Sensitivity of seed germination to water stress in high-altitude populations of a threatened palm species.

Divergence in seed germination patterns among populations of the same species is important for understanding plant responses to environmental gradients and potential plant sensitivity to climate change. In order to test responses to flooding and decreasing water potentials, over 3 years we germinated and grew seeds from three habitats of Euterpe edulis Mart. occurring along an altitudinal gradient. Seed germination and root growth were evaluated under different water availability treatments: control, flood, -0.4 MPa, -0.8 MPa, in the years 2012, 2013 and 2014, and in the final year of the experiment (2014) at -1.0 MPa and -1.5 MPa. Seeds from the montane habitat did not germinate in the flooding treatment. Seed germination of all three habitats decreased in the -1.5 MPa treatment and the montane habitat had lowest germination in this treatment. Time required for half of the seeds to germinate increased up to -0.8 MPa. Seeds from montane habitats germinated more slowly in all treatments. The only difference in seed germination synchrony was an increase in the submontane population under the flooding treatment. However, synchrony decreased at the lowest water potentials. Roots of the montane population were more vigorous in most treatments, except at -0.8 MPa. The unusual ability of these seeds to germinate at low water potentials might be related to early seed germination at the onset of the rainy season, which potentially decreases seed predation pressure. Seeds of the montane population were more sensitive to both types of water stress. A predicted increase in the frequency and intensity of extreme high rainfall or drought events may predispose early stages of this population to adverse factors that might negatively affect population viability with elevational in future climate change scenarios.

Genomic Traces of the Fruit Fly Anastrepha obliqua Associated with Its Polyphagous Nature.

Anastrepha obliqua (Macquart) (Diptera: Tephritidae) is an important pest in the neotropical region. It is considered a polyphagous insect, meaning it infests plants of different taxonomic families and readily colonizes new host plants. The change to new hosts can lead to diversification and the formation of host races. Previous studies investigating the effect of host plants on population structure and selection in Anastrepha obliqua have focused on the use of data from the mitochondrial DNA sequence and microsatellite markers of nuclear DNA, and there are no analyses at the genomic level. To better understand this issue, we used a pooled restriction site-associated DNA sequencing (pooled RAD-seq) approach to assess genomic differentiation and population structure across sympatric populations of Anastrepha obliqua that infest three host plants-Spondias purpurea (red mombin), Mangifera indica (mango) of the family Anacardiaceae and Averrhoa carambola (carambola) of the family Oxalidaceae-in sympatric populations of the species Anastrepha obliqua of Inter-Andean Valley of the Cauca River in southwestern Colombia. Our results show genomic differentiation of populations from carambola compared to mango and red mombin populations, but the genetic structure was mainly established by geography rather than by the host plant. On the other hand, we identified 54 SNPs in 23 sequences significantly associated with the use of the host plant. Of these 23 sequences, we identified 17 candidate genes and nine protein families, of which four protein families are involved in the nutrition of these flies. Future studies should investigate the adaptive processes undergone by phytophagous insects in the Neotropics, using fruit flies as a model and state-of-the-art molecular tools.

Genetic diversity and differentiation of nine populations of the hard clam (Meretrix petechialis) assessed by EST-derived microsatellites

BACKGROUND: Meretrix petechialis is one of the commercially important marine bivalves. In this study, we selected six highly polymorphic EST-derived microsatellite markers to assess the genetic diversity and population differentiation on nine wild populations of Meretrix petechialis. RESULTS: The number of alleles detected per-locus ranged from 4 to 30 (mean NA = 27.5) with a total of 165 alleles. The mean value of observed and expected heterozygosities varied from 0.717 to 0.861 and from 0.797 to 0.856, respectively. Meanwhile, the result of Neighbor-joining and overall FST = 0.214 (P < 0.01) reveled that M. petechialis populations from GX are the farthest populations, a certain degree of genetic variation among individuals in each population and the genetic differentiation is significant. CONCLUSIONS: GX population has high genetic diversity among individual, and there are certain differences in genetic characteristics among different populations. This study will provide a basis for the domestication and cultivation of genetic diversity of M. petechialis population and the protection of clam germplasm resources.

Exploring the drivers of population structure across desert snakes can help to link micro and macroevolution.

