MSc Project by Martijn van Berlo (3282309) In cooperation with Ivan Scotti (EcoFoG) and Hans ter Steege (Naturalis)

UTRECHT

UNIVERSITY

October 7, 2013

Phylogeography of the Neotropical tree species

Eperua falcata (Fabaceae) across the Guianas

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Abstract Eperua falcata (Fabaceae) is arguably the most abundant canopy tree species in the lowland tropical rainforests of the Guianas. Despite this, no study has earlier been conducted to reveal its genetic diversity and dispersal history across the area. To reveal phylogeographic patterns of this species a sampling of leaf material was conducted in Guyana, Suriname and French Guiana. Genetic structure was analysed using two non-coding chloroplast DNA (cpDNA) intergenic spacer regions (trnH–psbA and trnC–ycf6). An AMOVA analysis performed on both markers separately showed fairly strong differences in results from both markers. However, when analyses for both markers were considered together they, combined with a comparison of pairwise FST, pairwise genetic differences and the distribution of haplotypes, revealed a clear pattern of genetic diversity across the Guianas. This pattern suggests that E. falctata originates in or near central Guyana and then dispersed to Suriname and French Guiana. This dispersal appears to have coincided with one or multiple founder events. These migrations have possibly taken place as a result Quaternary glacial and interglacial forest composition changes, contractions and expansions, caused by reduced rainfall patterns. E. falcata may then have benefitted from reduced competition, and dispersed rapidly through the area. Summary for general audience Eperua falcata, commonly named Soft Wallaba, Baboen Walaba and Wapa in Guyana, Suriname and French Guiana respectively, is arguably the most abundant tree species of the Guianas. It is a species that is relatively well adapted to drought and poor soil conditions. Despite the high abundance in the Guianas and widespread use of its timber, no research has earlier been done to reveal its genetic diversity and dispersal history across the area. To answer for this lack of available information, a phylogeographic study of the species was conducted. Phylogeography is the study that combines biogeography, or the spatial and temporal distribution of a species, with its genetics. This can then provide insights into the species’ origins, its historic dispersal patterns, and how populations of this species are genetically linked. This information can then contribute to fundamental knowledge about species evolution and biodiversity. But it can also be used in conservation strategies based on genetic diversity in an ever more threatened tropical rainforest environment. For the purpose of this study, leaf material was collected from individual trees at various locations in all three aforementioned countries. From this leaf material, DNA was extracted and sequenced in order to reveal its genetic code. These genetic codes were then compared, statistically analysed, and put into geographical perspective. The patterns revealed by this study suggest that the species E. falcata originated in or near central Guyana, and then dispersed to Suriname and French Guiana in one or multiple steps. This dispersal pattern may be explained by historic events such as the ice ages. It is thought that during these ice ages, many of the areas now covered by tropical rainforest were then covered by forests of different composition, or more open vegetation types. During these periods, E. falcata, being able to thrive in conditions where other rainforest tree species are unable to grow, may have taken its chance and dispersed to its present geographic range.

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Contents Chapter Page

1. Introduction 3

2. Materials and Methods 4 2.1. Species description 4 2.2. Sampling 5 2.3. Laboratory methods 7 2.4. Analysis 8

3. Results 10 3.1. AMOVA 10 3.2. Population pairwise FST 12 3.3. Population pairwise genetic distance 14 3.4. Haplotype diversity 16 3.5. Haplotype distribution 18

4. Discussion 20 4.1. trnH-psbA 20 4.2. ycf6-trnC 20 4.3. Overall diversity and dispersal 21

5. Recommendations 26

6. Aknowledgements 26

7. References 27

8. Appendix 32

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1. Introduction Neotropical rainforests contain the highest diversity of trees in the world with sometimes nearly 300 species of trees per hectare (Phillips et al., 1994; De Oliveira and Mori, 1999; Gentry 1988; ter Steege et al., 2003; Fine and Ree 2006; Hubbell et al., 2008). Despite the scientific and practical importance of understanding the mechanisms leading to this pattern of biodiversity, and the debate about its origins, relatively few studies have evaluated the factors underlying tree species diversification in the Neotropics (Dick, 2010; Hoorn et al., 2010; Pennington and Dick, 2010). To explain the great diversification in the tropics, several hypotheses have been proposed. But until present day, no general model has been accepted (Willig et al., 2003; Mittelbach et al., 2007). Processes that cause population divergence are also likely to lead to speciation. Therefore phylogeography, the study of historical processes that may have led to the contemporary geographic distribution of individuals, could provide new insights into the evolutionary history and processes of tree species dispersal and formation by combining a species’ biogeography with its genetics. (Avise, 2000; Knowles and Maddison 2002, Hickerson et al., 2010). Phylogenetic changes within and between populations of tree species can result from a number of processes, an important factor of which is spatial isolation by geographic distance and its subsequent disruption of gene flow (Tuomisto et al., 2003). This, combined with the knowledge that even after controlling for geographical distance, lower latitudinal populations show greater genetic differentiation within species, makes it especially interesting to look at species level phylogeography (Eo et al., 2008). Some studies that have investigated the phylogeography of tree species in the Amazon have indicated high levels of genetic divergence between geographically isolated populations (Aide and Rivera, 1998; Hardesty et al., 2010; Lemes et al., 2010; Scotti-Saintagne et al., 2012; Scotti-Saintagne et al., 2013). A phylogeographic approach based on chloroplast DNA (cpDNA) variation can provide information about historical patterns of genetic divergence of tree species across the Guianas. CpDNA is a reasonably slowly mutating, maternally inherited, haploid genome in most angiosperms (Birky, 1995; McCauley 1995). Polymorphisms in the chloroplast genome are therefore likely to reveal major historical events such as dispersal patterns, genetic bottlenecks, etc. This approach has proven to be very effective in recent phylogeographic research on various species in the tropics (e.g. Dauby et al., 2010; Lowe et al., 2010; Scotti-Saintange et al., 2012; Scotti-Saintagne et al., 2013). Earlier research on biogeography and phylogeny of the genus Eperua conducted by Rozendaal (2004) using trnL-F, matK and ITS showed some strong incongruences in the phylogeny of Eperua falcata. She speculated this might be the result of a low resolution of the cpDNA spacer regions used in her study. The inaccuracy of the use of these markers and ITS for intraspecific phylogenetic research was later confirmed by Shaw et al. (2005). The main goal of this study is to examine genetic diversity and dispersal patterns of the Neotropical tree species E. falcata across the Guianas using two cpDNA intergenic spacer regions, trnH-psbA and ycf6-trnC. The study was conducted by investigating the distribution of genetic diversity through a genetic survey at both the population stand level and the regional level across the Guianas. Not only can the results of this study provide insight into the evolutionary diversity of this species, they can also contribute to knowledge about general tree phylogeography in the Guianas. On a broader level they can aid in providing a conceptual framework for developing conservation strategies aiming to protect species and genetic diversity as a whole (Milliken et al., 2010; Moritz, 1994). More sustainable forest management based on population genetics could protect species and biodiversity, and help to ensure the continuation of the evolutionary processes that maintain genetic diversity (Dick, 2010; Moritz and Faith, 1998; Moritz, 2002; Newton et al., 1999). With increasing concerns about climatic changes and the fate of tropical rainforests across the world, this is of particular importance (Hammond et al., 2007; Hubbell et al., 2008).

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2. Materials and methods

2.1. Species description E. falcata is the most common and widely distributed species of the genus Eperua (ter Steege et al., 2013, in press). It is also one of the, or arguably, the most abundant canopy tree species across the Guianas (Johnston and Gillman, 1995; Johnston, 1998; Lescure and Boulet, 1985; ter Steege and Zondervan, 2000; ter Steege et al., 2004; ter Steege et al., 2013, in press). Common names for the species are Soft Wallaba in Guyana, Baboen Walaba in Suriname and Wapa in French Guiana. Individuals of this species can grow to 40 m tall and can reach a trunk diameter of up to 80 cm. It is easily recognisable by its compound paripinnate leaves with clearly falcate leaflets and its large woody falcate pods hanging by long threads (Cowan, 1975; Polak, 1992) (Figure 1). Seed dispersal occurs by exploding seed pods, hurling seeds tens of metres away from the parent tree (Hammond and Brown, 1995). It often occurs in clumped populations on nutrient-poor, well drained white sand soils. E. falcata is often concentrated in so called Wallaba forests where it is one of the dominant tree species (Fanshawe, 1952; ter Steege and Cornelissen 1989; ter Steege et al., 1993). In Addition, it is also positively associated with seasonally flooded forests (Baraloto et al., 2007). The wood of E. falcata is considered to be a useful timber product. The species Eperua falcata is known to contain chemical compounds and relatively large amounts of resin which make it very resistant to insects and fungal decay (Amusant et al., 2009; Vink, 1965). This in combination with its highly concentrated, widespread abundance and lack of logging restrictions make it a cheap, durable and therefore useful timber tree for various construction applications (Guyana Forestry Commission, 2004).

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2.2. Sampling

This study is based on the sampling of 138 individuals of Eperua falcata in Guyana, Suriname and French Guiana from the locations shown in Table 1 and Figure 2. These samples consisted of silica-dried leaf tissue obtained from individual trees in the field. In both Guyana and Suriname, ten samples were taken from each population at random distance intervals and geographic directions, with the only distance criterion being a small likelihood of sampling parent and sibling trees given the dispersal mode of exploding pods. All samples from French-Guiana were provided by Ivan Scotti (EcoFoG).

Figure 1: Schematic drawing of Eperua falcata growth habit (Polak, 1992)

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Table 1: Sampling sites per country

Country Site no. Site name Coordinates (decimal)

Guyana 1

2

3

Mabura Hill

Pibiri Research Site

Iwokrama

5.0183455°, -58.6216908° 5.1827751°, -58.7013732° 4.6111188°, -58.7332415°

Suriname 4

5

6

7

8

Mata

Zanderij

Powakka

Brownsberg

Cropi

5.4500451°, -55.3310629° 5.4192646°, -55.1886232°

5.4443929°, -55.0746241° 4.9510346°, -55.1846924° 5.5164444°, -54.3239021°

French-Guiana 9

10

11

12

13

14

15

16

17

18

19

20

21

22

23

24

25

26

27

28

29

Bafog

Acarouany

Organabo Sable Blanc

La Mana, Embouchure crique Korossibo

La Mana, Saut Lézard

La Mana, Saut Fracas

St. Elie

Paracou

Crique Plomb

PK 72 Est

Montagne Signes

Mont la Trinité-Aimara

Montagne Trésor

Saut Deux Roros, Sinnamary Aval

Nouragues

Piste Belizon

Camp Cisame

Approuague, Saut Mapaou

St. Lavillette

Route de l’est, Autour de la Montagne Couronnée

Oyapok

4.1997532°, -52.3730852° 5.6101451°, -53.7717395° 5.5478200°, -53.4580204° 5.1681207°, -53.6618925° 4.9716509°, -53.7859440° 4.7780598°, -53.6616316° 5.2799965°, -53.0799955° 5.2793627°, -52.9232195° 4.9966886°, -52.9230571° 5.0989672°, -52.7686169° 5.0752500°, -52.7112443° 4.6717018°, -53.2829373° 4.6128835°, -52.2707395° 4.3658290°, -52.8969419° 4.0957057°, -52.6774124° 4.2854495°, -52.3918225° 4.1997532°, -52.3730852° 4.1828611°, -52.3349966° 4.1155287°, -52.2087409° 4.0571106°, -52.0861712° 3.5296950°, -52.0571561°

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2.3. Laboratory methods

All laboratory work was conducted at the CIRAD Campus Agronomique in Kourou, French Guiana. The dried leaf tissue samples were cut into small pieces, flash-frozen in liquid nitrogen and ground to a powder using purpose built racks, tubes and metal pellets in a mill. The cpDNA was extracted using a DNeasy 96 Plant Kit (Qiagen, Valencia, CA, USA). In the cp genome, two intergenic spacer regions, trnH-psbA and ycf6 -trnC (Shaw et al., 2005), were tested for polymorphism within and between populations. The polymerase chain reaction (PCR) followed Duminil et al. (2006), with annealing temperature and elongation time as described in Grivet et al. (1999). Amplifications were carried out in a Perkin Elmer thermal cycler (Perkin Elmer, Waltham, MA, USA), with an initial denaturation step of 4 min at 94°C, followed by 30 cycles [45 s of denaturation at 94°C, 45 s of annealing (temperature dependent on the primer pairs) and 1-4 min of extension at 72°C, (depending on the size of the amplified fragments) ], with a final extension step of 72°C for 10 min. Enzymes and buffers were obtained from Life Technologies, Gibco BRL (Life Technologies, Carlsbad, CA, USA). Sequencing of the samples obtained in Guyana and Suriname was conducted using an Applied Biosystems 3130xl capillary sequencer (Life Technologies, Carlsbad, CA, USA). The samples obtained in French Guiana were sequenced using a 96-capillary automatic sequencer MegaBACE 1000 (GE Healthcare, Madison, WI, USA).

Figure 2: Sampling locations in Guyana, Suriname and French Guiana

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2.4. Data analysis All sequences were aligned and edited using CodonCode Aligner 4.0.4 (Codoncode Corporation, Dedham, MA, USA). Any ambiguities were corrected by hand. Haplotype diversity and locations were determined using the R statistical package. Due to the high proportion of missing data in especially the ycf6-trnC spacer region, the structure of the populations was investigated using a hierarchical approach implemented through locus by locus Analysis of Molecular Variance (AMOVA) (Excoffier et al., 1992) using Arlequin 3.5 (Excoffier et al., 2005). The sequences were entered into Arlequin, where all loci were analysed separately to compensate for missing data. The AMOVA partitions the total variance observed into covariance components that result from the variance among groups, among populations within groups and within populations. Next, the covariance components are used to calculate the following fixation indices, /-statistics: FCT: proportion of genetic variance among groups. FSC: proportion of genetic variance between populations within groups. FST: proportion of genetic variance between groups and populations overall. These are analogous to Wright’s F-statistics (Excoffier et al., 2005), where a value of zero indicates no differentiation or complete panmixis, and a value of one indicates complete genetic differentiation or no gene flow whatsoever. The significance of the fixation indices was tested using a nonparametric permutation approach with 1023 replicates (Excoffier et al., 2005). The populations were grouped according to country, comparable soil conditions and geographical proximity. Very small populations of 2 samples or less were added to the geographically closest group to form a total of 11 groups (Table 2). Table 2: Populations arranged in groups as used in the AMOVA analysis.

