99% 5-HTP; and (3) semi-purified dry extract granules
standardized to contain 95% 5-HTP. In this case, it is clear that some ingredients represented as
Griffonia seed extract may be purified 5-HTP, ranging from 95% to 99% purity. According to the
manufacturer, 5-HTP helps improve mood, depression, anxiety, insomnia and migraines (BannerBio
Nutraceuticals Inc., 2020).

In Canada, the chemical substance 5-HTP (5-Hydroxy-L-tryptophan) extracted from the seeds of G.
simplicifolia is classified as an active ingredient of licensed Natural Health Products (NHPs), requiring
pre-marketing authorization. Licensed 5-HTP NHPs may be marketed with indications for use
including “Helps to promote healthy mood balance,” “Helps to relieve symptoms of fibromyalgia,”
“Helps to reduce the severity and duration of migraine headaches when taken as a preventative
(prophylactic),” “To be used with a program of reduced intake of dietary calories and increased
physical activity (if possible) to help in weight management by reducing carbohydrate cravings,” and
“Used as a sleep aid” (Natural and Non-prescription Health Products Directorate, 2018). 5-HTP is
also included as an active ingredient in Canada’s “Cognitive Function Products” monograph (Natural
and Non-prescription Health Products Directorate, 2019). There are licensed NHPs in the Canadian
market containing extracts of G. simplicifolia seed standardized to contain 10% or 20% of 5-HTP as
well as NHPs containing isolated 5-HTP as an active ingredient (Natural and Non-prescription Health
Products Directorate, 2020).

In the European Union, G. simplicifolia seed extract is authorized for use as a skin-conditioning
component of cosmetic products (European Commission, 2020). In the United States, extracts of G.
simplicifolia seed, standardized to levels of 5-HTP as well as isolated 5-HTP are used as components
of dietary supplement products.

Interestingly, Goraya & Ved (2017) report that a market for the use of imported G. simplicifolia
seeds by rural householders in India has also developed with an estimated annual demand of less
than 10 tonnes. They do not elaborate on why rural householders in India are purchasing imported
G. simplicifolia seeds. Another source reports that G. simplicifolia has been domesticated and
planted in the area of Darbhanga, Bihar, India, listing it among the “Important Medicinal plants of
Darbhanga” (Jyotsna & Kumar, 2015).

3.4 Seed and leaf chemistry

Unlike many species in the Fabaceae that fix nitrogen through a symbiotic relationship with soil
bacteria, G. simplicifolia is not considered to fix atmospheric nitrogen (Tropical Plants Database,
2020). As the 5-HTP and 5-HT levels in G. simplicifolia leaves are low compared to the seeds (Fellows
& Bell, 1970), the leaves and twigs are not only eaten by livestock (Barnes, 1998), but also wildlife,
including elephants (Theuerkauf, et al., 2000) and larvae of six Charaxes butterfly species (see:
www.nhm.ac.uk). Interestingly, 5-HTP levels in leaves are high during the early part of the fruiting
season and 5-HT levels are high in leaves at the end of the fruiting season (Figure 5).
[Insert Figure 5. The seasonal variation in the indole amino-acids (including 5-HTP and 5-HT) content of
mature G. simplicifolia leaves in Ghana (redrawn from Fellows & Bell, 1970)]

In contrast to low indole amino-acids in the leaves, there are very high levels of 5-HTP in G.
simplicifolia seeds, with 5-HTP of 14% of seed mass (Bell, 1976) or as high as ~20% (w/w) (Lemaire &
Adosraku, 2002). In a comparison of 5-HTP levels in 10 G. simplicifolia populations from Ghana and
Liberia, Giurleo et al (in press) found similar 5-HTP levels (from 110.23 mg/g to137.04 mg/g).
Although minute quantities of 5-HTP are found in seeds in other plant families, these high 5-HTP
levels are unique to the genus Griffonia (Fellows & Bell, 1970). Janzen (1980) suggested chemical
protection of seeds against beetle predation by 5-HTP, which is supported by his earlier
experimental study that showed that 1% 5-HTP was effective against the predatory cowpea beetle
Callosobruchus maculatus Fab. and a 5% 5-HTP was lethal (Janzen, et al., 1977). However, Asiamah
(2003) found that fresh G. simplicifolia seeds collected at different sites in Ghana contained 14.18 -
18.24% w/w of 5-HTP seed weight, but that 5-HTP levels of harvested seeds decreased at a rate of
4.62% w/w per month when stored under tropical conditions. This reduction in 5-HTP levels is likely
to increase the vulnerability of unharvested seed to fungal and insect attack.