To understand the underlying mechanisms generating population genetic divergence and structure is a critical step towards understanding how biodiversity evolves at both micro- and macroevolutionary scales. At the population-level, geographic isolation as well as adaptation to local environmental conditions can generate different patterns of spatial genetic variation among populations. Specific organismal traits as well as the characteristics of the environment might influence the process under which populations become spatially structured. In a From the Cover article in this issue of Molecular Ecology, Myers et al. (2019) present an integrative approach to investigate if the Cochise filter barrier (CFB), lying between the Sonoran and Chihuahuan Deserts, and the surrounding river networks were relevant in driving the population structure of 13 snake species. While local environmental conditions seem to predominantly contribute to lineage divergence, traditionally studied vicariant barriers seem to have played a minor role in shaping population structure across the studied species. This study brings insights into how population-level processes could contribute to the formation of incipient species, which ultimately might affect the speciation rates measured at macroevolutionary scales. Hence, Myers et al. (2019) not only represents an integrative study aiming to understand the drivers of population genetic divergence, but also a potentially important contribution to our ongoing challenge in linking micro- and macroevolution.

Flood tolerance in two tree species that inhabit both the Amazonian floodplain and the dry Cerrado savanna of Brazil.

Comparing plants of the same species thriving in flooded and non-flooded ecosystems helps to clarify the interplay between natural selection, phenotypic plasticity and stress adaptation. We focussed on responses of seeds and seedlings of Genipa americana and Guazuma ulmifolia to substrate waterlogging or total submergence. Both species are commonly found in floodplain forests of Central Amazonia and in seasonally dry savannas of Central Brazil (Cerrado). Although seeds of Amazonian and Cerrado G. americana were similar in size, the germination percentage of Cerrado seeds was decreased by submergence (3 cm water) and increased in Amazonian seeds. The seeds of Amazonian G. ulmifolia were heavier than Cerrado seeds, but germination of both types was unaffected by submergence. Three-month-old Amazonian and Cerrado seedlings of both species survived 30 days of waterlogging or submersion despite suffering significant inhibition in biomass especially if submerged. Shoot elongation was also arrested. Submersion triggered chlorosis and leaf abscission in Amazonian and Cerrado G. ulmifolia while waterlogging did so only in Cerrado seedlings. During 30 days of re-exposure to non-flooded conditions, G. ulmifolia plants that lost their leaves produced a replacement flush. However, they attained only half the plant dry mass of non-flooded plants. Both submerged and waterlogged G. americana retained their leaves. Consequently, plant dry mass after 30 days recovery was less depressed by these stresses than in G. ulmifolia. Small amounts of cortical aerenchyma were found in roots 2 cm from the tip of well-drained plants. The amount was increased by flooding. Waterlogging but not submergence promoted hypertrophy of lenticels at the stem base of both species and adventitious rooting in G. ulmifolia. Despite some loss of performance in dryland plants, flood tolerance traits were present in wetland and dryland populations of both species. They are part of an overall stress-response potential that permits flexible acclimation to locally flooded conditions.

The Effect of Balancing Selection on Population Differentiation: A Study with HLA Genes.

Balancing selection is defined as a class of selective regimes that maintain polymorphism above what is expected under neutrality. Theory predicts that balancing selection reduces population differentiation, as measured by FST. However, balancing selection regimes in which different sets of alleles are maintained in different populations could increase population differentiation. To tackle the connection between balancing selection and population differentiation, we investigated population differentiation at the HLA genes, which constitute the most striking example of balancing selection in humans. We found that population differentiation of single nucleotide polymorphisms (SNPs) at the HLA genes is on average lower than that of SNPs in other genomic regions. We show that these results require using a computation that accounts for the dependence of FST on allele frequencies. However, in pairs of closely related populations, where genome-wide differentiation is low, differentiation at HLA is higher than in other genomic regions. Such increased population differentiation at HLA genes for recently diverged population pairs was reproduced in simulations of overdominant selection, as long as the fitness of the homozygotes differs between the diverging populations. The results give insight into a possible "divergent overdominance" mechanism for the nature of balancing selection on HLA genes across human populations.

Possible glimpses into early speciation: the effect of ovarian fluid on sperm velocity accords with post-copulatory isolation between two guppy populations.

