Deciphering patterns of transoceanic dispersal: the evolutionary origin and biogeography of coastal lizards (Cryptoblepharus) in the Western Indian Ocean region

Aim Cryptoblepharus is a genus of small arboreal or rock‐dwelling scincid lizards, widespread through the Indo‐Pacific and Australian regions, with a disjunct outlier in the Malagasy region. The taxonomy within this genus is controversial, with different authors ranking the different forms (now some 36) at various levels, from different species to subspecies of a single species, Cryptoblepharus boutonii. We investigated the biogeography and genetic differentiation of the Cryptoblepharus from the Western Indian Ocean region, in order to understand their origin and history. Location Western Indian Ocean region. Methods We analysed sequences of mitochondrial DNA (partial 12s and 16s rRNA genes, 766 bp) from 48 specimens collected in Madagascar, Mauritius, the four Comoros islands and East Africa, and also in New Caledonia, representing the Australo‐Pacific unit of the distribution. Results Pairwise sequence divergences of c. 3.1% were found between the New Caledonian forms and the ones from the Western Indian Ocean. Two clades were identified in Madagascar, probably corresponding to the recognized forms cognatus and voeltzkowi, and two clades were identified in the Comoro islands, where each island population formed a distinct haplotype clade. The East African samples form a monophyletic unit, with some variation existing between Pemba, Zanzibar and continental Tanzania populations. Individuals from Mauritius form a divergent group, more related to populations from Moheli and Grand Comore (Comoros islands) than to the others. Main conclusions The level of divergence between the populations from the Western Indian Ocean and Australian regions and the geographic coherence of the variation within the Western Indian Ocean group are concordant with the hypothesis of a colonization of this region by a natural transoceanic dispersal (from Australia or Indonesia). The group then may have diversified in Madagascar, from where it separately colonized the East African coast, the Comoros islands (twice), and Mauritius. The genetic divergence found is congruent with the known morphological variation, but its degree is much lower than typically seen between distinct species of reptiles.     

Tracking island colonization history and phenotypic shifts in Indian Ocean bulbuls (Hypsipetes: Pycnonotidae)

Molecular phylogenies of island organisms provide useful systems for testing hypotheses of convergent or parallel evolution, since selectively neutral molecular characters are likely to be independent of phenotype, and the existence of similar environments on multiple isolated islands provides numerous opportunities for populations to evolve independently under the same constraints. Here we construct a phylogenetic hypothesis for Hypsipetes bulbuls of the western Indian Ocean, and use this to test hypotheses of colonization pattern and phenotypic change among islands of the region. Mitochondrial sequence data were collected from all extant taxa of the region, combined with sequence data from relevant lineages in Asia. Data are consistent with a single Hypsipetes colonization of the western Indian Ocean from Asia within the last 2.6 Myr. The expansion of Hypsipetes appears to have occurred rapidly, with descendants found across the breadth of its western Indian Ocean range. The data suggest that a more recent expansion of Hypsipetes madagascariensis from Madagascar led to the colonization of Aldabra and a secondary colonization of the Comoros. Groupings of western Indian Ocean Hypsipetes according to phenotypic similarities do not correspond to mtDNA lineages, suggesting that these similarities have evolved by convergence or parallelism. The direction of phenotypic change cannot be inferred with confidence, since the primary expansion occurred rapidly relative to the rate of mtDNA substitution, and the colonization sequence remains uncertain. However, evidence from biogeography and comparison of independent colonization events are consistent with the persistence of a small grey continental bulbul in India and Madagascar, and multiple independent origins of large size and green plumage in insular island populations of the Comoros, Mascarenes and Seychelles. © 2005 The Linnean Society of London, Biological Journal of the Linnean Society, 2005, 85 , 271–287.     