Group No. Populations included in group Group No. Populations included in group

1 Pibiri

Mabura Hill

Iwokrama

7 St. Elie

Paracou

Crique Plomb

PK 72 Est

Montagne Signes 2 Zanderij

Powakka

Mata 8 Montagne Trésor

3 Brownsberg 9 Mont la Trinité-Aimara

4 Cropi 10 Saut Deux Roros, Sinnamary Aval

Nouragues

Piste Belizon

Camp Cisame

Approuague, Saut Mapaou

St. Lavillette

Route de l’est, Autour de la Montagne Couronnée

5 Acarouany

Bafog

Organabo Sable Blanc

6 La Mana, Embouchure crique Korossibo

La Mana, Saut Lézard

La Mana, Saut Fracas

11 Oyapok

A matrix of pairwise FST based on Weir and Cockerham (1984) was generated to determine short term genetic distances between populations. The null distribution of pairwise FST values under the hypothesis of no difference between the populations was generated by permuting haplotypes between populations. The P-value of the test is the proportion of permutations leading to a FST value larger or equal to the observed one. This representation makes it possible to quickly discern possible genetic affinities between populations.

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Also a matrix displaying average numbers of pairwise differences (π) between and within sampled populations, as well as Nei’s net (DA) number of nucleotide differences between populations, was generated. DA was calculated as: DA = πXY – (πX + πY) / 2 (Nei and Li, 1979). Since there were too many inconsistencies in the availability of medium to high quality sequences from the two intergenic spacer regions used in this study, for many individuals, one of both markers was unavailable in a sufficiently high quality to be combined. Therefore both markers were analysed separately. Significance for all tests was set at a probability equal to or lower than 0.05. Lastly, from the abundance of different haplotypes identified among samples from every country, a haplotype α-diversity was calculated using an R code for Fisher’s alpha available from the Amazon Tree Diversity Network (2013) in the R statistical package (R Core Team, 2013). The calculated haplotype α-diversity could then be used as a measure for haplotype diversity corrected for sample size.

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3. Results

3.1. AMOVA The locus by locus AMOVA for the trnH-psbA marker yielded highly significant overall population structure among groups, among populations within groups and within populations on all populations, with most of the explained variation occurring among groups. The highest differentiation was found within populations (Table 3). Table 3: Locus by locus Analysis of Molecular Variance (AMOVA) based on the trnH-psbA intergenic spacer region (as a

weighted average over 89 loci).

Source of variation Sum of squares

Variance components

% variation

Among groups 384.779 3.12437 Va 77.43712 Among populations within groups 27.321 0.23251 Vb 5.76264 Within populations 70.275 0.67784 Vc 16.80024 Total 482.375 4.03472

Average F-Statistics over all loci: Fixation Indices FCT : 0.77437 FSC : 0.25540 FST : 0.83200 Significance tests (1023 permutations) (significant if P < 0.05) Va and FCT P(rand. value > obs. value) =

P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

Vb and FSC P(rand. value > obs. value) = P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

Vc and FST P(rand. value < obs. value) = P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

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Sequences obtained from the ycf6-trnC spacer region were generally of lower quality and showed very high numbers of ambiguities and differences in length. In some cases, even the forward and reverse sequences were so different from each other, that they could not be merged and therefore had to be discarded. Note that for this marker no samples of sufficiently high quality from the populations of Acarouany and Nouragues were available for analysis. Locus by locus AMOVA yielded highly significant fixation indices for this marker. However, the fixation indices low values, indicating only minor overall genetic differentiation. Most of the explained variation, as well as the highest differentiation occurred within populations (Table 4). Table 4: Locus by locus Analysis of Molecular Variance (AMOVA) based on the ycf6-trnC intergenic spacer region (as a

weighted average over 850 loci).

Source of variation Sum of squares Variance components

% variation

Among groups 860.320 3.96432 Va 9.12196 Among populations within groups 685.532 1.67693 Vb 3.85864 Within populations 3106.290 37.81779 Vc 87.01940 Total 4652.142 43.45903

Average F-Statistics over all loci: Fixation Indices FCT : 0.09122 FSC : 0.04246 FST : 0.12981 Significance tests (1023 permutations) (significant if P < 0.05) Va and FCT P(rand. value > obs. value) =

P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

Vb and FSC P(rand. value > obs. value) = P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

Vc and FST P(rand. value < obs. value) = P(rand. value = obs. value) = P-value =

0.00000 0.00000 0.00000

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3.2. Population pairwise FST The matrix of population pairwise FST shows that the populations in Guyana were most strongly differentiated from all others. Within Guyana, the population from Mabura Hill showed the highest levels of differentiation, even when compared to other populations from Guyana (Figure 3). Most population comparisons outside of Guyana yielded values lower than or equal to zero. It should be noted that some values above and all values below 0.5 were not statistically significant. All values lower than zero in this matrix were statistically highly insignificant and can therefore be treated as zero (no rejection of H0). (see appendix table 1, page 32, for pairwise FST values and appendix table 2, page 36, for accompanying P-values).

Figure 3: Matrix of pairwise FST based on the trnH-psbA intergenic spacer region. Here only the dark blues for the Pibiri,

Mabura Hill and Iwokrama populations are significant (P < 0.05).

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The matrix of pairwise FST of the ycf6-trnC marker showed that some populations appear to be strongly differentiated from others. While this differentiation was not structured through a certain geographic region, many values are statistically significant (Figure 4). Since pairwise FST values were relative they should normally be between zero and one. All values lower than zero in this matrix were statistically highly insignificant and can therefore be treated as zero (no rejection of H0) (see appendix table 3, page 42, for pairwise FST values and appendix table 4, page 46, for accompanying P-values).

Figure 4: Matrix of pairwise FST based on the yfc6-trnC intergenic spacer region.

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3.3. Population pairwise genetic distance When comparing the average number of pairwise differences, the population that was most different from all others is that of Mabura Hill. Here there is little genetic variation within the population, but it is highly different from all other populations throughout the Guianas. The highest difference within populations was found at the Pibiri Research Site and in the Iwokrama forest. These factors contributed to the high genetic variation within Guyana compared to Suriname and French Guiana (Figure 5) (see appendix table 5, page 52, for all genetic distance values and appendix table 6, page 56, for all accompanying P-values).

Figure 5: Average number of pairwise genetic differences and Nei’s distance based on the trnH-PsbA intergenic spacer

region.

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The analysis of the average number of pairwise genetic differences for the ycf6-trnC marker indicated more variation than the analysis of trnH-psbA (Figure 6). Contrasting to the trnH-psbA marker, here most of the variation within a population was found at Mabura Hill. This population was also most different from all others. Also more variation was found within and between populations in Suriname and French Guiana (see appendix table 7, page 60, for all genetic distance values and appendix table 8, page 64, for all accompanying P-values).

Figure 6: Average number of pairwise genetic differences and Nei’s distance based on the ycf6-trnC intergenic spacer

region.

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3.4. Haplotype diversity In the trnH-psbA marker most of the sampled individuals belonged to the same haplotype. The trnH-psbA intergenic spacer region yielded 33 different haplotypes among the individuals within the sampled populations. Of these haplotypes, haplotype no. 2 (shown in blue) had by far the highest frequency (Figure 7).

Figure 7: The number of different haplotypes of the trnH-psbA marker and their frequency. Haplotype no. 2,

the most common haplotype, is shown in blue.

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The medium to high quality sequences of the ycf6-trnC intergenic spacer region showed more variation among a smaller amount of individuals with a total of 56 different haplotypes among the sampled individuals. Of these different haplotypes again one, haplotype no. 7, had by far the highest frequency (Figure 8).

Figure 8: The number of different haplotypes of the ycf6-trnC marker and their frequency. Haplotype no. 7,

the most common haplotype, is shown in blue.

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3.5. Haplotype distribution The distribution of haplotype diversity for the trnH-psbA marker showed a pattern across the Guianas. Haplotype diversity was highest in Suriname and Guyana, and far lower in French Guiana. In Guyana, a total of 10 different haplotypes among 13 samples was found, yielding a haplotype α-diversity of 19.85559. The haplotype diversity in Suriname was slightly lower with 18 different haplotypes among 34 samples yielding a haplotype α-diversity of 15.50515. In French-Guiana the lowest number of different haplotypes was found with only 7 different haplotypes among 91 samples, yielding a haplotype α-diversity of 1.767417 (Figure 9). The frequency of occurrence of haplotype no. 2 was larger in French Guiana than in Suriname, and this haplotype was totally absent in Guyana.

Figure 9: Haplotype diversity of the trnH-psbA marker. All populations containing individuals of haplotype no. 2 are

shown in blue. For every country N represents the total number of samples, R represents the total number of different

haplotypes, and α represents the haplotype α-diversity.

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For the ycf6-trnC marker, by far the highest haplotype diversity was found in Guyana, with diversity being much lower in French Guiana and lowest in Suriname. In Guyana a total of 15 different haplotypes was found among 16 samples, yielding haplotype α-diversity of 117.4462. In Suriname 16 different haplotypes were found among 37 samples, yielding the lowest haplotype α-diversity of 10.70951. In French-Guiana 27 different haplotypes among a total of 56 samples were found, yielding a somewhat higher haplotype α-diversity of 20.50726. Haplotype no. 7, the most abundant haplotype for this marker, occurred only once in Guyana. Its frequency of occurrence increased in Suriname and was largest in French Guiana (Figure 10).

Figure 10: Haplotype diversity of the ycf6-trnC marker. All populations containing individuals of haplotype no. 7 are

shown in blue. For every country N represents the total number of samples, R represents the total number of different

haplotypes, and α represents haplotype α-diversity.

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4. Discussion

4.1. trnH-psbA For the trnH-psbA marker, the AMOVA analysis yielded strong overall genetic differentiation within and between all populations. A matrix of pairwise FST shows that the strongest differentiation based on marker originates in Guyana, while no statistically significant values are found in populations outside of Guyana. The high FCT value of 0.77437 indicates strong differentiation among the selected groups. This is supported by the matrix of population pairwise FST and the matrix of population pairwise genetic distances. Both matrices indicate that populations from Guyana differ most strongly from all others and show the highest diversity within populations. Quite remarkable is the extremely high FST value of 0.832, indicating individuals overall share only very little genetic diversity. This value however does not explain the largest part of the variation. The fact that very low pairwise FST values and genetic distances were found among almost all populations in Suriname and French Guiana suggests that the mutation rate of this marker is rather low. In fact the mutation rate appears to be too low to infer any further patterns from Suriname down to French Guiana from pairwise FST and pairwise genetic distances from the trnH-psbA marker alone. However the distribution of haplotypes and the α-values calculated from them does reveal a genetic pattern with haplotype diversity being highest in Guyana, lower in Suriname and lowest in French Guiana. This confirms the the results from both matrices as it shows that diversity in Guyana is highest, and decreases in Suriname and French Guiana. The occurrence of haplotype no. 2 of the trnH-psbA marker also suggests a pattern of decreasing genetic diversity from west to east as its frequency increases strongly from west to east. In Guyana, haplotype no. 2 is totally absent, while in French Guiana, most individuals are of this haplotype.

4.2. Ycf6-trnc The high haplotype diversity of the ycf6-trnC intergenic spacer region suggests that it has a higher mutation rate than that of trnH-psbA. However, it might also be an artefact caused by the lower quality sequences available from this marker. Therefore caution is exercised in drawing conclusions from these results. Yet its pattern appears to be consistent with that of the other marker, indicating high diversity in Guyana, and decreasing diversity towards the east. The values yielded by the AMOVA analysis for this marker show lower, yet highly significant F-statistics, indicating a lower degree of overall differentiation or, as mentioned before, a high rate of mutation. Remarkably, most of the explained variation originates between groups and populations overall, and not among groups as in the trnH-psbA marker. This indicates a lower degree of group structure within geographically related areas. The matrix of pairwise FST appears to be unusable for inferring clear genetic patterns from the ycf6-trnC marker. Although yielding many statistically significant values, again genetic differentiation is not restricted to any geographically related areas.

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Despite this, a noticeable pattern can still be observed when looking at pairwise genetic distance. Although the pattern here is not as clear as that from trnH-psbA, again the highest variation within populations can be found in Guyana, and these populations also differ most strongly from all others. This is supported by the distribution of haplotypes. Here again, one haplotype, namely haplotype no. 7 was by far most abundant and its frequency of occurrence was highest in French Guiana and lowest in Guyana, where it occurs only once. This is also clearly reflected in the very high haplotype α-diversity found in Guyana compared to French Guiana and Suriname. This suggests a genetic origin in Central Guyana, and a subsequent dispersal to the east.

4.3. Overall diversity and dispersal With the AMOVA calculation for one marker showing strong and for the other showing weak overall differentiation both markers at first glance appear to yield contrasting results. Would more high quality sequences have been available it would have been possible to combine the two markers and in that way mediate the results in one AMOVA calculation, matrix of pairwise FST and matrix of average pairwise differences and genetic distance in a single go. Now that both analyses are only available separately it is only possible to qualitatively infer a combined pattern from them. The apparent low number of populations and large geographical gaps between populations that were sampled in Guyana and Suriname, combined with the unconventional statistical methods, do however not oppose the revelation of a genetic pattern. For AMOVA analysis, the populations were grouped according to country, comparable soil conditions and geographical proximity. Since no environmental data were available for many samples from French Guiana, the current grouping in that country based on geographical proximity seemed most logical. If environmental data such as soil condition would have been available, grouping based on such factors could have yielded slightly different AMOVA results. However, due to the low relative haplotype diversity in French Guiana, this effect would not have been very large. With one marker showing strong overall differentiation and an apparent low degree of gene flow, and the other weak overall differentiation and a much higher degree of gene flow, both still show a similar pattern. Both markers indicate a relatively large amount of genetic distance within populations from Guyana and when these populations are compared to those outside of Guyana. These genetic distances suggest that genetic diversity is highest in Guyana and decreases in Suriname and French Guiana. From the haplotype distributions and α-diversities of the two separate markers it is very easy to infer a clear pattern as both show similar results. In conventional analysis these intergenic spacer regions would have been merged yielding intermediate results, but still a very clear pattern of genetic diversity. When qualitatively combined the haplotype distributions and α-diversities show a pattern of decreasing haplotype diversity from west to east. This pattern is most easily inferred from the presence of haplotype No. 2 of trnH-psbA and haplotype No. 7 of ycf6-trnC. This pattern suggests that the species Eperua falcata originated in, or near central Guyana and then dispersed east through Suriname into French Guiana. This dispersal may have taken place in several steps and was likely combined with one or more founder effects (sensu Mayr, 1954). These findings appear to be supported by the distribution of Eperua falcata across the Guiana shield. This distribution shows very high abundance in central Guyana and Suriname, and decreasing abundance in eastern Suriname and French-Guiana (Figure 11) (ter Steege et al., 2013, in press).