3.5 Challenges of complex interactive effects on G. simplicifolia populations

On its own, commercial harvesting of seeds from wild plant populations can be sustainable. In
Africa, for example, matrix population modelling of marula (Sclerocarya birrea (A. Rich.) Hochst.)
showed that 92% of fruits/seeds could be harvested without long-term population declines
(Emanuel, et al., 2005). And in South Africa, without browsing by livestock, 90% of baobab
(Adansonia digitata L.) fruits could be harvested (or 33% of fruits with moderate livestock numbers)
without population decline (Venter & Witkowski, 2013).

Although Kew Science’s Plants of the World Online classifies the conservation status of G.
simplicifolia as not threatened (POWO, 2019), other sources report the species as threatened in
Benin (Neuenschwander & Adomou, 2017) or endangered (EN) according to IUCN criteria (Adamou,
2005). Why be concerned at all about G. simplicifolia populations with this species being a vigorous
re-sprouter that is less habitat specific than the three other species in the genus? After all,
Hawthorne (1993), based on extensive field experience in Ghana, ranked G. simplicifolia as a "Green
Star" species (i.e: a widely distributed species not of conservation concern), later upgrading this
species to “Blue Star” status, meaning that G. simplicifolia is intermediate in terms of rarity (W.
Hawthorne, pers. comm., March 2020) (and see:
https://herbaria.plants.ox.ac.uk/bol/oxford/bioquality).

We suggest that the situation has changed in the almost 30 years since Hawthorne's (1993)
assessment. And that the interactive effects of multiple forms of disturbance summed up below and
in Figure 7 need to be taken into account. There are six interacting reasons why the long-term
future of wild G. simplicifolia populations should be an issue of conservation concern:

 Rapid loss of forest habitats: Forest loss in Ghana, which is the main source country for G.
simplicifolia, is one of the highest rates of forest loss globally. Decades after G. simplicifolia
 exports started, Ghana was losing between 1.2-2% of forest per year (Benhin & Barbier, 2001;
Hansen, et al., 2009). A more recent study shows that even forests within Ghana's Forest
Reserves are in decline at a similar rate (of 1.1 to 2% per year) due to clearing for tree crops,
agricultural crops or settlements (Acheampong, et al., 2019). In addition to habitat loss and
disturbance, the fires that accompany forest conversion to farmland also thin out G.
simplicifolia densities. In Cote d'Ivoire, for example, de Rouw (1993) found that while forest
plants (of a range of species, including G. simplicifolia) resprouted after a mild initial burning
(5.8 plants/m2), a burn of normal intensity reduced plant densities (to 1.4 plants/m2). Over a
29-year period, dense forest cover in the areas studied by Acheampong, et al. (2019) was
reduced by 33.2% and the deforestation rate is likely to increase in the future (Koranteng et
al., 2016). Forest loss and fragmentation are also high in other West African countries where
G. simplicifolia occurs (Bowker et al., 2016; Hansen et al., 2020). Although G. simplicifolia
resprouts and can be abundant along forest margins (Ofosu-Bambo, et al., 2019), population
numbers are affected by human disturbance. For example, in Atiwa range forest reserve,
Ghana, where G. simplicifolia was the most abundant liana, its relative density (RD) and
Importance Value Index (IVI) declined from an RD of 7.44 and IVI of 12.50 in undisturbed
forest to an RD of 0.68 and IVI of 2.02 in heavily disturbed forest (Kpontsu, 2011).