Phylogeny of Gaertnera Lam. (Rubiaceae) based on multiple DNA markers: evidence of a rapid radiation in a widespread, morphologically diverse genus

Phylogenetic relationships among 28 of the 68 species of the paleotropical genus Gaertnera (Rubiaceae) and two related genera were inferred from nucleotide sequence variation in four nuclear DNA (nDNA) markers: the internal transcribed spacers of nuclear rDNA (ITS), the large and small copies of phosphoenolpyruvate carboxylase (PepC-large and PepC-small), and triose phosphate isomerase (Tpi). Phylogenetic analysis of the combined nDNA dataset suggested that Gaertnera is monophyletic, but genetic variation among species was insufficient to reconstruct well-supported relationships within the genus. This was counter to expectations based on the very distinct morphologies and widespread distribution of the genus (West Africa to Sulawesi). Molecular clock analyses suggested variable dates of origin for Gaertnera depending upon the calibration method used. The most plausible calibration implies that Gaertnera migrated to Africa during the early Tertiary, possibly via a boreotropical land bridge and suggests that Gaertnera started to radiate 5.21 +/- 0.14 million years ago. This implies that range expansion in the group has occurred via a number of long-distance dispersal events rather than vicariance. The molecular clock estimate in turn estimated an unusually rapid lineage diversification rate within the radiation of 0.717-0.832 species/million years, comparable to those estimated for radiations on oceanic islands. Although low interspecific competition levels may have contributed to the diversification of Gaertnera on Mauritius, the mechanisms driving the rapid radiation of the group in other parts of its range remain elusive.     

Short-term progressive early diagenesis in density bands of recent corals:Porites Colonies,Mauritius Island,Indian Ocean

Summary The growth history of some recentPorites colonies of Mauritius Island (Indian Ocean) was dated by sclerochronological methods. Couples of high-density and low-density bands represent the annual growth rate of the corals and allow the growth pattern of every year in the corallum to be counted. The growth and structure of the skeletons ofPorites solida andPorites lutea were investigated. Older parts of the aragonitic skeleton in these 10 to 20 year old corals show various secondary microstructures resulting from alterations and thickenings of the elements of the skeleton. The primary needle-like aragonite crystals are absent in older parts of the corallum and blocky aragonitic cements can occur. Pores and primary skeletal elements are overgrown by new microstructures. These microstructures are caused by secondary cementation and exhibit frontal zones (Stirnzonen), zigzag-like and pseudolamellar-structures. The lamellar structures can be compared with similar structures in the exoskeleton of some Rugosa. A very short early diagenesis within the recent corals is responsible for the thickening and alteration of skeletal elements. It occurs only 4 to 5 years after the formation of the skeleton and tends to increase in importance in older parts of the corallum. Nevertheless, there is no proof for any alteration of aragonite to calcite.     

How marine currents influenced the widespread natural overseas dispersal of reptiles in the Western Indian Ocean region

In a recent contribution to this journal, Wilmé et al. (2016) proposed that the giant tortoises of the islands of the Western Indian Ocean (WIO: Aldabra, the Mascarenes, and the Granitic Seychelles) might have originated from translocation by early Austronesian sailors. Prompted by this paper we review recent literature and show that natural overseas dispersal was remarkably widespread in the colonization history of terrestrial reptiles in the WIO region. Almost 90% of the successful colonization events are supported by prevailing marine surface currents. However, these currents may change over geological (and evolutionary) time‐scales, and eddies and counter‐currents may facilitate transport against the main current direction. We review the cases of the extant and extinct WIO giant tortoises and suggest that the current distribution of all lineages can be convincingly explained by overseas dispersal.     

A time‐calibrated phylogenetic approach to assessing the phylogeography,colonization history and phenotypic evolution of snakes in the Japanese Izu Islands

Aim We infer the biogeography and colonization history of a dispersal‐limited terrestrial vertebrate, the Japanese four‐lined ratsnake (Elaphe quadrivirgata), to reveal the number of times mainland populations have invaded the Izu Archipelago of Japan, the mainland sources of these colonists, and the time‐scale of colonization. We compare these results with those of past studies in an attempt to uncover general biogeographical patterns. Moreover, we briefly examine the significance of colonization history when evaluating the evolution of body size and melanism of the Izu Island E. quadrivirgata populations. Location The Izu Islands (Oshima, Toshima, Niijima, Shikine, Kozu, Tadanae and Mikura), a volcanic archipelago off the Pacific coast of central Japan. Methods We obtained DNA sequences of the mitochondrial cytochrome b gene (1117 base pairs) from 373 individual snakes sampled from seven of the Izu Islands and 25 mainland localities. We employed partitioned Bayesian phylogenetic analyses assuming a relaxed molecular clock to estimate phylogenetic relationships among extant haplotypes and to give an explicit temporal scale to the timing of clade divergence, colonization history and tempo of body‐size evolution. Moreover, we employed model‐based biogeographical analysis to calculate the minimum number of times E. quadrivirgata colonized the Izu Islands. Results We found evidence that three separate regions of the Izu Archipelago have been colonized independently from mainland ancestors within the past 0.58–0.20 Ma. The Izu Peninsula plus Oshima and Mikura were both colonized independently from lineages inhabiting eastern mainland Japan. The Toshima, Niijima, Shikine, Kozu and Tadanae populations all derive from a single colonization from western mainland Japan. Oshima has been subject to three or possibly four colonizations. Main conclusions These results support the hypothesis that the extreme body‐size disparity among island populations of this ratsnake evolved in situ. Moreover, the fact that the dwarf, melanistic population inhabiting Oshima descends from multiple mainland colonization events is evidence of an extremely strong natural selection pressure resulting in the rapid evolution of this unique morphology. These results contrast with theoretical predictions that natural selection pressures should play a decreased role on islands close to the mainland and/or subject to frequent or recent immigration.     