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Figure 11: Distribution of Eperua falcata across the Guiana Shield where darker shades of grey indicate higher relative

abundances (ter Steege et al., 2013, in press).

Figure 12: A fairly complete overview of locations from which botanical collections of E.

falcata exist. Areas covered by white sand soil are marked yellow. It is important to note

that this overview does not provide any information on the abundance of E. falcata on

these locations (ter Steege unpublished data).

23

While E. falcata is quite abundant across the entire Guiana Shield, it is a (co-)dominant tree species in many white sand areas (ter Steege et al., 1993). White sand soils are characterized as being nutrient poor and well drained, leading to nutrient deficiencies and drought stress that relatively few species are adapted to (Fanshawe, 1952). At present day, overall tree alpha diversity of the Guyana Shield decreases from west to east. Also the eastern Guiana Shield is substantially wetter in terms of rainfall than the western part, which is usually an important factor for tree species diversity (Wright, 1992). However, there is no apparent correlation between alpha diversity and rainfall across the Guianas, suggesting rainfall is not a limiting factor in this area (ter Steege et al., 2000). The difference in tree diversity, and possible cause of the E. falcata abundance pattern, can therefore likely be attributed to soil conditions. In Guyana and Suriname, large areas are covered by white sand soils (Figure 12). Competition by other tree species outside these white sand areas could simply much higher, leading to the present pattern of biodiversity (Wright, 1992). While the present Neotropical tree diversity is thought of to have originated mostly during the Tertiary, present species distribution and genetic diversity patterns are likely to have been caused by more recent historical events, regeneration dynamics and species interactions. In glacial and interglacial periods during the Quaternary, forested areas have contracted and expanded several times (Bush, 1994; Charles-Dominique et al., 1998; Hoorn et al., 2010; Veullemier, 1971). During glacial periods large areas of this region are thought to have been subjected to lowered annual mean temperatures and up to 40% reduced rainfall and were subsequently covered by different forest types and savannahs (Figure 13) (Hooghiemstra and van der Hammen, 1998; van der Hammen and Hooghiemstra, 2000). However the Quaternary forest developments in both the Amazon and the Guianas are still a matter of much debate. Some studies suggest that most of the areas presently covered by moist evergreen broadleaf forests actually remained forested during these periods, but may have changed in terms of canopy density and tree species composition (Figure 14)(Beerling and Mayle, 2006; Colinvaux et al., 1996; Colinvaux et al., 2000; Cowling et al., 2001). Evidence for multiple different scenarios has been found and the true biogeographic history of E. falcata is highly likely to have been influenced by such events.

Figure 13: Maps showing the modern distribution of rain forest

in relation to the present day 2500, 2000 and 1500 mm

precipitation isoline. After a reduction of 25% and 40%, new

1500 mm iso-contours show the potential area with rainforest.

The situation during the Last Glacial Maximum (LGM), some

20,000 before present, might have been intermediate

between the -25% and -40% scenarios. (Hooghiemstra and van

der Hammen, 1998)

24

White sand forests in the Guianas are not continuous throughout the entire region and to some degree analogous to seasonally dry forests and Amazonian white sand forests (Anderson, 1981). They therefore possibly constitute present day refugia or island patches to species that are otherwise outcompeted outside of these areas (Macedo and Prance, 1978). The largest continuous areas of white sand soil on the Guiana Shield are found in Guyana. Although E. falcata is not endemic to these white sand areas, it is often a (co-)dominant canopy tree species, after which the forest type is often named (Wallaba forest). These areas can therefore be seen as edaphic islands, particularly hospitable to species such as E. falcata. Hence, E. falcata, being a species well adapted to edaphic conditions, possibly followed the migration pattern from west to east by benefitting from decreased competition outside the contracted wet forests during dryer periods. During glacial forest changes and contractions, it may have been able to remain present in the changing environment. It may then have rapidly dispersed prior to post-glacial forest expansion, where conditions permitted it. After moist forests expanded, the abundance of E. falcata outside white sand areas presumably became more fragmented due to increased competition by other tree species. This process may have taken several steps and took place in conjunction with one or more founder events, causing the present pattern of genetic diversity (Figure 15).

Figure 14: Maps showing possible spatial and temporal vegetation cover of the Amazon and surrounding areas. These

show that the Guianas largely remained forested during the Last Glacial Maximum (LGM), although the forest may

have had a different composition or canopy density (Beerling and Mayle, 2006).

25

Figure 15: Reconstruction of the possible vegetation pattern in the Amazon basin and surroundings during the LGM, some

20,000 BP, assuming a general reduction of 40% in rainfall (van der Hammen en Hooghiemstra, 2000). The map is elaborated

with red arrows that show the direction of dispersal as inferred from the data gathered in this study. The areas marked in red

roughly show the present distribution of E. falcata where darker shades of red indicate areas that are thought of to have been

colonized earlier in geological history. The dispersal of E. falcata is likely to have taken place at some time prior to, or after the

LGM, and may have taken several steps. Also, conditions during the LGM may have been less dry than suggested on this map.

26

5. Recommendations Because of time limitations unfortunately a number of critical analyses for a more accurate and elaborate phylogeographic study could not be done. For proper phylogeographic analysis a minimum spanning network and a haplotype diversity map, as well as divergence time dating should be added to increase the scientific rigor of the study. However, a combination of software problems and low quality data made it impossible to do this within the amount of time set for this study. For future research I recommend that prior to the study, the student or researcher in question first gets him or herself familiar with the field of phylogeography, the analyses and software involved, and builds up a network of people who are well established in the field. This could vastly increase productivity as less time is wasted on getting to know all the different aspects of this kind of research and the usage of the software packages involved. In my view, the time used for just getting familiar with the software could have been much better spent on the actual contents, thereby yielding scientifically stronger conclusions. To pinpoint both the spatial and temporal the origins of Eperua falcata, a more elaborate sampling of individuals from Guyana en Suriname is recommended. When based on a larger number of genetic markers and quantitative methods, this might then reveal more precise regional and temporal dispersal patterns and possible origins of speciation (Cruzan and Templeton, 2000).

6. Aknowledgements I wish to thank Ted Feldpausch and Nikée Groot of the University of Leeds, the Guyana Forestry Commission and the Iwokrama International Centre for providing me the means of going into the forested interior of Guyana for the necessary fieldwork. I am grateful to Raven Kartoikromo of Natuurbeheer Suriname for his assistance during the fieldwork in Suriname and to Stinasu for providing me access to the Brownsberg National Park. I am also very grateful to Ivan Scotti (EcoFoG) for his help with DNA extractions and sequencing and for providing me with all the lab equipment needed for this in Kourou, French-Guiana. And most of all for providing me with all the Eperua falcata DNA sequences he had available from French-Guiana, thereby more than doubling the size of my dataset. Most thanks go to my supervisor Hans ter Steege for allowing me to conduct this study and always being available when I needed him. His extensive network has brought me into contact with all people and institutions mentioned above, and made this whole study possible. Last, but not least, I am thankful to the Van Eeden Fonds and Miquel Fonds for the financial support they have provided to me.

27

7. References Aide, T.M., Rivera, E., 1998. Geographic patterns of genetic diversity in Poulsenia armata (Moraceae): implications for the theory of Pleistocene refugia and the importance of riparian forest. Journal of Biogeography 25, 695-705. Amazon Tree Diversity Network, 2013. R code to calculate Fisher’s alpha. Available on http://testweb.science.uu.nl/Amazon/atdn/pagina39.html, accessed on September 27, 2013.

Amusant, N., Moretti, C., Richard, B., Prost, E., Nuzillard, J.M., Thévenon, M.F., 2007. Chemical compounds from Eperua falcata and Eperua grandiflora heartwood and their biological activities againstwood destroying fungus (Coriolus versicolor). Holz Roh Werkst 65, 23–28. Anderson, A.B., 1981. White-sand vegetation of Brazilian Amazonia. Biotropica 13 No. 3, 199-210. Avise, J.C., 2000. Phylogeography: the history and formation of species. Harvard University Press, Cambridge, Massachusetts. Baraloto, C., Morneau, F., Bonal, D., Blanc, L., Ferry, B. 2007. Seasonal water stress tolerance and habitat associations within four neotropical tree genera. Ecology 88 No. 2, 478-489. Beerling, D.J., Mayle, F.E., 2006. Contrasting effects of climate and CO2 on Amazonian ecosystems since the last glacial maximum. Global Change Biology 12 No. 10, 1977-1984. Birky, C.W., 1995. Uniparental inheritance of mitochondrial and chloroplast genes: Mechanisms and evolution. Proc. Natl. Acad. Sci. USA 92, 11331-11338. Bush, M.B., 1994. Amazonian Speciation: A Necessarily Complex Model. Journal of Biogeography 21 No. 1, 5-17. Charles-Dominique, P., Blanc, P., Larpin, D., Ledru, M., Riéra, B., Sarthou, C., Servant, M., Tardy, C., 1998. Forest perturbations and biodiversity during the last ten thousand years in French Guiana. Acta Oecologica 19 No. 3, 295-302. Colinvaux, P.A., De Oliveira, P.E., Moreno, J.E., Miller, M.C., Bush, M.B., 1996. A long pollen record from lowland Amazonia: forest and cooling in glacial times. Science 274 No. 5284, 85-88. Colinvaux, P.A., De Oliveira, P.E., Bush, M.B., 2000. Amazonian and neotropical plant communities on glacial time-scales: the failure of the aridity and refuge hypotheses. Quaternary Science Reviews 19, 141-169. Cowan, R.S., 1975. A Monograph of the Genus Eperua (Leguminosae: Caesalpnioideae). Smithsonian Institution Press, City of Washington. Cowling, S.A., Maslin, M.A., Sykes, M.T., 2001. Paleovegetation simulations of lowland Amazonia and implications for neotropical allopatry and speciation. Quaternary Research 55 No. 2, 140-149. Cruzan, M.B.,Templeton, A.R., 2000. Paleoecology and coalescence: phylogeographic analysis of hypotheses from the fossil record". Trends in Ecology and Evolution 15 No. 12, 491–496.

28

Dauby. G., Duminil, J., Heuertz, M., Hardy, O.J., 2010. Chloroplast DNA Polymorphism and Phylogeography of a Central African Tree Species Widespread in Mature Rainforests: Greenwayodendron suaveolens (Annonaceae). Tropical Plant Biol. 3, 4–13. De Oliveira, A.A., Mori, S.A., 1999. A central Amazonian terra firme forest. I. High tree species richness on poor soils. Biodiversity and Conservation 8, 1219–1244. Dick, C.W., 2010. Phylogeography and Population Structure of Tropical Trees. Tropical Plant Biol. 3, 1-3. Duminil, J., Caron, H., Scotti, I., Cazal, S.O. & Petit, R.J., 2006. Blind population genetics survey of tropical rainforest trees. Molecular Ecology 15, 3505–3513. Eo, S.H., Wares, J.P., Carroll, J.P., 2008. Population divergence in plant species reflects latitudinal biodiversity gradients. Biology Letters 4, 382-384. Excoffier, L., Smouse, P.E., Quattro, J.M., 1992. Analysis of Molecular Variance Inferred From Metric Distances Among DNA Haplotypes: Application to Human Mitochondrial DNA Restriction Data. Genetics 131, 479-491. Excoffier, L., Laval, G., Schneider, S., 2005. Arlequin version 3: An integrated software package for population genetics data analysis. Evolutionary Bioinformatics Online 1, 47-50. Fanshawe, D.B., 1952. The vegetation of British Guiana. A preliminary review. Institute Paper No 29. Imperial Forestry Institute, University of Oxford, Oxford. Fine, P.V.A., Ree, R.H., 2006. Evidence for a Time-Integrated Species-Area Effect on the Latitudinal Gradient in Tree Diversity. The American Naturalist 168 No. 6, 796-804. Gentry, A.H., 1988. Changes in Plant Community Diversity and Floristic Composition on Environmental and Geographical Gradients. Annals of the Missouri Botanical Garden 75 No. 1, 1-34. Grivet, D., Heinze, B., Vendramin, G.G., Petit, R.J., 2001. Genome walking with consensus primers: application to the large single copy region of chloroplast DNA. Molecular Ecology Notes 1, 345–349. Guyana Forestry Commission, 2004. Guyana woods, 21 species. Guyana. Hammond, D.S., Brown, V.K., 1995, Seed Size of Woody Plants in Relation to Disturbance, Dispersal, Soil Type in Wet Neotropical Forests. Ecology, 76 No. 8, 2544-2561. Hammond, D.S., Gond, V., de Thoisy, B., Forget, P-M., DeDijn, B.P.E., 2007. Causes and Consequences of a Tropical Forest Gold Rush in the Guiana Shield, South America. A Journal of the Human Environment 36 No. 8, 661-670. Hardesty, B.D., Dick, C.W., Hamrick, J.L., Degen, B., Hubell, S.P., Bermingham, E., 2010. Geographic Influence on Genetic Structure in the Widespread Neotropical Tree Simarouba amara (Simaroubaceae). Tropical Plant Biol. 3, 28–39. Hickerson, M.J., Carstens, B.C., Cavender-Bares, J., Crandall, K.A., Graham, C.H., Johnson, J.B., Rissler, L., Victoriano, P.F., Yoder, A.D., 2010. Phylogeography’s past, present, and future: 10 years after Avise, 2000. Molecular Phylogenetics and Evolution 54, 291-301.