 Plant-pollinator disruption: In the case of G. simplicifolia, understanding plant–pollinator

relationships is vital for identifying plant species at risk of reproductive failure. We suggest
that like many Bauhinia species (which are also in the Cercideae), G. simplicifolia is largely
self-incompatible. Pollination by vertebrates is relatively uncommon in angiosperms.
Pollination by birds or bats, for example, only occurs in 3–11% of species of lowland tropical
forests (Devy & Davidar, 2003; Fleming, et al., 2009). Both bird and bat pollination are
widespread in Caesalpinioideae, and birds and bats are the main pollinators of the ancient
tribe Cercideae to which G. simplicifolia belongs (Banks & Rudall, 2016). Arroyo (1981) lists G.
speciosa as bird pollinated and it is highly likely that this also applies to G. simplicifolia. At a
global scale, three bird families (Trochilidae (hummingbirds), Nectariniidae (sunbirds) and
Meliphagidae (honey-eaters) are the main specialist flower pollinators (Proctor, et al., 1996).
In West Africa, G. simplicifolia flowers from July till November and the pods ripen from
August onwards. We suggest that sunbirds, weavers and possibly barbets would be the most
likely pollinators of G. simplicifolia. In Ghana, for example, a quantitative study showed that
populations of Olive sunbirds, the most common species (and most likely pollinator), had
declined 53% between 1995-2015 (Arcilla, et al., 2015). Habitat loss and illegal logging affects
several of the bird populations (such as sunbirds) that pollinate G. simplicifolia flowers (Figure
6). Given the similarities in flower form of G. simplicifolia (formerly Schotia simplicifolia),
additional insights can be gained from bird pollination studies of Schotia brachypetala Sond.
by Symes & Yoganand (2013), (Figure 6).
 Figure 6. A. Mature G. simplicifolia pod from a plant growing in an cleared of forest. Although G.
simplicifolia pods can contain up to four seeds, seed number/pod is reduced by pollination
failure. B. Griffonia (formerly Schotia) simplicifolia flowers, typical of a bird pollinated species. C.
A Yellow rumped tinker barbet attracted to Schotia brachypetala flowers; D. A Collared sunbird
pollinating flowers of Schotia brachypetala, which is in the same legume sub-family as G.
simplicifolia. As understorey bird abundance has declined by >50% in Ghanaian forests (Arcilla
et al., 2015), we hypothesise that this decline may be causing pollination failure and reduced
yields of G. simplicifolia seeds.

The negative effects of introduced invasive species (Broussonetia papyrifera (L.) L'Hér. ex
Vent., Chromolaena odorata L. R.M. King & H. Rob.) which create deep shade and in the case
of C. odorata, the likelihood of allelopathic effects that may inhibit G. simplicifolia seed
germination, which is know from other species (Rusdy and Ako, 2017). Both shading and
allelopathy are likely to reduce regeneration of G. simplicifolia in remaining forests (Addo-
Fordjour, et al., 2009b; De Rouw, 1993);
 Grazing by livestock or lopping for fodder, as G. simplicifolia is palatable (Barnes, 1998;
Bosch, 2008), reducing future seed production;

 Commercial scale G. simplicifolia seed collection: For the past 50 years, billions of seeds have
been exploited from wild G. simplicifolia populations. Considering the biological
characteristics of this species (with a low number of seeds per pod (1-4 seeds) and short
distance (ballistic) seed dispersal from a relatively short (3 m high) species), this seed off-take
may affect future regeneration. People certainly are now the main "seed predators". To put
what plant population biologists would term "seed predation" (in this case, by people) into
perspective, an ASNAPP (Agribusiness in Sustainable Natural African Plant Products) survey of
key exporters in Ghana showed that from 2005 to 2013, G. simplicifolia exports totalled 5550
metric tonnes (or between 9.1 billion to 13.5 billion seeds) over a 9-year period. Based on a
seed mass of where 100 seeds weighed between 41g - 61g (or 0.41 - 0.61g per seed), this
represents the loss of 616.67 tonnes of seeds per year (or 1 - 1.5 billion seeds/year) from the
wild populations in Ghana. On the basis of available trade data, we suggest that total G.
simplicifolia seed exports for West Africa could be 1000 – 2000 tonnes/year, as Juliani (2013)
suggested that Ghana exports 2000 tonnes/year, of which 30-40% came from neighbouring
 countries such as Cote d’Ivoire and Liberia. Over an extended period, this scale of seed
exploitation is likely to impact recruitment of seedlings and the sustainability of wild
populations, particularly if other factors are taken into account (Figures 6 and 7). Although
many studies have been done in Ghana on Relative Densities (RD) and Importance Value
Indices (IVI) of G. simplicifolia (Addo-Fordjour, et al., 2009a; 2013b, 2015 b; Kpontsu, 2011),
with Hermann (2011) counting seedling numbers in Nkrabia Forest Reserve, Ghana (where
there were 5008.8 G. simplicifolia stems/hectare inside the forest), we are not aware of
studies carried out on current population structure (age/size structure) in West Africa, or on
changes to G. simplicifolia population structure.