Immigration, species radiation and extinction in a highly diverse songbird lineage: white-eyes on Indian Ocean islands

Molecular phylogenetic hypotheses of species-rich lineages in regions where geological history can be reliably inferred may provide insights into the scale of processes driving diversification. Here we sample all extant or recently extinct white-eye (Zosterops) taxa of the southwest Indian Ocean, combined with samples from all principal continental lineages. Results support a high dispersal capability, with at least two independent continental sources for white-eyes of the region. An early (within 1.8 million years ago) expansion into the Indian Ocean may have originated either from Asia or Africa; the three resulting lineages show a disparate distribution consistent with considerable extinction following their arrival. Africa is supported as the origin of a later expansion into the region (within 1.2 million years ago). On two islands, a pair of Zosterops species derived from independent immigrations into the Indian Ocean co-occur or may have formerly co-occurred, providing strong support for their origin by double-island colonization rather than within-island (sympatric or microallopatric) speciation. On Mauritius and La Réunion, phylogenetic placement of sympatric white-eyes allow us to rule out a scenario in which independent within-island speciation occurred on both islands; one of the species pairs must have arisen by double colonization, while the other pair is likely to have arisen by the same mechanism. Long-distance immigration therefore appears to be responsible for much of the region's white-eye diversity. Independent immigrations into the region have resulted in lineages with mutually exclusive distributions and it seems likely that competition with congeneric species, rather than arrival frequency, may limit present-day diversity.     

Revisiting the phylogeography of Asellus aquaticus in Europe: insights into cryptic diversity and spatiotemporal diversification

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The influence of protected area management on the status of coral reefs at Misali Island,Tanzania following the 1998 bleaching event in the western Indian Ocean

The reefs of Misali Island, Tanzania suffered serious coral mortality as a result of the 1998 coral bleaching event in the western Indian Ocean. Four years after this event, Frontier‐Tanzania assessed reef substrate in Misali Island Marine Conservation Area (MIMCA) using scuba surveys. Substrate cover was compared between sites within and outside the nonextraction zone (NEZ) of MIMCA and recovery rates were estimated through reference to previous studies. Coral recovery at Misali Island following the bleaching event has generally been slow, but nonextractive management of MIMCA has enhanced coral recovery within the NEZ. The absence of fishing activities in the NEZ has sustained herbivorous fish populations and thus reduced algal overgrowth; however, outside the NEZ, diving and anchoring activities have caused further coral damage and led to increased levels of debris. Awareness‐raising programmes could lessen these problems and the NEZ appeared to be well‐placed to capitalize on nonmanageable factors that are thought to mitigate the effects of coral bleaching. In addition, the implementation of flexible zoning schemes could improve recovery rates after future bleaching events.     