29

Hooghiemstra, H., van der Hammen, T., 1998. Neogene and Quaternary development of the neotropical rain forest: the forest refugia hypothesis, and a literature overview. Earth-Science Reviews 44 No. 3, 147-183. Hoorn, C., Wesselingh, F.P., ter Steege, H., Bermudez, M.A., Mora, J., Sevink, J., Sammartín, I., Sanchez-Mesequer, A., Anderson, C.L., Figueiredo, J.P., Jaramillo, C., Riff, D., Negri, F.R., Hooghiemstra, H., Lundberg, J., Stadler, T., Särkinen, T., Antonelli, A., 2010. Amazonia Through Time: Andean Uplift, Climate Change, Landscape Evolution, and Biodiversity Science 330, 927-931. Hubbell, S.P., He, F., Condit, R., Borda-de-A, L., Kellner, J., ter Steege, H., 2008. How many tree species are there in the Amazon and how many of them will go extinct? PNAS 105 Suppl. 1, 11498-11504. Johnston, M., 1998. Tree Population studies in low-diversity forests, Guyana. II. Assessments on the distribution and abundance of non-timber forest products. Biodiversity and Conservation 7, 73-86. Johnston, M., Gillman, M., 1995. Tree Population studies in low-diversity forests, Guyana. I. Floristic composition and stand structure. Biodiversity and Conservation 4, 339-362. Knowles, L.L., Maddison, W.P., 2002. Statistical Phylogeography. Molecular Ecology 11, 2623-2635. Lescure, J.P., Boulet, R., 1985. Relationships Between Soil and Vegetation in a Tropical Rain Forest in French Guiana. Biotropica 17, No. 2, 155-164. Lemes, M.R., Dick, C.W., Navarro, C., Lowe, A.J., Cavers, S., Gribel, R., 2010. Chloroplast DNA Microsatellites Reveal Contrasting Phylogeographic Structure in Mahogany (Swietenia macrophylla King, Meliaceae) from Amazonia and Central America. Tropical Plant Biol. 3, 40-49. Lowe, A.J., Harris, D., Dormontt, E., Dawson, I.K., 2010. Testing Putative African Tropical Forest Refugia Using Chloroplast and Nuclear DNA Phylogeography. Tropical Plant Biol. 3, 50–58. Mayr, E., 1954. Change of genetic environment and evolution. In: Huxley, J., Hardy, A.C., Ford, E.B., Evolution as a process, 157-180. Allen and Unwin, London. McCauley D.E., 1995. The use of chloroplast DNA polymorphism in studies of gene flow in plants. TREE 10, No. 5, 198-202. Milliken, W., Zappi, D., Sasaki, D., Hopkins, M., Pennington, R.T., 2010. Amazon vegetation: how much don’t we know and how much does it matter? Kew Bulletin 65, 691-709. Macedo, M., Prance, G.T., 1978. Notes on the vegetation of Amazonia II. The dispersal of plants in Amazonian white sand campinas: the campinas as functional islands. Brittonia 30 No. 2, 203-215. Mittelbach, G.G., Schemske, D.W., Cornell, H.V., Allen, A.P., Brown, J.M., Bush, M.B., Harrison, S.P., Hurlbert, A.H., Knowlton, N., Lessios, H.A., McCain, C.M., McCune, A.R., McDade, L.A., McPeek, M.A., Near, T.J.,. Price, T.D., Ricklefs, R.E., Roy, K., Sax. D.F., Schluter, D., Sobel, J.M., Turelli, M., 2007. Evolution and the latitudinal diversity gradient: speciation, extinction and biogeography. Ecology Letters 10, 315–331. Moritz, C., 1994. Defining ‘Evolutionarily Significant Units’ for conservation. TREE 9 No. 10, 373-375.

30

Moritz, C., Faith, D.P., 1998. Comparative phylogeography and the identification of genetically divergent areas for conservation. Molecular Ecology 7, 419-429. Moritz, C., 2002. Strategies to Protect Biological Diversity and the Evolutionary Processes That Sustain It. Syst. Biol. 51, 238–254. Nei, M., Li, W.H., 1979. Mathematical model for studying genetic variation in terms of restriction endonucleases. Proc.Natl.Acad.Sci.USA 76:5269-5273. Newton, A.C., Allnutt, T.R., Gillies, A.C.M., Lowe, A.J., Ennos, R.A., 1999. Molecular phylogeography, intraspecific variation and the conservation of tree species. TREE 14 No. 4, 140-145. Phillips, O.L., Hall, P., Gentry, A.H., Sawyer, S.A., Vasquez, R., 1994. Dynamics and species richness of tropical rain forests. Proc. Natl. Acad. Sci. USA 91,2805-2809. Polak, A.M., 1992. Major Timber Trees of Guyana. A Field Guide. The Tropenbos Foundation, Wageningen. R Core Team, 2013. R: A language and Environment for Statistical Computing. R Foundation for Statistical Computing. Vienna, Austria. Available on http://www.R-project.org Rozendaal, D., 2004. Eperua (Fabaceae) Evolution and Biogeography. MSc-project report. Group of Plant Systematics, Faculty of Biology, Utrecht University, Utrecht. Scotti-Saintagne, C., Dick, C.W., Caron, H., Venramin, G.G., Troispoux, V., Sire, P., Casalis, M., Buonamici, A., Valencia, R., Lemes, M.R., Gribel, R., Scotti, I., 2012. Amazon diversification and cross- Andean dispersal of the widespread Neotropical tree species Jacaranda copaia (Bignoniaceae). Journal of Biogeography 40 No. 4, 707-719. Scotti-Saintagne, C., Dick, C.W., Caron, H., Venramin, G.G., Guichoux, E., Buonamici, A., Duret, C., Sire, P., Valencia, R., Lemes, M.R., Gribel, R., Scotti, I., 2013. Phylogeography of a species complex of lowland Neotropical rain forest trees (Carapa, Meliaceae). Journal of Biogeography 40 No. 4, 676-692. Shaw, J., Lickey, E.B., Beck, J.T., Farmer, S.B., Liu, W., Miller, J., Siripun, K.C., Winder, C.T., Schilling, E.E., Small, R.L., 2005. The Tortoise and the Hare II: Relative Utility of 21 Noncoding Chloroplast DNA Sequences for Phylogenetic Analysis. American Journal of Botany 92 No. 1, 142-166. ter Steege, H., Cornelissen, H.C., 1989. Distribution and Ecology of Vascular Epiphytes in Lowland Rain Forest of Guyana. Biotropica 21 No. 4, 331-339. ter Steege, H., Jetten, V.G., Polak, A.M., Werger, M.J.A., 1993. Tropical rain forest types and soil factors in a watershed area in Guyana. Journal of Vegetation Science 4, 705-716. ter Steege, H., Sabatier, D., Castellanos, H., van Andel, T., Duivenvoorden, J., Adalardo de Oliveira, A., Ek, R., Lilwah, R., Maas, P., Mori, S., 2000. An analysis of the floristic composition and diversity of Amazonian forests including those of the Guiana Shield. Journal of Tropical Ecology 16, 801-828. ter Steege, H., Zondervan, G., 2000. A preliminary analysis of large-scale forest inventory data of the Guiana Shield. Plant diversity in Guyana. With recommendations for a National Protected Area Strategy (Tropenbos Series 18). Wageningen, The Tropenbos Foundation, 35-54.

31

ter Steege, H., Pitman, N., Sabatier, D., Castellanos, H., van der Hout, P., Daly, D.C., Silveira, M., Phillips, O., Vasquez, R., van Andel, T., Duivenvoorden, J., de Oliveira, A.A., Ek, R., Lilwah, R., Thomas, R., van Essen, J., Baider, C., Maas, P., Mori, S., Terborgh, J., Núñez Vargas, P., Mogollo, H., Morawetz, W., 2003. A spatial model of tree α-diversity and tree density for the Amazon. Biodiversity and Conservation 12, 2255-2277. ter Steege, H., Bánki, O.S., van Andel, T.R., Behari-Ramdas, J., Ramharakh, G., 2004. Plant diversity of the Brownsberg Nature Park, Suriname. Report of the Nov-Dec 2003 Expedition. NHN-Utrecht Branch, Utrecht University, Utrecht. ter Steege et al., 2013. Hyper-dominance in the Amazonian Tree Flora. Science, in press. Tuomisto, H., Ruokolainen, K., Yli-Halla, M., 2003. Dispersal, Environment, and Floristic Variation of Western Amazonian Forests. Science 299, 241-244. van der Hammen, T., Hooghiemstra, H., 2000. Neogene and Quaternary history of vegetation, climate, and plant diversity in Amazonia. Quaternary Science Reviews 19 No. 8, 725-742 Vink, A.T., 1965. Surinam Timbers. Surinam Forest Service, Ministry of Development Surinam, Paramaribo. Vuilleumier, B.S., 1971. Pleistocene Changes in the Fauna and Flora of South America. Science, New Series 173 No. 3999, 771-780. Weir, B.S., Cockerham, C.C., 1984. Estimating F-statistics for the Analysis of Population Structure. Evolution 38 No. 6, 1358-1370. Willig, M.R., Kaufman, D.M., Stevens, R.D., 2003. Latitudinal Gradients of Biodiversity: Pattern, Process, Scale and Synthesis. Annu. Rev. Ecol. Evol. Syst. 34, 273-309. Wright, S.J., 1992. Seasonal Drought, Soil Fertility and Tree Species Density of Tropical Forest Plant Communities. TREE 7 No. 8, 260-263.

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8. Appendix Table 1: Population pairwise FST values based on the trnH-psbA marker. All significant values are marked yellow. Significance level P ≤ 0.05. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri 0

Mabura Hill 0.70588 0

Iwokrama -0.30496 0.6993 0

Zanderij 0.74351 0.99215 0.73864 0

Powakka 0.71717 1 0.71045 -0.17718 0

Mata 0.78082 1 0.77559 0 0 0

Brownsberg 0.70588 1 0.69794 -0.0687 0 0 0

Cropi 0.66622 0.98135 0.65971 -0.02773 -0.14754 0.05138 -0.03448

Acarouany 0.52941 1 0.5198 -0.31765 0 0 0

Bafog 0.81221 1 0.80759 0.02946 0 0 0

Organabo 0.52941 1 0.5198 -0.31765 0 0 0

La Mana. S Lezard 0.33333 1 0.31884 -1 0 0 0

La Mana. S Fracas 0.66667 1 0.65942 -0.10891 0 0 0

La Mana. cr Korossibo 0.6129 1 0.60477 -0.17483 0 0 0

St Elie 0.6129 1 0.60477 -0.17483 0 0 0

Paracou 0.6129 1 0.60477 -0.17483 0 0 0

Mt Singes 0.81221 1 0.80759 0.02946 0 0 0

PK 72 Est 0.52941 1 0.5198 -0.31765 0 0 0

Cr Plomb 0.6129 1 0.60477 -0.17483 0 0 0

Tresor 0.61915 0.89425 0.61905 -0.08423 -0.18919 -0.03733 -0.09272

Mt Trinite 0.83562 1 0.83149 0.05352 0 0 0

Camp Cisame 0.52941 1 0.5198 -0.31765 0 0 0

Approuague 0.52941 1 0.5198 -0.31765 0 0 0

Nouragues 0.52941 1 0.5198 -0.31765 0 0 0

St Lavillette 0.52941 1 0.5198 -0.31765 0 0 0

Mt couronnee 0.66667 1 0.65942 -0.10891 0 0 0

Piste Belizon 0.52941 1 0.5198 -0.31765 0 0 0

saut deux roros 0.6129 1 0.60477 -0.17483 0 0 0

Oyapok 0.76068 1 0.75507 -0.01818 0 0 0

33

Table 1 continued

Cropi Acarouany Bafog Organabo La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo St Elie

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi 0

Acarouany -0.30435 0

Bafog 0.09091 0 0

Organabo -0.30435 0 0 0

La Mana. S Lezard -1 0 0 0 0

La Mana. S Fracas -0.08108 0 0 0 0 0

La Mana. cr Korossibo -0.15385 0 0 0 0 0 0

St Elie -0.15385 0 0 0 0 0 0 0

Paracou -0.15385 0 0 0 0 0 0 0

Mt Singes 0.09091 0 0 0 0 0 0 0

PK 72 Est -0.30435 0 0 0 0 0 0 0

Cr Plomb -0.15385 0 0 0 0 0 0 0

Tresor -0.15438 -0.32663 -0.01617 -0.32663 -1 -0.12821 -0.18919 -0.18919

Mt Trinite 0.12409 0 0 0 0 0 0 0

Camp Cisame -0.30435 0 0 0 0 0 0 0

Approuague -0.30435 0 0 0 0 0 0 0

Nouragues -0.30435 0 0 0 0 0 0 0

St Lavillette -0.30435 0 0 0 0 0 0 0

Mt couronnee -0.08108 0 0 0 0 0 0 0

Piste Belizon -0.30435 0 0 0 0 0 0 0

saut deux roros -0.15385 0 0 0 0 0 0 0

Oyapok 0.02778 0 0 0 0 0 0 0

34

Table 1 continued

Paracou Mt Singes PK 72 Est Cr Plomb Tresor Mt Trinite Camp Cisame Approuague

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog

Organabo

La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo

St Elie

Paracou 0

Mt Singes 0 0

PK 72 Est 0 0 0

Cr Plomb 0 0 0 0

Tresor -0.18919 -0.01617 -0.32663 -0.18919 0

Mt Trinite 0 0 0 0 0 0

Camp Cisame 0 0 0 0 -0.32663 0 0

Approuague 0 0 0 0 -0.32663 0 0 0

Nouragues 0 0 0 0 -0.32663 0 0 0

St Lavillette 0 0 0 0 -0.32663 0 0 0

Mt couronnee 0 0 0 0 -0.12821 0 0 0

Piste Belizon 0 0 0 0 -0.32663 0 0 0

saut deux roros 0 0 0 0 -0.18919 0 0 0

Oyapok 0 0 0 0 -0.05119 0 0 0

35

Table 1 continued

Nouragues St Lavillette

Mt couronnee

Piste Belizon

saut deux roros Oyapok

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog

Organabo

La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Tresor

Mt Trinite

Camp Cisame

Approuague

Nouragues 0

St Lavillette 0 0

Mt couronnee 0 0 0

Piste Belizon 0 0 0 0

saut deux roros 0 0 0 0 0

Oyapok 0 0 0 0 0 0

36

Table 2: P-values of population pairwise FST gained from the trnH-psbA marker, based on 1023 permutations. All significant values (P ≤ 0.05) are marked yellow. Note that, due to its size, the table is spread across six pages.