 Destructive harvest methods: Sustaining this scale commercial harvest is also influenced by
the fact that seed collectors in Cote d'Ivoire (Ake-Assi, pers. comm., 1989), Ghana (Bosch,
2008) and Liberia (Hwang, 2017; Hwang, et al., 2018) are commonly known to chop down G.
simplicifolia stems to collect the pods. Griffonia simplicifolia is a vigorous resprouter, so the
chopping of stems does not kill the plants. But we suggest that it may have a "ripple effect"
for two reasons. Firstly, stem regrowth after cutting delays time to reproductive maturity,
reducing seed (or fruit) production (see Hwang, 2017; Hwang, et al., 2018). Secondly, the
resulting lower seed (or fruit) production shifts this form of destructive harvest to other sites
affecting more plants over a larger area.

As Mandle, et al. (2015) point out, studies of multiple interactive factors on plant populations are
rare. Such studies are needed in the case of G. simplicifolia, where multiple factors are influencing
this species. In their seminal study of the combined effects of fire, herbivory and harvest on the
Mountain date palm (Phoenix loureiroi Kunth.), a widely distributed, re-sprouting species in South
Asia, Mandle et al (2015) showed the necessity of using integral projection models (IPMs) of plant
population dynamics to better understand interactive impacts and more effectively manage
harvested plant populations. With G. simplicifolia facing the range of interactive impacts
summarized in Figure 7, there is a need for use of integral projection models (IPMs) of plant
population dynamics could help to investigate the effects of seed harvest and habitat disturbance
on G. simplicifolia pollination and breeding systems.

Figure 7. A conceptual model showing biological factors that confer resilience on wild G. simplicifolia
populations (blue arrows) and multiple interactive factors that collectively could be driving long-term decline
in G. simplicifolia across Range States where large-scale commercial seed exploitation occurs (white arrows).
In general, links between biodiversity factors (forest loss, bird pollination) and sustainable trade in
G. simplicifolia seed exports need to be better understood. For example, in 2012, the PROSPER
programme, identified G. simplicifolia as “the NTFP that showed the most potential for large-scale
commercialization that could be beneficial for forest-dependent households, providing them a
complementary source of income during the dry season” (Tetra Tech ARD, 2017). Yet PROSPER
stopped supporting work on G. simplicifolia for two reasons. Firstly, due to weaknesses in BOTPAL,
the harvest and trade association. And secondly, due to their belief “that Griffonia is found only in
secondary forests with tenuous link to biodiversity objectives” (Tetra Tech ARD, 2017). This was
unfortunate because:

(i) G. simplicifolia is not only found in secondary forests. For example, in Atewa Range Forest
Reserve, Ghana, for example, G. simplicifolia was the dominant liana species in the hill-bottom
habitat, followed by Parquetina nigrescens (Afzel.) Bullock, Landolphia dulcis (Sabine ex. G. Don.)
Pinchon and Alafia barteri Oliv. (IVI = 23.5, 21.6, 20.4 and 19.3, respectively). In hill-slope habitat, G.
simplicifolia was the second most important liana species after Motandra guineensis (Thonn.) A. DC.
(IVI = 35.1) (with G. simplicifolia having an IVI of 23.1) and in the plateau habitat, was the most
abundant liana species (G. simplicifolia (IVI =39.6)) after Acacia kamerunensis Gand. (IVI=25.1),
Calycobolus africanus (G. Don.) Heine (IVI = 24.5) and Salacia africana (Willd.) DC. (IVI = 19.1) (Addo-
Fordjour & Rahmad, 2015b). In the Tano Offin and Afram Headwaters Forest Reserves in Ghana, G.
simplicifolia was the most abundant liana species in both edge habitats and the most abundant
species in forest interior habitat (Addo-Fordjour & Owusu-Boadi, 2016);