Biogeography and phylogeny of the New Zealand cicada genera (Hemiptera: Cicadidae) based on nuclear and mitochondrial DNA data

Aim Determine the geographical and temporal origins of New Zealand cicadas. Location New Zealand, eastern Australia and New Caledonia. Methods DNA sequences from 14 species of cicadas from New Zealand, Australia, and New Caledonia were examined. A total of 4628 bp were analysed from whole genome extraction of four mitochondrial genes (cytochrome oxidase subunits I and II, and ribosomal 12S and 16S subunits) and one nuclear gene (elongation factor‐1 alpha). These DNA sequences were aligned and analysed using standard phylogenetic methods based primarily on the maximum likelihood optimality criterion. Dates of divergences between clades were determined using several molecular clock methods. Results New Zealand cicadas form two well‐defined clades. One clade groups with Australian taxa, the other with New Caledonian taxa. The molecular clock analyses indicate that New Zealand genera diverged from the Australian and New Caledonian genera within the last 11.6 Myr. Main conclusions New Zealand was likely colonized by two or more invasions. One NZ lineage has its closest relatives in Australia and the other in New Caledonia. These invasions occurred well after New Zealand became isolated from other land masses, therefore cicadas must have crossed large bodies of water to reach New Zealand.     

Evolution of subterranean diving beetles (Coleoptera: Dytiscidae: Hydroporini, Bidessini) in the arid zone of Australia

Calcrete aquifers in arid inland Australia have recently been found to contain the world's most diverse assemblage of subterranean diving beetles (Coleoptera: Dytiscidae). In this study we test whether the adaptive shift hypothesis (ASH) or the climatic relict hypothesis (CRH) is the most likely mode of evolution for the Australian subterranean diving beetles by using a phylogeny based on two sequenced fragments of mitochondrial genes (CO1 and 16S-tRNA-ND1) and linearized using a relaxed molecular clock method. Most individual calcrete aquifers contain an assemblage of diving beetle species of distantly related lineages and/or a single pair of sister species that significantly differ in size and morphology. Evolutionary transitions from surface to subterranean life took place in a relatively small time frame between nine and four million years ago. Most of the variation in divergence times of the sympatric sister species is explained by the variation in latitude of the localities, which correlates with the onset of aridity from the north to the south and with an aridity maximum in the Early Pliocene (five mya). We conclude that individual calcrete aquifers were colonized by several distantly related diving beetle lineages. Several lines of evidence from molecular clock analyses support the CRH, indicating that all evolutionary transitions took place during the Late Miocene and Early Pliocene as a result of aridification.     

Red crabs in rainforest on Christmas Island,Indian Ocean: No effect on the soil seed bank

Red land crabs (Gecarcoidea natalis) are both abundant and widespread in intact rainforest on Christmas Island, Indian Ocean, occurring at densities of ~ 1 crab m^–2. We used an on-going exclusion experiment to test the hypothesis that through their activities as seed predators and soil disturbers, these omnivores are important determinants of the density and relative species abundance of seeds in the soil seed bank of undisturbed primary rainforest. After 6.5 y of continuous crab exclusion, there was no significant difference in the density of seeds in the top 3 cm of soil between the control (3671 ± 517 seeds/0.19 m²) and exclusion plots (3285 ± 801 seeds/0.19 m²), nor was there any significant treatment difference in relative species composition. There were also no significant treatment differences when Muntingia calabura, which accounted for 91.9% of all seeds in the seed bank, was excluded from the analyses. We attribute the lack of a significant impact of red crabs to their probable inability to handle the very small seeds which occurred in the seed bank, and the low level of soil disturbance at the study site. We conclude that the observed impact of red crabs on the abundance and diversity of successional species in natural light gaps on Christmas Island is due to their impact on germinating seedlings alone, rather than any additional impact on the soil seed bank.     

Comparative phylogeography of livebearing fishes in the genera Poeciliopsis and Poecilia (Poeciliidae: Cyprinodontiformes) in central Mexico

Aim To test the hypothesis that the vicariant event responsible for north–south divergences in two clades of the fish genus Poeciliopsis Regan was also responsible for north–south divergences in the fish Poecilia butleri Jordan. Location Central Mexico. Methods Parsimony, distance, maximum likelihood and Bayesian phylogenetic analyses of a mitochondrial gene. Molecular clock test and Bayesian analyses of divergence time. Results We report concordant phylogeographical patterns between two clades in the genus Poeciliopsis (i.e. the other formed by P. latidens Garman and P. fasciata Meek, and the other formed by P. presidionis Jordan and P. turneri Miller) and the clade of Poecilia butleri, with northern and southern individuals within each clade grouping into separate lineages. There is also evidence for slower substitution rates in Poecilia compared with Poeciliopsis. After taking into account these substitution rate discrepancies with Bayesian relaxed molecular clock analyses, north–south divergences in Poecilia butleri were equivalent to those reported for Poeciliopsis latidens‐fasciata and P. presidionis‐turneri. Main conclusions The same Plio‐Pleistocene vicariant event associated with geological activity of the Trans‐Mexican Volcanic Belt (TMVB) appears to have caused divergence in three different freshwater fish lineages. This study is an example of how comparative phylogeography can strengthen inferences about vicariant events in regions of high biological diversity and complex geological history such as the TMVB.     