Pibiri Mabura Hill Iwokrama Zanderij

Pibiri *

Mabura Hill 0.03809 ± 0.0060 *

Iwokrama 0.65820 ± 0.0153 0.04883 ± 0.0062 *

Zanderij 0.00000 ± 0.0000 0.00293 ± 0.0016 0.00293 ± 0.0016 *

Powakka 0.00293 ± 0.0016 0.00098 ± 0.0010 0.00000 ± 0.0000 0.99902 ± 0.0002

Mata 0.00293 ± 0.0016 0.00000 ± 0.0000 0.00195 ± 0.0014 0.99902 ± 0.0002

Brownsberg 0.00977 ± 0.0033 0.00586 ± 0.0022 0.01562 ± 0.0044 0.99902 ± 0.0002

Cropi 0.00391 ± 0.0019 0.00000 ± 0.0000 0.00195 ± 0.0014 0.77246 ± 0.0116

Acarouany 0.06445 ± 0.0077 0.05762 ± 0.0075 0.05273 ± 0.0064 0.99902 ± 0.0002

Bafog 0.00195 ± 0.0014 0.00098 ± 0.0010 0.00195 ± 0.0014 0.44043 ± 0.0163

Organabo 0.05566 ± 0.0074 0.04883 ± 0.0059 0.05859 ± 0.0079 0.99902 ± 0.0002

La Mana. S Lezard 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. S Fracas 0.02344 ± 0.0054 0.00781 ± 0.0028 0.02734 ± 0.0053 0.99902 ± 0.0002

La Mana. cr Korossibo 0.02734 ± 0.0065 0.02051 ± 0.0038 0.02930 ± 0.0046 0.99902 ± 0.0002

St Elie 0.03125 ± 0.0057 0.02246 ± 0.0053 0.02441 ± 0.0043 0.99902 ± 0.0002

Paracou 0.02637 ± 0.0042 0.02148 ± 0.0043 0.03320 ± 0.0066 0.99902 ± 0.0002

Mt Singes 0.00000 ± 0.0000 0.00000 ± 0.0000 0.00098 ± 0.0010 0.44922 ± 0.0206

PK 72 Est 0.06055 ± 0.0066 0.04590 ± 0.0059 0.06738 ± 0.0070 0.99902 ± 0.0002

Cr Plomb 0.02148 ± 0.0040 0.02734 ± 0.0065 0.04102 ± 0.0059 0.99902 ± 0.0002

Tresor 0.00293 ± 0.0016 0.00000 ± 0.0000 0.00195 ± 0.0014 0.99902 ± 0.0002

Mt Trinite 0.00195 ± 0.0014 0.00000 ± 0.0000 0.00195 ± 0.0014 0.39941 ± 0.0162

Camp Cisame 0.06641 ± 0.0101 0.06250 ± 0.0067 0.03613 ± 0.0047 0.99902 ± 0.0002

Approuague 0.06348 ± 0.0065 0.03125 ± 0.0062 0.08301 ± 0.0081 0.99902 ± 0.0002

Nouragues 0.07715 ± 0.0078 0.04688 ± 0.0058 0.08496 ± 0.0093 0.99902 ± 0.0002

St Lavillette 0.07520 ± 0.0076 0.04785 ± 0.0064 0.06738 ± 0.0098 0.99902 ± 0.0002

Mt couronnee 0.02637 ± 0.0056 0.01172 ± 0.0030 0.03125 ± 0.0052 0.99902 ± 0.0002

Piste Belizon 0.07031 ± 0.0063 0.04785 ± 0.0075 0.06348 ± 0.0066 0.99902 ± 0.0002

saut deux roros 0.02344 ± 0.0052 0.01660 ± 0.0034 0.02051 ± 0.0043 0.99902 ± 0.0002

Oyapok 0.00000 ± 0.0000 0.00098 ± 0.0010 0.00391 ± 0.0023 0.99902 ± 0.0002

37

Table 2 continued

Powakka Mata Brownsberg Cropi Acarouany

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka *

Mata 0.99902 ± 0.0002 *

Brownsberg 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Cropi 0.75684 ± 0.0132 0.43750 ± 0.0117 0.99902 ± 0.0002 *

Acarouany 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Bafog 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.37695 ± 0.0147 0.99902 ± 0.0002

Organabo 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. S Lezard 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. S Fracas 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. cr Korossibo 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

St Elie 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Paracou 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Singes 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.37012 ± 0.0122 0.99902 ± 0.0002

PK 72 Est 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Tresor 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.34375 ± 0.0128 0.99902 ± 0.0002

Camp Cisame 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Approuague 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Nouragues 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt couronnee 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Piste Belizon 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Oyapok 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.43555 ± 0.0115 0.99902 ± 0.0002

38

Table 2 continued

Bafog Organabo La Mana. S Lezard La Mana. S Fracas La Mana. cr Korossibo

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog *

Organabo 0.99902 ± 0.0002 *

La Mana. S Lezard 0.99902 ± 0.0002 0.99902 ± 0.0002 *

La Mana. S Fracas 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

La Mana. cr Korossibo 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

St Elie 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Paracou 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Singes 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

PK 72 Est 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Tresor 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Camp Cisame 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Approuague 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Nouragues 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt couronnee 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Piste Belizon 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Oyapok 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

39

Table 2 continued

St Elie Paracou Mt Singes PK 72 Est Cr Plomb

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog

Organabo

La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo

St Elie *

Paracou 0.99902 ± 0.0002 *

Mt Singes 0.99902 ± 0.0002 0.99902 ± 0.0002 *

PK 72 Est 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Tresor 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Camp Cisame 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Approuague 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Nouragues 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt couronnee 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Piste Belizon 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Oyapok 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

40

Table 2 continued

Tresor Mt Trinite Camp Cisame Approuague Nouragues

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog

Organabo

La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Tresor *

Mt Trinite 0.99902 ± 0.0002 *

Camp Cisame 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Approuague 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Nouragues 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt couronnee 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Piste Belizon 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Oyapok 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

41

Table 2 continued

St Lavillette Mt couronnee Piste Belizon saut deux roros Oyapok

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Acarouany

Bafog

Organabo

La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Tresor

Mt Trinite

Camp Cisame

Approuague

Nouragues

St Lavillette *

Mt couronnee 0.99902 ± 0.0002 *

Piste Belizon 0.99902 ± 0.0002 0.99902 ± 0.0002 *

saut deux roros 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Oyapok 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

42

Table 3: Population pairwise FST values based on the ycf6-trnC marker. All significant values are marked yellow. Significance level P ≤ 0.05. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri 0

Mabura Hill 0.23245 0

Iwokrama 0.16441 0.16441 0

Zanderij 0.45365 0.26917 0.01513 0

Powakka 0.79451 0.48661 0.17708 0.01171 0

Mata 0.56419 0.32395 0.03442 -0.04562 0.08547 0

Brownsberg 0.36618 0.12272 -0.00846 -0.09818 0.16438 0.03628 0

Cropi 0.69307 0.37563 0.08887 -0.02251 0.61582 0.05314 0.08449

Organabo 0.08899 -0.23835 -0.72103 -0.86811 0.875 -0.71963 -0.7257

Bafog 0.71437 0.40654 0.11822 0.01508 0.59203 0.11215 0.12207

La Mana. Saut Lezard 0.08038 -0.24201 -0.75885 -0.95536 0.66667 -0.91667 -0.77556

La Mana. Saut Fracas 0.60369 0.28032 0.00676 -0.09394 0.82423 0.00054 0.00025

La Mana. cr Korossibo 0.41622 0.11673 -0.1851 -0.2933 0.71106 -0.24742 -0.19337

St Elie 0.47362 0.16267 0.12597 0.21231 0.77566 0.45455 0.16431

Paracou 0.08685 -0.24018 -0.73973 -0.91331 0.81818 -0.84 -0.75219

Mt Singes 0.41572 0.1173 -0.18241 -0.2876 0.70711 -0.23389 -0.19318

PK 72 Est 0.40535 0.10341 -0.19558 -0.3004 0.84155 -0.22612 -0.20193

Cr Plomb 0.08685 -0.24018 -0.73973 -0.91331 0.81818 -0.84 -0.75219

Mt Trinite 0.57609 0.36319 0.10765 0.0407 0.22965 0.08659 0.10705

Tresor 0.67557 0.43427 0.14431 0.04913 0.26256 0.09509 0.14052

Camp Cisame 0.41783 0.11649 -0.17055 -0.26412 0.79579 -0.19288 -0.17969

Mt couronnee 0.41998 0.11587 -0.17931 -0.27807 0.84155 -0.22124 -0.18427

St Lavillette 0.08685 -0.24201 -0.73973 -0.91331 0.81818 -0.84 -0.75219

saut deux roros 0.41847 0.11537 -0.17304 -0.26538 0.79579 -0.18834 -0.17868

Piste Belizon 0.41394 0.11795 -0.16431 -0.24417 0.76507 -0.13936 -0.16514

Approuague 0.41768 0.11647 -0.17484 -0.26703 0.80769 -0.19589 -0.17783

Oyapok 0.66729 0.34066 0.06742 -0.03926 0.83333 0.04296 0.06148

43

Table 3 continued

Cropi Organabo Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo St Elie Paracou

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi 0

Organabo 0.47826 0

Bafog 0.42487 0.2973 0

La Mana. Saut Lezard -0.09091 1 -0.13043 0

La Mana. Saut Fracas 0.56364 0.73333 0.46845 0.55556 0

La Mana. cr Korossibo 0.25843 0.6 0.20819 -1 0.56364 0

St Elie 0.66178 0.03684 0.67604 0.00543 0.50383 0.36761 0

Paracou 0.29412 1 0.13333 1 -0.33333 0.33333 0.01081 0

Mt Singes 0.29412 0.33333 0.23718 -1 0.38462 -0.5 0.36519 0

PK 72 Est 0.46429 1 0.34414 1 0.11111 0.6 0.2523 0

Cr Plomb 0.29412 1 0.13333 1 -0.33333 0.33333 0.01081 0

Mt Trinite 0.13029 -0.55541 0.16433 -0.6811 -0.09889 -0.14925 0.10955 -0.71576

Tresor 0.16873 -0.37082 0.2037 -0.5586 0.03872 -0.08797 0.55346 -0.64895

Camp Cisame 0.48344 0.14286 0.38724 -0.2 -0.21212 0.2 0.36441 -1

Mt couronnee 0.46429 1 0.34414 1 0.11111 0.66667 0.36513 0

St Lavillette 0.29412 1 0.13333 1 -0.33333 0.33333 0 0

saut deux roros 0.47297 -0.2 0.38724 -0.2 0.02439 0.2 0.36602 -1

Piste Belizon 0.49754 -0.07692 0.42554 -0.55556 0.33945 0.2 0.36919 -1

Approuague 0.47826 0.5 0.37467 0 -0.14286 0.25 0.36842 -1

Oyapok 0.58323 0.875 0.49319 0.66667 0.12621 0.63585 0.63441 -1

44

Table 3 continued

Mt Singes PK 72 Est Cr Plomb Mt Trinite Tresor Camp Cisame

Mt couronnee

St Lavillette

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes 0

PK 72 Est 0.42857 0

Cr Plomb 0 0 0

Mt Trinite -0.14334 -0.34459 -0.71576 0

Tresor -0.07507 -0.15082 -0.64895 0.1517 0

Camp Cisame 0 -0.2 -1 -0.15768 -0.10791 0

Mt couronnee 0.5 0 0 -0.18583 -0.14439 0 0

St Lavillette 0 0 0 -0.73363 -0.64895 -1 0 0

saut deux roros 0.16667 -0.5 -1 -0.15541 -0.10791 -0.2 0 -1

Piste Belizon 0.18182 0 -1 -0.11528 -0.20095 0 0 -1

Approuague 0.2 -1 -1 -0.15916 -0.12172 -0.25 -0.33333 -1

Oyapok 0.60177 -1.16216 -1 0.06038 0.0691 0.22559 -0.29032 -1

45

Table 3 continued

saut deux roros

Piste Belizon Approuague Oyapok

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Mt Trinite

Tresor

Camp Cisame

Mt couronnee

St Lavillette

saut deux roros 0

Piste Belizon 0 0

Approuague -0.42857 0 0

Oyapok 0.22559 0.40853 0.10891 0

46

Table 4: P-values of population pairwise FST gained from the ycf6-trnC marker, based on 1023 permutations. All significant values (P ≤ 0.05) are marked yellow. Note that, due to its size, the table is spread across six pages.

Pibiri Mabura Hill Iwokrama Zanderij

Pibiri *

Mabura Hill 0.01953 ± 0.0036 *

Iwokrama 0.15137 ± 0.0102 0.06934 ± 0.0075 *

Zanderij 0.01562 ± 0.0034 0.01172 ± 0.0033 0.34277 ± 0.0127 *

Powakka 0.00195 ± 0.0014 0.00000 ± 0.0000 0.00293 ± 0.0016 0.26172 ± 0.0167

Mata 0.01270 ± 0.0034 0.00293 ± 0.0016 0.33594 ± 0.0134 0.87012 ± 0.0096

Brownsberg 0.01758 ± 0.0042 0.12109 ± 0.0122 0.40137 ± 0.0155 0.54883 ± 0.0110

Cropi 0.00781 ± 0.0028 0.00879 ± 0.0025 0.07129 ± 0.0093 0.38965 ± 0.0150

Organabo 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Bafog 0.00684 ± 0.0023 0.00000 ± 0.0000 0.00098 ± 0.0010 0.43652 ± 0.0211

La Mana. Saut Lezard 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. Saut Fracas 0.03320 ± 0.0047 0.02832 ± 0.0052 0.48340 ± 0.0139 0.29785 ± 0.0152

La Mana. cr Korossibo 0.09180 ± 0.0091 0.21094 ± 0.0086 0.53125 ± 0.0147 0.16602 ± 0.0093

St Elie 0.09082 ± 0.0096 0.09570 ± 0.0107 0.18066 ± 0.0131 0.20215 ± 0.0146

Paracou 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Singes 0.09570 ± 0.0072 0.16309 ± 0.0136 0.56543 ± 0.0140 0.20898 ± 0.0073

PK 72 Est 0.09961 ± 0.0110 0.27344 ± 0.0130 0.77051 ± 0.0143 0.16992 ± 0.0090

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.00684 ± 0.0027 0.00195 ± 0.0014 0.01074 ± 0.0026 0.17773 ± 0.0142

Tresor 0.00098 ± 0.0010 0.00098 ± 0.0010 0.01465 ± 0.0031 0.02148 ± 0.0053

Camp Cisame 0.11719 ± 0.0084 0.26367 ± 0.0136 0.30176 ± 0.0141 0.25684 ± 0.0132

Mt couronnee 0.10254 ± 0.0057 0.23145 ± 0.0135 0.45215 ± 0.0140 0.16406 ± 0.0104

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.12402 ± 0.0089 0.25293 ± 0.0159 0.37207 ± 0.0128 0.19629 ± 0.0085

Piste Belizon 0.10840 ± 0.0082 0.15430 ± 0.0107 0.39062 ± 0.0111 0.23730 ± 0.0121