(i) it is likely that due to bird pollination, G. simplicifolia reproduction from seed can be closely
linked with forest biodiversity and conservation, with massive declines in understory bird
populations a significant issue (see Arcilla et al., 2015);

(iii) from 2012 and 2014, the PROSPER programme, with support from ASNAPP, stimulated the best
practical work on sustainable harvest and improved quality along supply chains (Acquaye et. al.,
2014; Juliani, 2013, Hwang, 2017). Yet this innovative work on G. simplicifolia was stopped while the
project still had three years to run (Tetra Tech ARD, 2017).

We know from quantitative studies in Ghana and Cote d'Ivoire that G. simplicifolia is resilient,
however, so entire populations are not wiped out by conversion of forest to farmland (see Addo-
Fordjour, et al., 2019). In the Atiwa Range Forest Reserve, Ghana, for example, while many liana
species did not survive high levels of disturbance, G. simplicifolia plants survived, although much
smaller numbers (IVI = 2.02) compared to undisturbed forest (IVI = 12.5) and moderately disturbed
forest (IVI = 7.03) (Kpontsu, 2011). In Bobiri forest reserve, Ghana, where G. simplicifolia is one of
the commonest liana species, G. simplicifolia populations were much lower where the forest
reserve habitat was changed by conversion under the Tropical Shelterbelt System of timber
production. Also known as the taungya system, derived from the Burmese words (taung = hill, ya =
cultivation), this system was introduced to Ghana during the colonial period. Land is cleared of
forest, then farmed, while desirable timber species are established. Under this system, G.
simplicifolia had a Relative Density (RD) of 10) compared to Selective Logging (with a G. simplicifolia
Relative Density (RD) of 11.4), with G. simplicifolia recovering after logging (a G. simplicifolia
Relative Density (RD) of 23.6) (Addo-Fordjour, et al., 2009a).

3.6 A future challenge: the economics of 5-HTP from wild harvest vs. metabolic engineering

As Lin et al (2014) point out, the main reason why 5-HTP is commercially produced through
extraction from G. simplicifolia seeds is because it currently is cheaper than synthetic methods.
Declining wild G. simplicifolia seed production from smaller populations (Figure 6) could make wild
harvest less competitive, however, as local villagers go to remoter areas to harvest smaller
quantities of seeds. And costs of wild collection increase. In Liberia, for example, Juliani (2013)
found that the collection, drying and sale of G. simplicifolia seed was only profitable for the local
association (BOTPAL) with a minimum FOB price of US$4.00/kg. Yet the highest price offered to
BOTPAL at that time was only US$2.30/kg. More recently, innovative techniques for microbial
synthesis of 5-HTP are becoming more efficient, with microbial biosynthesis showing promise to 5-
HTP production on a large scale (Lin et al., 2014; Wang et al., 2018). China is not only the major
importer of G. simplicifolia seeds from West Africa for extraction and re-export of 5-HTP. Chinese
scientists are also leaders in biosynthesis of active ingredients found in heavily harvested wild plant
populations such as G. simplicifolia (Lin et al., 2014; Wang et al., 2018) and Rhodiola rosea L. (Bai et
al., 2014). Through the L-tryptophan biosynthetic pathway (L-tryptophan module) and the use of
recombinant Escherichia coli Mig., Wang et al., (2018) could extract 5-HTP that was up to 94.3%
pure with a yield of 40% 5-HTP from biosynthesis. If scaled-up microbial biosynthesis of 5-HTP from
glucose or glycerol results in collapsing exports of G. simplicifolia from West Africa, this would be a
bitter pill for local villagers and African traders to swallow. And a lesson on the importance of
resource management at multiple scales to maintain high yields from wild plant populations.