Identifying mechanisms of reproductive isolation is key to understanding speciation. Among the putative mechanisms underlying reproductive isolation, sperm-female interactions (post-mating-prezygotic barriers) are arguably the hardest to identify, not least because these are likely to operate at the cellular or molecular level. Yet sperm-female interactions offer great potential to prevent the transfer of genetic information between different populations at the initial stages of speciation. Here, we provide a preliminary test for the presence of a putative post-mating-prezygotic barrier operating between three populations of Trinidadian guppies (Poecilia reticulata), an internally fertilizing fish that inhabits streams with different levels of connectivity across Trinidad. We experimentally evaluate the effect of female ovarian fluid on sperm velocity (a predictor of competitive fertilization success) according to whether males and females were from the same (native) or different (foreign) populations. Our results reveal the potential for ovarian fluid to act as a post-mating-prezygotic barrier between two populations from different drainages, but also that the strength of this barrier is different among populations. This result may explain the previous finding that, in some populations, sperm from native males have precedence over foreign sperm, which could eventually lead to reproductive isolation between these populations.

Genetic assessment for the endangered black lion tamarin Leontopithecus chrysopygus (Mikan, 1823), Callitrichidae, Primates.

This is the first study analyzing genetic diversity in captive individuals of the endangered black lion tamarin, Leontopithecus chrysopygus, and also comparing genetic diversity parameters between wild populations and captive groups using the same set of molecular markers. We evaluated genetic diversity and differentiation for the Brazilian and European captive groups and a wild population through 15 polymorphic microsatellite markers. The genetic diversity levels were similar among Brazilian captive, European captive and wild animals from the National Forest of Capão Bonito. Expected heterozygosity showed values ranging from 0.403 to 0.462, and significant differences were not observed among the populations. Different allele frequencies were observed among the groups, which showed the presence of distinct private alleles. The PCoA analysis evidenced three main clusters suggesting that the captive Brazilian and European groups are markedly differentiated both from one another and from the wild population of Capão Bonito. Likewise, the most likely number of genetic clusters (K) revealed by Structure was three. Such a structure is probably the result of the strength of drift and non-random reproduction in these small and isolated groups. Despite this differentiation, all groups still have similar genetic diversity levels, comparable to other callitrichids. The data obtained herein are important to increasing knowledge of the genetics of tamarins and supporting breeding programs to prevent loss of genetic diversity and inbreeding depression.

Migration by seed dispersal of ACCase-inhibitor-resistant Avena fatua in north-western Mexico.

BACKGROUND: Biotypes of Avena fatua resistant to ACCase-inhibiting herbicides have been reported in the States of Baja California (BC) and Sonora (SON), Mexico. We hypothesised that resistant biotypes present in SON (Valle de Hermosillo and Valle del Yaqui) are derived from a resistant population from BC (Valle de Mexicali) via gene flow through the transport and exchange of contaminated wheat seed. This study aimed to determine (1) the resistance of A. fatua to ACCase-inhibiting herbicides in populations from BC and SON, (2) the mutation at the site of action and (3) the genetic structure and gene flow among populations. RESULTS: DNA sequencing showed that all biotypes shared the same mutation (Leu × Ile at codon 1781). Microsatellites showed evidence of a genetic bottleneck in SON, and spatial analysis of molecular variance grouped one biotype from the Valle de Mexicali with two biotypes from the Valle de Hermosillo. Migration analysis suggested gene flow from the Valle de Mexicali to the Valle de Hermosillo, but not to the Valle del Yaqui. CONCLUSIONS: The presence of resistant biotypes of A. fatua in the Valle de Hermosillo, SON, are likely derived from seeds from BC, possibly through the transport of contaminated wheat seeds. © 2016 Society of Chemical Industry.

The imperiled fish fauna in the Nicaragua Canal zone.

Large-scale infrastructure projects commonly have large effects on the environment. The planned construction of the Nicaragua Canal will irreversibly alter the aquatic environment of Nicaragua in many ways. Two distinct drainage basins (San Juan and Punta Gorda) will be connected and numerous ecosystems will be altered. Considering the project's far-reaching environmental effects, too few studies on biodiversity have been performed to date. This limits provision of robust environmental impact assessments. We explored the geographic distribution of taxonomic and genetic diversity of freshwater fish species (Poecilia spp., Amatitlania siquia, Hypsophrys nematopus, Brycon guatemalensis, and Roeboides bouchellei) across the Nicaragua Canal zone. We collected population samples in affected areas (San Juan, Punta Gorda, and Escondido drainage basins), investigated species composition of 2 drainage basins and performed genetic analyses (genetic diversity, analysis of molecular variance) based on mitochondrial cytb. Freshwater fish faunas differed substantially between drainage basins (Jaccard similarity = 0.33). Most populations from distinct drainage basins were genetically differentiated. Removing the geographic barrier between these basins will promote biotic homogenization and the loss of unique genetic diversity. We found species in areas where they were not known to exist, including an undescribed, highly distinct clade of live bearing fish (Poecilia). Our results indicate that the Nicaragua Canal likely will have strong impacts on Nicaragua's freshwater biodiversity. However, knowledge about the extent of these impacts is lacking, which highlights the need for more thorough investigations before the environment is altered irreversibly.