Approuague 0.09668 ± 0.0100 0.25488 ± 0.0142 0.35254 ± 0.0134 0.19141 ± 0.0136

Oyapok 0.02246 ± 0.0047 0.00195 ± 0.0014 0.14844 ± 0.0105 0.38965 ± 0.0172

47

Table 4 continued

Powakka Mata Brownsberg Cropi Organabo

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka *

Mata 0.19727 ± 0.0108 *

Brownsberg 0.02637 ± 0.0056 0.38184 ± 0.0170 *

Cropi 0.00000 ± 0.0000 0.01758 ± 0.0037 0.19336 ± 0.0099 *

Organabo 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Bafog 0.00000 ± 0.0000 0.00293 ± 0.0016 0.00293 ± 0.0016 0.00000 ± 0.0000 0.24902 ± 0.0150

La Mana. Saut Lezard 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

La Mana. Saut Fracas 0.00000 ± 0.0000 0.43652 ± 0.0129 0.33301 ± 0.0152 0.00488 ± 0.0020 0.18945 ± 0.0109

La Mana. cr Korossibo 0.02539 ± 0.0045 0.27246 ± 0.0110 0.45215 ± 0.0149 0.07812 ± 0.0062 0.33301 ± 0.0135

St Elie 0.00488 ± 0.0020 0.03320 ± 0.0059 0.12891 ± 0.0133 0.02637 ± 0.0042 0.24023 ± 0.0130

Paracou 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Singes 0.02246 ± 0.0051 0.27637 ± 0.0135 0.40723 ± 0.0145 0.04004 ± 0.0066 0.71875 ± 0.0105

PK 72 Est 0.01855 ± 0.0036 0.30469 ± 0.0105 0.64551 ± 0.0112 0.02734 ± 0.0051 0.34473 ± 0.0142

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.00000 ± 0.0000 0.18359 ± 0.0122 0.06250 ± 0.0084 0.16406 ± 0.0133 0.34961 ± 0.0211

Tresor 0.00000 ± 0.0000 0.12598 ± 0.0091 0.07324 ± 0.0102 0.00098 ± 0.0010 0.46875 ± 0.0155

Camp Cisame 0.00391 ± 0.0019 0.31348 ± 0.0114 0.26172 ± 0.0144 0.04102 ± 0.0070 0.69141 ± 0.0119

Mt couronnee 0.01367 ± 0.0034 0.21777 ± 0.0106 0.26660 ± 0.0147 0.04199 ± 0.0077 0.36133 ± 0.0139

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.00586 ± 0.0022 0.30957 ± 0.0139 0.26367 ± 0.0093 0.05762 ± 0.0060 0.99902 ± 0.0002

Piste Belizon 0.01270 ± 0.0031 0.27441 ± 0.0138 0.30469 ± 0.0094 0.04004 ± 0.0049 0.65430 ± 0.0179

Approuague 0.01074 ± 0.0030 0.31152 ± 0.0151 0.23535 ± 0.0099 0.04395 ± 0.0058 0.30859 ± 0.0131

Oyapok 0.00000 ± 0.0000 0.12207 ± 0.0086 0.18750 ± 0.0148 0.00000 ± 0.0000 0.18652 ± 0.0114

48

Table 4 continued

Bafog La Mana. Saut Lezard La Mana. Saut Fracas

La Mana. cr Korossibo St Elie

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog *

La Mana. Saut Lezard 0.99902 ± 0.0002 *

La Mana. Saut Fracas 0.00195 ± 0.0014 0.22070 ± 0.0149 *

La Mana. cr Korossibo 0.29297 ± 0.0157 0.99902 ± 0.0002 0.12793 ± 0.0092 *

St Elie 0.01660 ± 0.0034 0.75098 ± 0.0124 0.07422 ± 0.0077 0.32422 ± 0.0133 *

Paracou 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Singes 0.24512 ± 0.0114 0.99902 ± 0.0002 0.15430 ± 0.0126 0.99902 ± 0.0002 0.27344 ± 0.0177

PK 72 Est 0.24023 ± 0.0153 0.33691 ± 0.0100 0.46680 ± 0.0135 0.36230 ± 0.0167 0.33008 ± 0.0124

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Mt Trinite 0.09473 ± 0.0082 0.77539 ± 0.0163 0.50488 ± 0.0150 0.42871 ± 0.0145 0.43262 ± 0.0170

Tresor 0.00293 ± 0.0016 0.90039 ± 0.0107 0.32715 ± 0.0144 0.37598 ± 0.0173 0.00781 ± 0.0024

Camp Cisame 0.07715 ± 0.0064 0.99902 ± 0.0002 0.72949 ± 0.0160 0.65625 ± 0.0131 0.29980 ± 0.0137

Mt couronnee 0.26270 ± 0.0101 0.28516 ± 0.0152 0.49316 ± 0.0146 0.31836 ± 0.0160 0.41797 ± 0.0181

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.08203 ± 0.0063 0.99902 ± 0.0002 0.66602 ± 0.0146 0.67383 ± 0.0122 0.29980 ± 0.0138

Piste Belizon 0.05469 ± 0.0087 0.63672 ± 0.0137 0.24512 ± 0.0128 0.33496 ± 0.0141 0.20801 ± 0.0116

Approuague 0.09082 ± 0.0089 0.66113 ± 0.0135 0.66797 ± 0.0130 0.70508 ± 0.0129 0.17285 ± 0.0117

Oyapok 0.00195 ± 0.0014 0.35449 ± 0.0136 0.38867 ± 0.0176 0.10352 ± 0.0109 0.03711 ± 0.0059

49

Table 4 continued

Paracou Mt Singes PK 72 Est Cr Plomb Mt Trinite

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo

St Elie

Paracou *

Mt Singes 0.64258 ± 0.0142 *

PK 72 Est 0.99902 ± 0.0002 0.31152 ± 0.0170 *

Cr Plomb 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Mt Trinite 0.99902 ± 0.0002 0.40723 ± 0.0148 0.95117 ± 0.0050 0.99902 ± 0.0002 *

Tresor 0.99902 ± 0.0002 0.37988 ± 0.0171 0.39746 ± 0.0127 0.99902 ± 0.0002 0.00293 ± 0.0016

Camp Cisame 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.43457 ± 0.0110

Mt couronnee 0.99902 ± 0.0002 0.33008 ± 0.0116 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

saut deux roros 0.99902 ± 0.0002 0.64648 ± 0.0153 0.99902 ± 0.0002 0.99902 ± 0.0002 0.44434 ± 0.0144

Piste Belizon 0.99902 ± 0.0002 0.32129 ± 0.0144 0.99902 ± 0.0002 0.99902 ± 0.0002 0.52441 ± 0.0123

Approuague 0.99902 ± 0.0002 0.65723 ± 0.0161 0.99902 ± 0.0002 0.99902 ± 0.0002 0.52832 ± 0.0135

Oyapok 0.99902 ± 0.0002 0.11328 ± 0.0119 0.99902 ± 0.0002 0.99902 ± 0.0002 0.22070 ± 0.0119

50

Table 4 continued

Tresor Camp Cisame Mt couronnee St Lavillette saut deux roros

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Mt Trinite

Tresor *

Camp Cisame 0.37598 ± 0.0135 *

Mt couronnee 0.38672 ± 0.0140 0.99902 ± 0.0002 *

St Lavillette 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

saut deux roros 0.38867 ± 0.0151 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 *

Piste Belizon 0.58398 ± 0.0150 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Approuague 0.45020 ± 0.0138 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002 0.99902 ± 0.0002

Oyapok 0.23633 ± 0.0151 0.10254 ± 0.0064 0.99902 ± 0.0002 0.99902 ± 0.0002 0.10156 ± 0.0095

51

Table 4 continued

Piste Belizon Approuague Oyapok

Pibiri

Mabura Hill

Iwokrama

Zanderij

Powakka

Mata

Brownsberg

Cropi

Organabo

Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo

St Elie

Paracou

Mt Singes

PK 72 Est

Cr Plomb

Mt Trinite

Tresor

Camp Cisame

Mt couronnee

St Lavillette

saut deux roros

Piste Belizon *

Approuague 0.99902 ± 0.0002 *

Oyapok 0.27734 ± 0.0150 0.29102 ± 0.0107 *

52

Table 5: Population pairwise differences for the trnH-psbA marker. Average number of pairwise differences between populations (πXY) is shown in green, with the darker shade of green showing significant values. The average number of pairwise differences within populations (πX) is shown in orange. The corrected average pairwise differences (Nei’s distance) (πXY-(πX+πY)/2) are shown in blue with the darker shade of blue showing significant values. Significance level P ≤ 0.05. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri 7.5 11.25 5.875 10.40625 9.64286 11.25 11.25

Mabura Hill 7.5 0 11.5 20.25 18 21 21

Iwokrama -1.79167 7.58333 7.83333 10.65625 9.85714 11.5 11.45

Zanderij 6.53125 20.125 6.61458 0.25 0.10714 0.125 0.125

Powakka 5.89286 18 5.94048 -0.01786 0 0 0

Mata 7.5 21 7.58333 0 0 0 0

Brownsberg 7.5 21 7.53333 0 0 0 0

Cropi 6.125 19.5 6.16667 -0.02083 -0.04762 0 0

Acarouany 7.5 21 7.58333 0 0 0 0

Bafog 7.5 21 7.58333 0 0 0 0

Organabo 7.5 21 7.58333 0 0 0 0

La Mana. S Lezard 7.5 21 7.58333 0 0 0 0

La Mana. S Fracas 7.5 21 7.58333 0 0 0 0

La Mana. cr Korossibo 7.5 21 7.58333 0 0 0 0

St Elie 7.5 21 7.58333 0 0 0 0

Paracou 7.5 21 7.58333 0 0 0 0

Mt Singes 7.5 21 7.58333 0 0 0 0

PK 72 Est 7.5 21 7.58333 0 0 0 0

Cr Plomb 7.5 21 7.58333 0 0 0 0

Tresor 6.27083 19.83333 6.35417 -0.09375 -0.22619 0 0

Mt Trinite 7.5 21 7.58333 0 0 0 0

Camp Cisame 7.5 21 7.58333 0 0 0 0

Approuague 7.5 21 7.58333 0 0 0 0

Nouragues 7.5 21 7.58333 0 0 0 0

St Lavillette 7.5 21 7.58333 0 0 0 0

Mt couronnee 7.5 21 7.58333 0 0 0 0

Piste Belizon 7.5 21 7.58333 0 0 0 0

saut deux roros 7.5 21 7.58333 0 0 0 0

Oyapok 7.5 21 7.58333 0 0 0 0

53

Table 5 continued

Cropi Acarouany Bafog Organabo La Mana. S Lezard

La Mana. S Fracas

La Mana. cr Korossibo St Elie

Pibiri 10.20833 11.25 11.25 11.25 11.25 11.25 11.25 11.25

Mabura Hill 19.83333 21 21 21 21 21 21 21

Iwokrama 10.41667 11.5 11.5 11.5 11.5 11.5 11.5 11.5

Zanderij 0.4375 0.125 0.125 0.125 0.125 0.125 0.125 0.125

Powakka 0.28571 0 0 0 0 0 0 0

Mata 0.33333 0 0 0 0 0 0 0

Brownsberg 0.33333 0 0 0 0 0 0 0

Cropi 0.66667 0.33333 0.33333 0.33333 0.33333 0.33333 0.33333 0.33333

Acarouany 0 0 0 0 0 0 0 0

Bafog 0 0 0 0 0 0 0 0

Organabo 0 0 0 0 0 0 0 0

La Mana. S Lezard 0 0 0 0 0 0 0 0

La Mana. S Fracas 0 0 0 0 0 0 0 0

La Mana. cr Korossibo 0 0 0 0 0 0 0 0

St Elie 0 0 0 0 0 0 0 0

Paracou 0 0 0 0 0 0 0 0

Mt Singes 0 0 0 0 0 0 0 0

PK 72 Est 0 0 0 0 0 0 0 0

Cr Plomb 0 0 0 0 0 0 0 0

Tresor -0.20833 0 0 0 0 0 0 0

Mt Trinite 0 0 0 0 0 0 0 0

Camp Cisame 0 0 0 0 0 0 0 0

Approuague 0 0 0 0 0 0 0 0

Nouragues 0 0 0 0 0 0 0 0

St Lavillette 0 0 0 0 0 0 0 0

Mt couronnee 0 0 0 0 0 0 0 0

Piste Belizon 0 0 0 0 0 0 0 0

saut deux roros 0 0 0 0 0 0 0 0

Oyapok 0 0 0 0 0 0 0 0

54

Table 5 continued

Paracou Mt Singes PK 72 Est Cr Plomb Tresor Mt Trinite Camp Cisame Approuague

Pibiri 11.25 11.25 11.25 11.25 11.60417 11.25 11.25 11.25

Mabura Hill 21 21 21 21 21.41667 21 21 21

Iwokrama 11.5 11.5 11.5 11.5 11.85417 11.5 11.5 11.5

Zanderij 0.125 0.125 0.125 0.125 1.61458 0.125 0.125 0.125

Powakka 0 0 0 0 1.35714 0 0 0

Mata 0 0 0 0 1.58333 0 0 0

Brownsberg 0 0 0 0 1.58333 0 0 0

Cropi 0.33333 0.33333 0.33333 0.33333 1.70833 0.33333 0.33333 0.33333

Acarouany 0 0 0 0 1.58333 0 0 0

Bafog 0 0 0 0 1.58333 0 0 0

Organabo 0 0 0 0 1.58333 0 0 0

La Mana. S Lezard 0 0 0 0 1.58333 0 0 0

La Mana. S Fracas 0 0 0 0 1.58333 0 0 0

La Mana. cr Korossibo 0 0 0 0 1.58333 0 0 0

St Elie 0 0 0 0 1.58333 0 0 0

Paracou 0 0 0 0 1.58333 0 0 0

Mt Singes 0 0 0 0 1.58333 0 0 0

PK 72 Est 0 0 0 0 1.58333 0 0 0

Cr Plomb 0 0 0 0 1.58333 0 0 0

Tresor 0 0 0 0 3.16667 1.58333 1.58333 1.58333

Mt Trinite 0 0 0 0 0 0 0 0

Camp Cisame 0 0 0 0 0 0 0 0

Approuague 0 0 0 0 0 0 0 0

Nouragues 0 0 0 0 0 0 0 0

St Lavillette 0 0 0 0 0 0 0 0

Mt couronnee 0 0 0 0 0 0 0 0

Piste Belizon 0 0 0 0 0 0 0 0

saut deux roros 0 0 0 0 0 0 0 0

Oyapok 0 0 0 0 0 0 0 0

55

Table 5 continued

Nouragues St Lavillette

Mt couronnee

Piste Belizon

saut deux roros Oyapok

Pibiri 11.25 11.25 11.25 11.25 11.25 11.25

Mabura Hill 21 21 21 21 21 21

Iwokrama 11.5 11.5 11.5 11.5 11.5 11.5

Zanderij 0.125 0.125 0.125 0.125 0.125 0.125

Powakka 0 0 0 0 0 0

Mata 0 0 0 0 0 0

Brownsberg 0 0 0 0 0 0

Cropi 0.33333 0.33333 0.33333 0.33333 0.33333 0.33333

Acarouany 0 0 0 0 0 0

Bafog 0 0 0 0 0 0

Organabo 0 0 0 0 0 0

La Mana. S Lezard 0 0 0 0 0 0

La Mana. S Fracas 0 0 0 0 0 0

La Mana. cr Korossibo 0 0 0 0 0 0

St Elie 0 0 0 0 0 0

Paracou 0 0 0 0 0 0

Mt Singes 0 0 0 0 0 0

PK 72 Est 0 0 0 0 0 0

Cr Plomb 0 0 0 0 0 0

Tresor 1.58333 1.58333 1.58333 1.58333 1.58333 1.58333

Mt Trinite 0 0 0 0 0 0

Camp Cisame 0 0 0 0 0 0

Approuague 0 0 0 0 0 0

Nouragues 0 0 0 0 0 0

St Lavillette 0 0 0 0 0 0

Mt couronnee 0 0 0 0 0 0

Piste Belizon 0 0 0 0 0 0

saut deux roros 0 0 0 0 0 0

Oyapok 0 0 0 0 0 0

56

Table 6: P-values of population pairwise differences of the trnH-psbA marker. PXY P-values are shown in green, with P ≤ 0.05 shown in a darker shade of green. Corrected PXY P-values are shown in blue, with P ≤ 0.05 shown in a darker shade of blue. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri * 0.03812 0.65885 0 0.00293 0.00293 0.00978