4.0 Conclusions

As a herbal alternative to Prozac®, the annual bulk G. simplicifolia seed extract market is worth
between US$ 31.7 million (Transparency Market Research (TMR), 2014) and US$ 50 - 100 million,
with demand growing at 7% per year (Yan & Lin, 2015). Meeting this demand requires the “human
predation” of an average of 616.67 tonnes of seeds per year (or 1 - 1.5 billion seeds/year) exported
from Ghana. In addition, rapid deforestation rates in exporting countries in West Africa are of
concern, as are other interacting factors on wild G. simplicifolia population dynamics. If supply
chains from wild stocks are to be sustained, we recommend improved resource management that is
informed by studies of G. simplicifolia pollination ecology, Integral Projection Models (IPM) and
improved monitoring of international trade.

5.0 Acknowledgements

This paper was prepared during the course of a project funded by the German Ministry of the
Environment, Nature Conservation and Nuclear Safety (BMU) and managed by the Federal Agency
for Nature Conservation (BfN) (FKZ 3519532056). Expert advice that contributed to this work was
received from Dr. Uwe Schippmann, former head of the Plant Conservation Division of BfN, Dr.
Kodzo Gbewonyo, President, BioResources International Inc. (Accra, Ghana) who provided the trade
data in Table 2, Dr. Chris Okunji (CEO, Global Phytotherapeutics Consulting LLC (Accra, Ghana)),
Kwabena Fordjour, (retired) Tek-Spec. Co. Ltd. (Accra, Ghana) and Eyiah Richard, Business and
Economic Consultant at Africa Naturals (Accra, Ghana). The photographs in Figures 2A, 5B and C
were provided by Dr. Jörn Germer, Institut für Tropische Agrarwissenschaften (Hans-Ruthenberg-
Institut), Universität Hohenheim (Stuttgart, Germany). Dr K. Yoganand kindly provided the
photographs showing Schotia pollination (Figure 5 C & D), Dr. Rodolpho Juliani the photo of the
Liberian trader (Figure 3C) and Daniel Acquaye (Director, ASNAPP Ghana) gave permission for use of
the photographs in Figure 2C, D & E. Professors Jeremy Midgley (University of Cape Town) and
Professor William Hawthorne are thanked for discussions with A.B-C on G. simplicifolia. Insights
from Professor Laurent Ake-Assi (Jardin Botanique of the Université de Cocody in Abidjan, Côte
d'Ivoire) during discussions there with A.B-C in October 1989 are also gratefully acknowledged.

Authors contributions

A.B. (Tony) Cunningham and J.A. Brinckmann carried out the research and wrote the manuscript,
supported by additional information from D.E.V. Harter.

Conflict of interest statement

The authors declare that there are no conflicts of interest.

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Authors contributions

This study was carried out by A.B (Tony) Cunningham (A.B-C) and Josef Brinckmann (JB) who
wrote the paper with input from David Harter. A.B-C also producing the maps and unless
otherwise acknowledged, took the photographs.

Conflict of interest statement

The authors declare that there are no conflicts of interest.