Woody climbers show greater population genetic differentiation than trees: Insights into the link between ecological traits and diversification.

The climbing habit is a key innovation in plants: climbing taxa have higher species richness than nonclimbing sister groups. We evaluated the hypothesis that climbing plant species show greater among-population genetic differentiation than nonclimber species. We compared the among-population genetic distance in woody climbers (eight species, 30 populations) and trees (seven species, 29 populations) coexisting in nine communities in a temperate rainforest. We also compared within-population genetic diversity in co-occurring woody climbers and trees in two communities. Mean genetic distance between populations of climbers was twice that of trees. Isolation by distance (increase in genetic distance with geographic distance) was greater for climbers. Climbers and trees showed similar within-population genetic diversity. Our longevity estimate suggested that climbers had shorter generation times, while other biological features often associated with diversification (dispersal and pollination syndromes, mating system, size, and metabolic rate) did not show significant differences between groups. We hypothesize that the greater population differentiation in climbers could result from greater evolutionary responses to local selection acting on initially higher within-population genetic diversity, which could be driven by neutral processes associated with shorter generation times. Increased population genetic differentiation could be incorporated as another line of evidence when testing for key innovations.

Morphological variation in vanishing Mexican desert fishes of the genus Characodon (Goodeidae).

This study investigated a rapidly vanishing group of fishes in the genus Characodon (including Characodon lateralis and Characodon audax) from the upper Rio Mezquital drainage in Mexico. Using specimens from museum collections, morphological variation was assessed to quantify body shape differentiation among historic (i.e. extirpated and extant) collection sites. In both sexes, body shape (particularly head shape, the proportion of the caudal peduncle and the position and size of dorsal and anal fins) varied significantly among populations and species. Variation among collection sites could at least partially be attributed to geography, as the presence of distinct hydrographic units and a major waterfall coincided with major body shape differences. These results are discussed in the light of previously published molecular genetic analyses, as they have direct implications for taxonomic problems and the need for conservation measures for these endangered fishes.

Differentiating cave Aspidoras catfish from a karst area of Central Brazil, upper rio Tocantins basin (Siluriformes: Callichthyidae)

Two cave populations of Aspidoras albater catfish with reduced eyes and pigmentation were recently found near the city of Posse, São Domingos karst area, Goiás State, representing the first known case of a troglomorphic callichthyid catfish. Cave specimens are described and compared to epigean specimens of A. albater, but morphometric differences between the epigean and the cave populations are not adequate to delimit as distinct species.

Duas populações de Aspidoras albater com olhos e pigmentação reduzidos foram descobertas recentemente em cavernas do município de Posse, área cárstica de São Domingos, Goiás, representando o primeiro caso conhecido de Callichthyidae troglomórfico. Os espécimes cavernícolas são descritos e comparados com espécimes epígeos de A. albater, mas as diferenças morfométricas entre eles não são suficientes para delimitá-los como espécies distintas.

Differentiating cave Aspidoras catfish from a karst area of Central Brazil, upper rio Tocantins basin (Siluriformes: Callichthyidae)

Two cave populations of Aspidoras albater catfish with reduced eyes and pigmentation were recently found near the city of Posse, São Domingos karst area, Goiás State, representing the first known case of a troglomorphic callichthyid catfish. Cave specimens are described and compared to epigean specimens of A. albater, but morphometric differences between the epigean and the cave populations are not adequate to delimit as distinct species.(AU)

Duas populações de Aspidoras albater com olhos e pigmentação reduzidos foram descobertas recentemente em cavernas do município de Posse, área cárstica de São Domingos, Goiás, representando o primeiro caso conhecido de Callichthyidae troglomórfico. Os espécimes cavernícolas são descritos e comparados com espécimes epígeos de A. albater, mas as diferenças morfométricas entre eles não são suficientes para delimitá-los como espécies distintas.(AU)