Mabura Hill 0.03812 * 0.04888 0.00293 0.00098 0 0.00587

Iwokrama 0.65885 0.04888 * 0.00293 0 0.00196 0.01564

Zanderij 0 0.00293 0.00293 * 1 1 1

Powakka 0.00293 0.00098 0 1 * 1 1

Mata 0.00293 0 0.00196 1 1 * 1

Brownsberg 0.00978 0.00587 0.01564 1 1 1 *

Cropi 0.00391 0 0.00196 0.45552 0.72239 0.56207 0.44282

Acarouany 0.06452 0.05767 0.05279 1 1 1 1

Bafog 0.00196 0.00098 0.00196 1 1 1 1

Organabo 0.05572 0.04888 0.05865 1 1 1 1

La Mana. S Lezard 1 1 1 1 1 1 1

La Mana. S Fracas 0.02346 0.00782 0.02737 1 1 1 1

La Mana. cr Korossibo 0.02737 0.02053 0.02933 1 1 1 1

St Elie 0.03128 0.02248 0.02444 1 1 1 1

Paracou 0.02639 0.02151 0.03324 1 1 1 1

Mt Singes 0 0 0.00098 1 1 1 1

PK 72 Est 0.06061 0.04594 0.06745 1 1 1 1

Cr Plomb 0.02151 0.02737 0.04106 1 1 1 1

Tresor 0 0 0.00098 0.74585 0.75171 0.3998 0.29521

Mt Trinite 0.00196 0 0.00196 1 1 1 1

Camp Cisame 0.06647 0.06256 0.03617 1 1 1 1

Approuague 0.06354 0.03128 0.08309 1 1 1 1

Nouragues 0.07722 0.04692 0.08504 1 1 1 1

St Lavillette 0.07527 0.0479 0.06745 1 1 1 1

Mt couronnee 0.02639 0.01173 0.03128 1 1 1 1

Piste Belizon 0.07038 0.0479 0.06354 1 1 1 1

saut deux roros 0.02346 0.01662 0.02053 1 1 1 1

Oyapok 0 0.00098 0.00391 1 1 1 1

57

Table 6 continued

Cropi Acarouany Bafog Organabo La Mana, S Lezard

La Mana, S Fracas

La Mana, cr Korossibo St Elie

Pibiri 0,00391 0,21114 0,00196 0,16813 1 0,02346 0,02737 0,03128

Mabura Hill 0 0,05767 0,00098 0,04888 1 0,00782 0,02053 0,02248

Iwokrama 0,00196 0,18866 0,00196 0,174 1 0,02737 0,02933 0,02444

Zanderij 0,48192 1 0,44086 1 1 1 1 1

Powakka 0,75758 1 1 1 1 1 1 1

Mata 0,43793 1 1 1 1 1 1 1

Brownsberg 1 1 1 1 1 1 1 1

Cropi * 1 0,37732 1 1 1 1 1

Acarouany 0,22776 * 1 1 1 1 1 1

Bafog 0,62268 1 * 1 1 1 1 1

Organabo 0,28055 1 1 * 1 1 1 1

La Mana, S Lezard 0 1 1 1 * 1 1 1

La Mana, S Fracas 0,38123 1 1 1 1 * 1 1

La Mana, cr Korossibo 0,36755 1 1 1 1 1 * 1

St Elie 0,38905 1 1 1 1 1 1 *

Paracou 0,34702 1 1 1 1 1 1 1

Mt Singes 0,62952 1 1 1 1 1 1 1

PK 72 Est 0,27859 1 1 1 1 1 1 1

Cr Plomb 0,33431 1 1 1 1 1 1 1

Tresor 0,55132 0,12023 0 0,1349 0,07038 0,24145 0,22776 0,20137

Mt Trinite 0,65591 1 1 1 1 1 1 1

Camp Cisame 0,24536 1 1 1 1 1 1 1

Approuague 0,22581 1 1 1 1 1 1 1

Nouragues 0,2131 1 1 1 1 1 1 1

St Lavillette 0,25806 1 1 1 1 1 1 1

Mt couronnee 0,38514 1 1 1 1 1 1 1

Piste Belizon 0,23656 1 1 1 1 1 1 1

saut deux roros 0,33138 1 1 1 1 1 1 1

Oyapok 0,56403 1 1 1 1 1 1 1

58

Table 6 continued

Paracou Mt Singes PK 72 Est Cr Plomb Tresor Mt Trinite Camp Cisame Approuague

Pibiri 0,02639 0 0,19648 0,02151 0,00293 0,00196 0,24731 0,20039

Mabura Hill 0,02151 0 0,04594 0,02737 0 0 0,06256 0,03128

Iwokrama 0,03324 0,00098 0,19355 0,04106 0,00196 0,00196 0,18475 0,22385

Zanderij 1 0,44966 1 1 1 0,3998 1 1

Powakka 1 1 1 1 1 1 1 1

Mata 1 1 1 1 1 1 1 1

Brownsberg 1 1 1 1 1 1 1 1

Cropi 1 0,37048 1 1 1 0,34409 1 1

Acarouany 1 1 1 1 1 1 1 1

Bafog 1 1 1 1 1 1 1 1

Organabo 1 1 1 1 1 1 1 1

La Mana, S Lezard 1 1 1 1 1 1 1 1

La Mana, S Fracas 1 1 1 1 1 1 1 1

La Mana, cr Korossibo 1 1 1 1 1 1 1 1

St Elie 1 1 1 1 1 1 1 1

Paracou * 1 1 1 1 1 1 1

Mt Singes 1 * 1 1 1 1 1 1

PK 72 Est 1 1 * 1 1 1 1 1

Cr Plomb 1 1 1 * 1 1 1 1

Tresor 0,19941 0 0,14858 0,18084 * 1 1 1

Mt Trinite 1 1 1 1 0 * 1 1

Camp Cisame 1 1 1 1 0,10655 1 * 1

Approuague 1 1 1 1 0,1652 1 1 *

Nouragues 1 1 1 1 0,07918 1 1 1

St Lavillette 1 1 1 1 0,12903 1 1 1

Mt couronnee 1 1 1 1 0,29814 1 1 1

Piste Belizon 1 1 1 1 0,09971 1 1 1

saut deux roros 1 1 1 1 0,24829 1 1 1

Oyapok 1 1 1 1 0,34018 1 1 1

59

Table 6 continued

Nouragues St Lavillette

Mt couronnee

Piste Belizon

saut deux roros Oyapok

Pibiri 0,22678 0,17791 0,02639 0,21017 0,02346 0

Mabura Hill 0,04692 0,0479 0,01173 0,0479 0,01662 0,00098

Iwokrama 0,20137 0,21701 0,03128 0,21114 0,02053 0,00391

Zanderij 1 1 1 1 1 1

Powakka 1 1 1 1 1 1

Mata 1 1 1 1 1 1

Brownsberg 1 1 1 1 1 1

Cropi 1 1 1 1 1 0,43597

Acarouany 1 1 1 1 1 1

Bafog 1 1 1 1 1 1

Organabo 1 1 1 1 1 1

La Mana, S Lezard 1 1 1 1 1 1

La Mana, S Fracas 1 1 1 1 1 1

La Mana, cr Korossibo 1 1 1 1 1 1

St Elie 1 1 1 1 1 1

Paracou 1 1 1 1 1 1

Mt Singes 1 1 1 1 1 1

PK 72 Est 1 1 1 1 1 1

Cr Plomb 1 1 1 1 1 1

Tresor 1 1 1 1 1 1

Mt Trinite 1 1 1 1 1 1

Camp Cisame 1 1 1 1 1 1

Approuague 1 1 1 1 1 1

Nouragues * 1 1 1 1 1

St Lavillette 1 * 1 1 1 1

Mt couronnee 1 1 * 1 1 1

Piste Belizon 1 1 1 * 1 1

saut deux roros 1 1 1 1 * 1

Oyapok 1 1 1 1 1 *

60

Table 7: Population pairwise differences for the ycf6-trnC marker. Average number of pairwise differences between populations (πXY) is shown in green, with the darker shade of green showing significant values. The average number of pairwise differences within populations (πX) is shown in orange. The corrected average pairwise differences (Nei’s distance) (πXY-(πX+πY)/2) are shown in blue with the darker shade of blue showing significant values. Significance level P ≤ 0.05. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri 129.6667 290.8333 160.8571 172.2593 141 147.0556 188.4286

Mabura Hill 86.06667 279.8667 246.4286 231.037 225.1333 229.6667 221.3571

Iwokrama 27.42857 37.9 137.1905 108.746 77.52857 88.47619 124.6122

Zanderij 67.88426 51.56204 0.60913 79.08333 39.72222 53.68519 87.60317

Powakka 75.96667 85 8.73333 -0.01944 0.4 15.5 63.7

Mata 66.88889 74.4 4.54762 -1.18981 -0.03333 30.66667 76.35714

Brownsberg 66.52381 24.35238 -1.05442 -9.00992 6.42857 3.95238 114.1429

Cropi 74.72222 84.78889 9.14286 1.21759 1.5 0.91667 7.28571

Organabo 77.5 86.06667 11.11905 2.79167 3 2.5 9.07143

Bafog 74.54762 84.94762 9.44558 1.99802 2.42857 1.66667 8.19388

La Mana. Saut Lezard 76.16667 85.4 9.40476 0.90278 1 0.66667 7.21429

La Mana. Saut Fracas 73.16667 81.31667 8.75 0.98611 2.16667 1.41667 6.77381

La Mana. cr Korossibo 76.16667 85.06667 9.40476 0.90278 1 0.66667 7.21429

St Elie 85.77778 63.06667 25.19048 22.55093 31.76667 29.88889 25.52381

Paracou 77.16667 85.73333 10.2619 1.79167 2 1.33333 8.07143

Mt Singes 76.16667 85.06667 9.40476 0.90278 1 0.66667 7

PK 72 Est 73.33333 81.31667 8.7619 0.84722 2 1.25 6.85714

Cr Plomb 77.16667 85.73333 10.2619 1.79167 2 1.33333 8.07143

Mt Trinite 72.94643 74.97143 9.42262 2.89087 4.94643 3.75893 8.30357

Tresor 74.25556 78.77222 8.94127 2.21389 4.12222 2.40556 7.60794

Camp Cisame 76.83333 84.65 10.19048 1.68056 2 1.16667 7.71429

Mt couronnee 77 84.98333 10.11905 1.79167 2 1.33333 8.07143

St Lavillette 77.16667 85.4 10.2619 1.79167 2 1.33333 8.07143

saut deux roros 77 84.31667 9.97619 1.625 2 1.25 7.78571

Piste Belizon 76.33333 84.4 9.83333 1.625 2 1.33333 7.85714

Approuague 76.66667 84.81667 10.04762 1.79167 2 1.33333 8.07143

Oyapok 77.16667 85.6 10.2619 1.79167 2 1.33333 8.07143

61

Table 7 continued

Cropi Organabo Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo St Elie Paracou

Pibiri 140.5556 142.3333 141.2381 141 138.3333 141.5 181.1111 142

Mabura Hill 225.7222 226 226.7381 225.3333 221.5833 225.5 233.5 225.6667

Iwokrama 78.7381 79.71429 79.89796 78 77.67857 78.5 124.2857 78.85714

Zanderij 41.75926 42.33333 43.39683 40.44444 40.86111 40.94444 92.59259 41.33333

Powakka 2.7 3.2 4.48571 1.2 2.7 1.7 62.46667 2.2

Mata 17.25 17.83333 18.85714 16 17.08333 16.5 75.72222 16.66667

Brownsberg 65.35714 66.14286 67.12245 64.28571 64.17857 64.78571 113.0952 65.14286