Systematics and phylogeography of the Taiwanese endemic minnow Candidia barbatus (Pisces: Cyprinidae) based on DNA sequence,allozymic, and morphological analyses

The phylogeographical patterns of the endemic minnow, Candidia barbatus, were examined using mitochondrial (mt) cytochrome b sequences, allozyme electrophoresis, and morphology. The 23 mtDNA haplotypes recovered were grouped into six lineages (A–F). According to a molecular clock of 1.7% per million years, we estimated that the colonization of C. barbatus to Taiwan took place three million years ago with several subsequent vicariance events. In the allozyme analysis, the Bayesian approach partitioned different populations into three clusters, with clusters I, II, and III mainly found in northern, central, and southern Taiwan, respectively. Morphologically, fish from north and south of the Tropic of Cancer had distinct numbers of lateral line scales and were separated into two morphs. The cohabitation of different mtDNA lineages with high allozymic and morphological similarities suggests that interbreeding amongst them is possible. Therefore, in spite of highly differentiated mtDNA, samples from different collecting sites should be considered a single species. Based on the current population distribution and levels of divergence, we propose that both orogenic activities and marine regression during glacial epoch were the two main forces shaping population differentiation of this species. © 2011 The Linnean Society of London, Zoological Journal of the Linnean Society, 2011, 161 , 613–632.     

Comparative phylogeography of endemic cyprinids in the south-west Iberian Peninsula: evidence for a new ichthyogeographic area

The comparative phylogeography and evolutionary history of three native cyprinid genera, Squalius (subfamily Leuciscinae), Chondrostoma (subfamily Leuciscinae) and Barbus (subgenus Luciobarbus ; subfamily Cyprininae), were examined focusing mainly in the South-Western region of the Iberian Peninsula, where recently described endemic species are present with considerably restricted distribution areas. In order to accomplish that the variation at the complete cytochrome b gene (1140 bp) was analysed for specimens from the South-Western region, and also for representatives of the three genera from all over the Iberian Peninsula. Data indicate different evolutionary histories, with distinct time periods of colonization between the two cyprinid subfamilies in the Iberian Peninsula. Four new Iberian ichthyogeographic areas are accordingly proposed based on congruent phylogeographic and geological evidences: the South-Western, the South-Eastern, the Atlantic and the Mediterranean. Evidence was provided for the older isolation of the South-Western area in the Miocene during the Endorheic Drainages phase, designating a clearly defined and distinct ichthyogeographic area. A new molecular clock calibration is proposed for the subgenus Luciobarbus .     

A revision of the ant genus Mystrium in the Malagasy region with description of six new species and remarks on Amblyopone and Stigmatomma (Hymenoptera,Formicidae, Amblyoponinae)

The genus Mystrium is revised for the Malagasy region. Six species, Mystrium barrybresslerisp. n., Mystrium labyrinthsp. n., Mystrium equessp. n., Mystrium mirrorsp. n., Mystrium shadowsp. n., and Mystrium janovitzisp. n. are described as new. Two existing names, Mystrium fallax Forel and Mystrium stadelmanni Forel, are synonymized with Mystrium voeltzkowi Forel and Mystrium mysticum Roger, respectively. All recognized species, including species outside of the Malagasy region, are assigned to one of the three newly proposed species groups. The associations between existing names and males are reexamined, and males of eight of the ten Malagasy species are described or redescribed. The taxonomic history of Mystrium highlights the importance of using unique identifiers when designating type specimens and the use of deposited vouchers in phylogenetic and ecological studies. Keys to species for workers, queens, and males are provided. Furthermore, a neotype for Mystrium mysticum is designated, as well as lectotypes for Mystrium camillae Emery, Mystrium rogeri Forel, Mystrium fallax Forel, Mystrium oberthueri Forel, Mystrium stadelmanni Forel, and Mystrium voeltzkowi Forel. Stigmatomma gingivale (Brown) is reassigned to Amblyopone as comb. rev. and Amblyopone awa Xu & Chu, Amblyopone kangba Xu & Chu, Amblyopone meiliana Xu & Chu, and Amblyopone zomae Xu & Chu are transferred to the genus Stigmatomma as comb. n.     