Genetic variability of Five Periplaneta americana L. (Dyctioptera: Blattidae) populations in southwestern Colombia using the AFLP molecular marker technique

The genetic structure of five Periplaneta americana (L.) populations from three cities (Cali, Popay¨¢n and Buenaventura) located in southwestern Colombia was estimated using the AFLP molecular marker technique. A set of 174 loci were analyzed, 120 of which were polymorphic. Genetic diversity was estimated at different geographic levels for the total population and between cities. Genetic diversity averaged 0.32. The largest gene flow was detected between two population from Cali (F ST = 0.088; Nm = 2.6) and the lowest from the third population in this city (Univalle) and Popay¨¢n and Buenaventura (F ST = 0.13; Nm = 1.6 and F ST = 0.12; Nm = 1.8, respectively). The individuals' low mobility, the population' geographic separation, and possibly genetic drift have resulted in the population of this cockroach species having a significant degree of structuring, both between cities (¦µst = 0.13; P < 0.001) and within them (¦µst = 0.87; P < 0.001). These results suggest a high degree of subdivision within the P. americana population studied.

A preliminary analysis of the population genetics and molecular phylogenetics of Onchocerca volvulus (Nematoda: Filarioidea) using nuclear ribosomal second internal transcribed spacer sequences

Nuclear internal transcribed spacer 2 (ITS2) rDNA sequences were used for a molecular phylogenetics analysis of five Onchocerca species. The sister species of the human parasite O. volvulus was found to be the cattle parasite O. ochengi and not O. gibsoni, contrary to chromosomal evidence. The genetic differentiation of two African populations (representing the two African strains) and a Brazilian population of O. volvulus was also studied. Phylogenetic and network reconstruction did not show any clustering of ITS2 alleles on geographic or strain grounds. Furthermore, population genetics tests showed no indication of population differentiation but suggested gene flow among the three populations.

Use of AFLPS to distinguish landraces of pejibaye (Bactris gasipaes) in brazilian Amazonia

Although the first inhabitants of western Amazonia domesticated pejibaye (Bactris gasipaes Kunth, Palmae) or peach palm for its fruits, today it is widely planted for its heart-of-palm. Like other domesticates, pejibaye presents a complex hierarchy of landraces developed before the conquest of the Americas. The existence of three landraces (Pará, Solimões, Putumayo) was proposed along the Amazonas and Solimões Rivers, Brazil, based on morphological characteristics. There are some questions remaining about the intermediate landrace being an artifact of the morphometric analysis. AFLPs were used to evaluate the relationships among samples of these putative landraces. DNA was extracted from 99 plants representing 13 populations maintained in the Pejibaye Germplasm Bank, Manaus, AM; six primer combinations generated 245 markers via PCR, which were scored in an ABI Prism 310 sequencer and analyzed with GeneScan Software; Jaccard similarities were estimated and a dendrogram was generated with UPGMA. Two groups of plants were observed in the dendrogram instead of three, and were similar at 0.795. Each group contained two subgroups, similar at 0.815. One group (n=41) contained 73% Pará landrace plants, with one subgroup (n=22) containing 91% Pará, and the other (n=19) containing 53% Pará. The other group (n=58) contained 53% Solimões and 40% Putumayo landrace plants, with one subgroup (n=21) containing 52% Solimões and 43% Putumayo, and the other (n=35) containing 57% Solimões and 37% Putumayo. The first group confirmed the Pará landrace. The second group suggested that the Solimões landrace does not exist, but that the Putumayo landrace extends along the Solimões River to Central Amazonia.

Os primeiros povos da Amazônia ocidental domesticaram a pupunha (Bactris gasipaes Kunth, Palmae) por seu fruto, embora hoje seja muito plantada por seu palmito. Como outros cultivos domesticados, a pupunha apresenta uma hierarquia complexa de raças primitivas criadas antes da conquista das Américas. A existência de três raças (Pará, Solimões, Putumayo) foi proposta ao longo dos rios Amazonas e Solimões, Brasil, com base em características morfológicas. Algumas dúvidas existem sobre a raça intermediária, pois podia ser um artefato da análise morfométrica. AFLPs foram usados para avaliar as relações entre amostras destas raças hipotéticas. DNA foi extraido de 99 plantas representando 13 populações mantidas no Banco de Germoplasma de Pupunha, Manaus, AM; seis combinações de 'primers' geraram 245 marcadores via PCR, que foram codificados num sequenciador ABI Prism 310 e analisados com o programa GeneScan; similaridades de Jaccard foram estimadas e um dendrograma foi criado com UPGMA. Dois grupos de plantas foram observados no dendrograma, em lugar de três, com similaridade de 0,795. Cada grupo continha dois subgrupos, similares a 0,815. Um grupo (n=41) continha 73% de plantas da raça Pará, com um subgrupo (n=22) contendo 91% Pará e o outro (n=19) contendo 53% Pará. O outro grupo (n=58) continha 53% de plantas da raça Solimões e 40% da Putumayo, com um subgrupo (n=21) contendo 52% Solimões e 43% Putumayo, e o outro (n=35) contendo 57% Solimões e 37% Putumayo. O primeiro grupo confirmou a raça Pará, mas o segundo grupo sugeriu que a raça Solimões não existe; em lugar desta raça, a raça Putumayo se extende ao longo do rio Solimões até a Amazônia Central.