Cropi 2 3.83333 5.04762 1.83333 3.33333 2.33333 62.77778 2.83333

Organabo 2.83333 0 5.28571 2 2.5 2.5 63.33333 3

Bafog 2.19048 3.42857 3.71429 3.28571 4.75 3.78571 63.85714 4.28571

La Mana. Saut Lezard 0.83333 2 1.42857 0 1.5 0.5 61.33333 1

La Mana. Saut Fracas 2 2.16667 2.55952 1.16667 0.66667 1.75 53 0.5

La Mana. cr Korossibo 0.83333 2 1.42857 0 0.91667 1 61.5 1.5

St Elie 31.27778 32.83333 31.5 30.83333 22.16667 30.5 61 61.66667

Paracou 1.83333 3 2.42857 1 0.16667 1 31.16667 0

Mt Singes 0.83333 2 1.42857 0 0.41667 -0.5 30.33333 1

PK 72 Est 1.83333 2 2.42857 1 0.16667 0.75 20.5 0

Cr Plomb 1.83333 3 2.42857 1 0.16667 1 31.16667 0

Mt Trinite 4.65476 5.94643 5.10714 3.94643 0.1756 3.82143 3.40476 3.44643

Tresor 3.85556 5.22222 4.30794 3.22222 2.48889 3.12222 31.95556 2.42222

Camp Cisame 1.83333 2 2.42857 1 -0.58333 0.5 30.33333 0

Mt couronnee 1.83333 1.5 2.42857 1 0.16667 1 30.16667 0

St Lavillette 1.83333 1 2.42857 1 0.16667 1 30.5 0

saut deux roros 1.75 1 2.42857 1 -0.33333 0.5 30.5 0

Piste Belizon 1.83333 3 2.42857 1 0.16667 1 31.16667 0

Approuague 1.83333 3 2.42857 1 -0.33333 0.5 30.66667 0

Oyapok 1.83333 3 2.42857 1 0.06667 0.8 31.03333 0

62

Table 7 continued

Mt Singes PK 72 Est Cr Plomb Mt Trinite Tresor Camp Cisame

Mt couronnee

St Lavillette

Pibiri 142 138.1667 142 158.5833 150.4667 143.1667 141.8333 142

Mabura Hill 226 221.25 225.6667 235.7083 230.0833 226.0833 224.9167 225.3333

Iwokrama 79 77.35714 78.85714 98.82143 88.91429 80.28571 78.71429 78.85714

Zanderij 41.44444 40.38889 41.33333 63.23611 53.13333 42.72222 41.33333 41.33333

Powakka 2.2 2.2 2.2 25.95 15.7 3.7 2.2 2.2

Mata 17 16.58333 16.66667 39.89583 29.11667 18 16.66667 16.66667

Brownsberg 65.07143 63.92857 65.14286 86.17857 76.05714 66.28571 65.14286 65.14286

Cropi 2.83333 2.83333 2.83333 26.45833 16.23333 4.33333 2.83333 2.83333

Organabo 3 2 3 26.75 16.6 3.5 1.5 1

Bafog 4.28571 4.28571 4.28571 27.76786 17.54286 5.78571 4.28571 4.28571

La Mana. Saut Lezard 1 1 1 24.75 14.6 2.5 1 1

La Mana. Saut Fracas 1.75 0.5 0.5 21.3125 14.2 1.25 0.5 0.5

La Mana. cr Korossibo 1 1.25 1.5 25.125 15 2.5 1.5 1.5

St Elie 61.83333 51 61.66667 54.70833 73.83333 62.33333 60.66667 61

Paracou 2 0 0 24.25 13.8 1.5 0 0

Mt Singes 2 1.75 2 25.5625 15.5 2.5 2 2

PK 72 Est 0.75 0 0 20.25 13.7 1.25 0 0

Cr Plomb 1 0 0 24.25 13.8 1.5 0 0

Mt Trinite 3.75893 -0.55357 3.44643 41.60714 37.475 25.4375 23.875 24

Tresor 3.12222 2.32222 2.42222 5.29365 22.75556 15.2 13.8 13.8

Camp Cisame 0 -0.25 0 3.13393 2.32222 3 1.5 1.5

Mt couronnee 1 0 0 3.07143 2.42222 0 0 0

St Lavillette 1 0 0 3.19643 2.42222 0 0 0

saut deux roros 0.5 -0.5 0 3.19643 2.32222 -0.5 0 0

Piste Belizon 1 0 0 3.44643 -0.07778 0 0 0

Approuague 0.5 -0.5 0 3.32143 2.32222 -0.5 -0.25 0

Oyapok 0.8 -0.1 0 3.39643 2.38222 -0.2 0 0

63

Table 7 continued

saut deux roros

Piste Belizon Approuague Oyapok

Pibiri 143.3333 144.6667 142.5 142.2

Mabura Hill 225.75 227.8333 225.75 225.7333

Iwokrama 80.07143 81.92857 79.64286 79.05714

Zanderij 42.66667 44.66667 42.33333 41.53333

Powakka 3.7 5.7 3.2 2.4

Mata 18.08333 20.16667 17.66667 16.86667

Brownsberg 66.35714 68.42857 66.14286 65.34286

Cropi 4.25 6.33333 3.83333 3.03333

Organabo 2.5 6.5 4 3.2

Bafog 5.78571 7.78571 5.28571 4.48571

La Mana. Saut Lezard 2.5 4.5 2 1.2

La Mana. Saut Fracas 1.5 4 1 0.6

La Mana. cr Korossibo 2.5 5 2 1.5

St Elie 62.5 65.16667 62.16667 61.73333

Paracou 1.5 3.5 1 0.2

Mt Singes 3 5.5 2.5 2

PK 72 Est 1 3.5 0.5 0.1

Cr Plomb 1.5 3.5 1 0.2

Mt Trinite 25.5 27.75 25.125 24.4

Tresor 15.2 14.8 14.7 13.96

Camp Cisame 2.5 5 2 1.5

Mt couronnee 1.5 3.5 0.75 0.2

St Lavillette 1.5 3.5 1 0.2

saut deux roros 3 5 1.75 1.5

Piste Belizon 0 7 4.5 3.7

Approuague -0.75 0 2 1

Oyapok -0.2 0 -0.2 0.4

64

Table 8: P-values of population pairwise differences of the ycf6-trnC marker. PXY P-values are shown in green, with P ≤ 0.05 shown in a darker shade of green. Corrected PXY P-values are shown in blue, with P ≤ 0.05 shown in a darker shade of blue. Note that, due to its size, the table is spread across four pages.

Pibiri Mabura Hill Iwokrama Zanderij Powakka Mata Brownsberg

Pibiri * 0.11828 0.30303 0.13783 0.00196 0.01271 0.13685

Mabura Hill 0.01955 * 0.04203 0.01173 0 0.00293 0.08504

Iwokrama 0.09286 0.07331 * 0.23851 0.00293 0.65396 0.40176

Zanderij 0.00489 0.02835 0.51026 * 0.26197 0.57674 0.34702

Powakka 0.00196 0 0.00293 0.51711 * 0.19746 0.01466

Mata 0.01271 0.00293 0.24438 0.68426 0.89541 * 0.72043

Brownsberg 0.00587 0.13001 0.40176 0.53568 0.01369 0.37341 *

Cropi 0.00782 0.0088 0.02248 0 0 0.0176 0.19355

Organabo 1 1 1 1 1 1 0

Bafog 0.00684 0 0.00098 0 0 0.00293 0.00196

La Mana. Saut Lezard 1 1 1 1 1 1 0

La Mana. Saut Fracas 0.03324 0.02639 0.14956 0.07722 0 0.0176 0.15347

La Mana. cr Korossibo 0.09189 0.1173 0.07527 0.0176 0.02542 0.07527 0.02053

St Elie 0.09091 0.03715 0.09873 0.0567 0.00489 0.03324 0.08993

Paracou 1 1 1 1 1 1 0

Mt Singes 0.0958 0.09189 0.03519 0.01662 0.02248 0.11926 0.02542

PK 72 Est 0.09971 0.16422 0.18182 0.04008 0.01857 0.09677 0.17986

Cr Plomb 1 1 1 1 1 1 0

Mt Trinite 0.00684 0.00196 0.02053 0.17791 0 0.22874 0.10362

Tresor 0.00098 0.00098 0.01564 0.01955 0 0.06745 0.07331

Camp Cisame 0.1173 0.14663 0.03519 0.00978 0.00391 0.12219 0.02835

Mt couronnee 0.10264 0.12023 0.06647 0.02053 0.01369 0.08602 0.03617

St Lavillette 1 1 1 1 1 1 0

saut deux roros 0.12414 0.16031 0.05474 0.00587 0.00587 0.08016 0.02444

Piste Belizon 0.1085 0.14076 0.05963 0.04106 0.01271 0.07331 0.04888

Approuague 0.09677 0.1349 0.0694 0.02053 0.01075 0.08993 0.01075

Oyapok 0.02248 0.00196 0.04106 0.00098 0 0.0088 0.18671

65

Table 8 continued

Cropi Organabo Bafog

La Mana. Saut Lezard

La Mana. Saut Fracas

La Mana. cr Korossibo St Elie Paracou

Pibiri 0.00782 1 0.00684 1 0.03324 0.16422 0.09091 1

Mabura Hill 0.0088 1 0 1 0.16422 0.90029 0.5347 1

Iwokrama 0.59824 1 0.00098 1 0.95112 0.85337 0.37634 1

Zanderij 0.39003 1 0.43695 1 0.35093 0.16618 0.31769 1

Powakka 0 1 0 1 0 0.04594 0.00489 1

Mata 0.0176 1 0.00587 1 0.43304 0.31672 0.03324 1

Brownsberg 0.46139 1 0.00293 1 0.68817 0.7566 0.3392 1

Cropi * 1 0.00196 1 0.00489 0.22776 0.02639 1

Organabo 1 * 0.24927 1 0.18964 0.33333 0.24047 1

Bafog 0 0.24927 * 1 0.0567 0.29326 0.01662 1

La Mana. Saut Lezard 1 1 1 * 0.22092 1 0.75171 1

La Mana. Saut Fracas 0.00489 0.18964 0.00196 0.22092 * 0.07234 0.09971 1

La Mana. cr Korossibo 0.04692 0.33333 0.06745 1 0.12805 * 0.32454 1

St Elie 0.02639 0.24047 0.01662 0.75171 0.10753 0.32454 * 1

Paracou 1 1 1 1 1 1 1 *

Mt Singes 0.04008 0.71945 0.07234 1 0.2913 1 0.2737 0.64321

PK 72 Est 0.02737 0.34506 0.02542 0.33724 0.46725 0.36266 0.3304 1

Cr Plomb 1 1 1 1 1 1 1 1

Mt Trinite 0.16422 0.34995 0.00782 0.77615 0.4477 0.05963 0.28935 1

Tresor 0 0.46921 0.00098 0.90127 0.03812 0.15249 0.00782 1

Camp Cisame 0.04106 0.69208 0.0303 1 0.9306 0.65689 0.3001 1

Mt couronnee 0.04203 0.36168 0.01564 0.28543 0.49365 0.31867 0.41838 1

St Lavillette 1 1 1 1 1 1 1 1

saut deux roros 0.05767 1 0.02248 1 0.72141 0.67449 0.3001 1

Piste Belizon 0.04008 0.65494 0.01857 0.63734 0.3001 0.33529 0.20821 1

Approuague 0.04399 0.3089 0.02151 0.66178 0.66862 0.70577 0.38221 1

Oyapok 0 0.18671 0.00196 0.35484 0.38905 0.10362 0.03715 1

66

Table 8 continued

Mt Singes PK 72 Est Cr Plomb Mt Trinite Tresor Camp Cisame

Mt couronnee

St Lavillette

Pibiri 0.22092 0.19355 1 0.00684 0.00098 0.18671 0.17986 1

Mabura Hill 0.7957 0.88563 1 0.00196 0.00098 0.87586 0.91105 1

Iwokrama 0.81036 0.93939 1 0.00489 0.01466 0.67937 0.89345 1

Zanderij 0.20919 0.47703 1 0.49853 0.22092 0.25709 0.16422 1

Powakka 0.0303 0.01857 1 0 0 0.00391 0.01369 1

Mata 0.3216 0.30499 1 0.22092 0.12219 0.31378 0.21799 1

Brownsberg 0.72336 0.91202 1 0.0303 0.07331 0.67449 0.73314 1

Cropi 0.22581 0.14272 1 0.16618 0.41056 0.04106 0.13685 1

Organabo 0.71945 0.34506 1 0.34995 0.46921 0.69208 0.36168 1

Bafog 0.24536 0.24047 1 0.20039 0.43695 0.2346 0.26295 1

La Mana. Saut Lezard 1 0.33724 1 0.77615 0.90127 1 0.28543 1

La Mana. Saut Fracas 0.06061 1 1 0.96579 0.73607 0.37048 1 1

La Mana. cr Korossibo 1 0.36266 1 0.53666 0.76246 0.65689 0.31867 1

St Elie 0.2737 0.3304 1 0.26491 0.02639 0.3001 0.41838 1

Paracou 0.64321 1 1 1 1 1 1 1

Mt Singes * 0.31183 1 0.43304 0.50147 1 0.3304 1

PK 72 Est 0.31183 * 1 1 1 1 1 1

Cr Plomb 1 1 * 1 1 1 1 1

Mt Trinite 0.07429 0.60997 1 * 0.07234 0.35973 1 1

Tresor 0.15543 0.19453 1 0.00098 * 0.7869 1 1

Camp Cisame 1 1 1 0.58358 0.13978 * 1 1

Mt couronnee 0.3304 1 1 0.60313 0.2043 1 * 1

St Lavillette 1 1 1 1 1 1 1 *

saut deux roros 0.64712 1 1 0.5259 0.17595 1 1 1

Piste Belizon 0.3216 1 1 0.26393 0.47019 1 1 1

Approuague 0.65787 1 1 0.35093 0.1828 1 1 1

Oyapok 0.11339 1 1 0.21408 0.02151 0.75464 1 1

67

Table 8 continued

saut deux roros

Piste Belizon Approuague Oyapok

Pibiri 0.2434 0.19844 0.19941 0.02248

Mabura Hill 0.86022 0.77224 0.86608 0.0479

Iwokrama 0.75367 0.82014 0.7957 0.5259

Zanderij 0.19648 0.23754 0.19159 0.39003

Powakka 0.00587 0.01271 0.01075 0

Mata 0.30987 0.27468 0.31183 0.4653

Brownsberg 0.55914 0.56696 0.51515 0.42913

Cropi 0.05767 0.04008 0.04399 0

Organabo 1 0.65494 0.3089 0.18671

Bafog 0.24731 0.11437 0.30596 0.0088

La Mana. Saut Lezard 1 0.63734 0.66178 0.35484

La Mana. Saut Fracas 0.38807 0.39101 0.61388 0.30205

La Mana. cr Korossibo 0.67449 0.33529 0.70577 0.10362

St Elie 0.3001 0.20821 0.17302 0.03715

Paracou 1 1 1 1

Mt Singes 0.64712 0.3216 0.65787 0.11339

PK 72 Est 1 1 1 1

Cr Plomb 1 1 1 1

Mt Trinite 0.38807 0.56598 0.55132 0.91691

Tresor 0.74194 0.84164 0.86119 0.63832

Camp Cisame 1 1 1 0.10264

Mt couronnee 1 1 1 1

St Lavillette 1 1 1 1

saut deux roros * 1 1 0.10166

Piste Belizon 1 * 1 0.27761

Approuague 1 1 * 0.2913

Oyapok 0.78788 0.2434 0.7654 *