Phylogeography of the introduced species Rattus rattus in the western Indian Ocean,with special emphasis on the colonization history of Madagascar

Aim To describe the phylogeographic patterns of the black rat, Rattus rattus, from islands in the western Indian Ocean where the species has been introduced (Madagascar and the neighbouring islands of Réunion, Mayotte and Grande Comore), in comparison with the postulated source area (India). Location Western Indian Ocean: India, Arabian Peninsula, East Africa and the islands of Madagascar, Réunion, Grande Comore and Mayotte. Methods Mitochondrial DNA (cytochrome b, tRNA and D‐loop, 1762 bp) was sequenced for 71 individuals from 11 countries in the western Indian Ocean. A partial D‐loop (419 bp) was also sequenced for eight populations from Madagascar (97 individuals), which were analysed in addition to six previously published populations from southern Madagascar. Results Haplotypes from India and the Arabian Peninsula occupied a basal position in the phylogenetic tree, whereas those from islands were distributed in different monophyletic clusters: Madagascar grouped with Mayotte, while Réunion and Grand Comore were present in two other separate groups. The only exception was one individual from Madagascar (out of 190) carrying a haplotype that clustered with those from Réunion and South Africa. ‘Isolation with migration’ simulations favoured a model with no recurrent migration between Oman and Madagascar. Mismatch distribution analyses dated the expansion of Malagasy populations on a time‐scale compatible with human colonization history. Higher haplotype diversity and older expansion times were found on the east coast of Madagascar compared with the central highlands. Main conclusions Phylogeographic patterns supported the hypothesis of human‐mediated colonization of R. rattus from source populations in either the native area (India) or anciently colonized regions (the Arabian Peninsula) to islands of the western Indian Ocean. Despite their proximity, each island has a distinct colonization history. Independent colonization events may have occurred simultaneously in Madagascar and Grande Comore, whereas Mayotte would have been colonized from Madagascar. Réunion was colonized independently, presumably from Europe. Malagasy populations may have originated from a single successful colonization event, followed by rapid expansion, first in coastal zones and then in the central highlands. The congruence of the observed phylogeographic pattern with human colonization events and pathways supports the potential relevance of the black rat in tracing human history.     

Seri Indian traditional knowledge and molecular biology agree: no express train for island‐hopping spiny‐tailed iguanas in the Sea of Cortés

Aim The role of human activities in species biogeography can be difficult to identify, but in some cases molecular techniques can be used to test hypotheses of human‐mediated dispersal. A currently accepted hypothesis states that humans mediated the divergence of two species of spiny‐tailed iguanas in the Ctenosaura hemilopha species complex, namely C. conspicuosa and C. nolascensis, which occupy islands in the Sea of Cortés between the peninsula of Baja California and mainland Mexico. We test an alternative hypothesis that follows the traditional knowledge of the Seri Indians and states that the divergence of these species was not mediated by humans. Location Mexico, including Baja California, Sonoran and Sinaloan coastal regions, and Isla San Esteban and Isla San Pedro Nolasco in the Sea of Cortés. Methods We analysed mitochondrial (cytochrome b and cytochrome c oxidase subunit III) DNA sequences from four species in the C. hemilopha species complex. Maximum parsimony and Bayesian inference were used to infer matriarchal genealogical relationships between the species and several outgroup taxa. Bayesian methods were used to estimate divergence times for the major nodes on the trees based on previously published, fossil‐calibrated priors. Results Our analysis indicated that lineages within the C. hemilopha species complex diverged long before human colonization of the Americas. The divergence of C. nolascensis and C. conspicuosa could not be attributed to Seri translocations. The matriarchal genealogy of the species complex currently defies a simple biogeographical interpretation. Main conclusions We conclude that humans did not mediate the divergence of C. nolascensis and C. conspicuosa. This conclusion is consistent with the traditional knowledge of the Seri people. These results demonstrate the utility of molecular techniques in investigating potential cases of human‐mediated dispersal of plants and animals, and reinforce the importance of considering traditional knowledge in the formation of scientific hypotheses and the interpretation of results.