Use of AFLPS to distinguish landraces of pejibaye (Bactris gasipaes) in brazilian Amazonia

Although the first inhabitants of western Amazonia domesticated pejibaye (Bactris gasipaes Kunth, Palmae) or peach palm for its fruits, today it is widely planted for its heart-of-palm. Like other domesticates, pejibaye presents a complex hierarchy of landraces developed before the conquest of the Americas. The existence of three landraces (Pará, Solimões, Putumayo) was proposed along the Amazonas and Solimões Rivers, Brazil, based on morphological characteristics. There are some questions remaining about the intermediate landrace being an artifact of the morphometric analysis. AFLPs were used to evaluate the relationships among samples of these putative landraces. DNA was extracted from 99 plants representing 13 populations maintained in the Pejibaye Germplasm Bank, Manaus, AM; six primer combinations generated 245 markers via PCR, which were scored in an ABI Prism 310 sequencer and analyzed with GeneScan Software; Jaccard similarities were estimated and a dendrogram was generated with UPGMA. Two groups of plants were observed in the dendrogram instead of three, and were similar at 0.795. Each group contained two subgroups, similar at 0.815. One group (n=41) contained 73% Pará landrace plants, with one subgroup (n=22) containing 91% Pará, and the other (n=19) containing 53% Pará. The other group (n=58) contained 53% Solimões and 40% Putumayo landrace plants, with one subgroup (n=21) containing 52% Solimões and 43% Putumayo, and the other (n=35) containing 57% Solimões and 37% Putumayo. The first group confirmed the Pará landrace. The second group suggested that the Solimões landrace does not exist, but that the Putumayo landrace extends along the Solimões River to Central Amazonia.

Os primeiros povos da Amazônia ocidental domesticaram a pupunha (Bactris gasipaes Kunth, Palmae) por seu fruto, embora hoje seja muito plantada por seu palmito. Como outros cultivos domesticados, a pupunha apresenta uma hierarquia complexa de raças primitivas criadas antes da conquista das Américas. A existência de três raças (Pará, Solimões, Putumayo) foi proposta ao longo dos rios Amazonas e Solimões, Brasil, com base em características morfológicas. Algumas dúvidas existem sobre a raça intermediária, pois podia ser um artefato da análise morfométrica. AFLPs foram usados para avaliar as relações entre amostras destas raças hipotéticas. DNA foi extraido de 99 plantas representando 13 populações mantidas no Banco de Germoplasma de Pupunha, Manaus, AM; seis combinações de 'primers' geraram 245 marcadores via PCR, que foram codificados num sequenciador ABI Prism 310 e analisados com o programa GeneScan; similaridades de Jaccard foram estimadas e um dendrograma foi criado com UPGMA. Dois grupos de plantas foram observados no dendrograma, em lugar de três, com similaridade de 0,795. Cada grupo continha dois subgrupos, similares a 0,815. Um grupo (n=41) continha 73% de plantas da raça Pará, com um subgrupo (n=22) contendo 91% Pará e o outro (n=19) contendo 53% Pará. O outro grupo (n=58) continha 53% de plantas da raça Solimões e 40% da Putumayo, com um subgrupo (n=21) contendo 52% Solimões e 43% Putumayo, e o outro (n=35) contendo 57% Solimões e 37% Putumayo. O primeiro grupo confirmou a raça Pará, mas o segundo grupo sugeriu que a raça Solimões não existe; em lugar desta raça, a raça Putumayo se extende ao longo do rio Solimões até a Amazônia Central.