Figure Click here to download Figure Figure 1 Distribution & exports.jpg
Figure Click here to download Figure Figure 2 Griffonia harvest & res
mgmt.jpg
Figure Click here to download Figure Figure 3 Griffonia seed supply chain West Africa to
Europe.jpg
Figure Click here to download Figure Figure 4 Griffonia immature pod cropped Kodzo.jpg
Figure Click here to download Figure Figure 5 supply chain Griffonia2.jpg
Figure Click here to download Figure Figure 6 Fellows & Bell 1970.jpg
Figure Click here to download Figure Figure 7 Griffonia & Schotia pollination arrow.jpg
Figure Click here to download Figure Figure 8 impacts on GRIFFONIA LR populations.jpg
Table Click here to download Table Table 1 JEP Griffonia.docx
Table 1. A summary of Griffonia simplicifolia sources, harvest methods and trends in use
across the geographic range of this species.
Range
Source Harvest methods
State
Benin Wild
Cameroon Wild
Côte In general, for all Range States,
Wild
d’Ivoire according to Bosch (2008): “For local
medicinal use Griffonia simplicifolia
is harvested in small quantities.
Although harvesting seeds from the
wild is usually fairly sustainable,
there are worrying reports of lianas
being cut down on a large scale to be
able to collect the seeds. Harvesting
Gabon Wild for fodder is best done at intervals of
12 weeks, as total herbage yields are
then considerably higher than when
harvested at 6-week intervals.”
Hwang (2017) quantitatively
documents destructive harvest of G.
simplicifolia by commercial collectors
and recommends methods for
sustainable harvest in Liberia.
Ghana Wild
According to Hwang, et al. (2018), in
Liberia there are reports of local
collectors harvesting “only the pods
by cutting down trees and vines of
Xylopia and Griffonia, respectively, to
collect pods from their branches.
This may be due to the lack of
knowledge of alternative sustainable
Liberia Wild harvesting practices and the benefits
thereof to sustainability and
conservation of the populations.”
Nigeria Wild
Togo Wild
Trends in use
Oral hygiene (Hounnankpon, et al.,
2017).
No Cameroon-specific information
found.
In Côte d’Ivoire (and Nigeria) “The
pulped bark is applied to syphilitic
sores. A leaf decoction is used as an
emetic, cough medicine and
aphrodisiac. A decoction of stems and
leaves is taken as a purgative to treat
constipation and is used externally as
an antiseptic wash to treat
suppurating wounds. Leaf sap is used
as eye drops to cure inflamed eyes and
is drunk or applied as an enema to
cure kidney problems. Stems and stem
bark are made into a paste that is
applied to decaying teeth, and a paste
made from the leaves is applied to
burns” (Bosch, 2008).
No Gabon-specific information found.
Raw “Kagya” seeds are taken orally for
treating stomachache (Appiah, et al.,
2019). “Kagya” leaves are traditionally
used in the treatment of boils, fresh
wounds, sores in the head and insect
stings (CSIR-Forestry Research
Institute of Ghana, 2020). The leaves
are also used to make a “substantive
dye” (a direct dye that does not
require mordants) for cotton fabrics
yielding a brown colour (Opoku-Asare,
et al., 2013). The stem is traditionally
used as chewing stick, fuelwood and
for rope making, and the leaves for
fodder for sheep and goats (UNDP,
2010).
No Liberia-specific information found.
In Nigeria, stems are chewed by men
like chewing stick as aphrodisiac;
powdered leaves mixed with pap
(Nigerian corn meal) to treat male
infertility (Borokini, et al., 2013).
In Togo, decoction of the leafy stems
of “Egbodouti” is taken for treatment
of malaria anaemia (Klotoé, et al.,
2018). The leaves are eaten like
vegetables, while various other plant
parts are used by traditional healers
for the manufacture of traditional
medicines (Novidzro, et al., 2019).
Table Click here to download Table Table 2 Griffonia & Med plant
exports.docx

Table 2. The total quantity and value of medicinal plants exported from Ghana (2005-2019)
compared to estimated quantities of G. simplicifolia seed harvested and shipped or exported from
Ghana. The total exports data from was kindly obtained from the Ghana Export Authority in 2020 by
ASNAPP (Daniel Acquaye, pers. comm., 2020). The Griffonia export data is based on ASNAPP data
compiled from 2005-2013 by ASNAPP (Ghana) from interviews with key exporters of Griffonia in
Ghana, so probably represents an underestimate of total Griffonia exports, as Juliani (2013)
reported that Ghana exported 2000 tonnes per year, 30-40% of which came from Liberia or Cote
d’Ivoire. No disaggregated data for Griffonia was available for the period 2014-2019.

YEAR TOTAL MEDICINAL TOTAL VALUE OF GRIFFONIA

PLANT EXPORTS MEDICINAL PLANT EXPORTS FROM
(metric tonnes) EXPORTS (US$) KEY EXPORTERS
2005 1111.192 2 461 984 700
2006 2041.133 2 080 704 450
2007 4916.142 4 066 879 600
2008 1421.900 2 408 905 700
2009 4854.123 8 147 137 400
2010 10904.265 21 126 397 900
2011 21261.098 16 580 440 700
2012 22391.413 14 198 430 500
2013 54048.906 43 116 561 600
2014 68337.679 51 991 308
2015 35527.540 28 995 008
2016 32303.243 26 512 063
2017 8989.405 9 605 712
2018 7986.951 9 432 808
2019 3353.212 5 726 419