Revising the Bantu tree

Phylogenetic methods offer a promising advance for the historical study of language and cultural relationships. Applications to date, however, have been hampered by traditional approaches dependent on unfalsifiable authority statements: in this regard, historical linguistics remains in a similar position to evolutionary biology prior to the cladistic revolution. Influential phylogenetic studies of Bantu languages over the last two decades, which provide the foundation for multiple analyses of Bantu sociocultural histories, are a major case in point. Comparative analyses of basic lexica, instead of directly treating written words, use only numerical symbols that express non-replicable authority opinion about underlying relationships. Building on a previous study of Uto-Aztecan, here we analyse Bantu language relationships with methods deriving from DNA sequence optimization algorithms, treating basic vocabulary as sequences of sounds. This yields finer-grained results that indicate major revisions to the Bantu tree, and enables more robust inferences about the history of Bantu language expansion and/or migration throughout sub-Saharan Africa. “Early-split” versus “late-split” hypotheses for East and West Bantu are tested, and overall results are compared to trees based on numerical reductions of vocabulary data. Reconstruction of language histories is more empirically based and robust than with previous methods.     

Networks uncover hidden lexical borrowing in Indo-European language evolution

Language evolution is traditionally described in terms of family trees with ancestral languages splitting into descendent languages. However, it has long been recognized that language evolution also entails horizontal components, most commonly through lexical borrowing. For example, the English language was heavily influenced by Old Norse and Old French; eight per cent of its basic vocabulary is borrowed. Borrowing is a distinctly non-tree-like process--akin to horizontal gene transfer in genome evolution--that cannot be recovered by phylogenetic trees. Here, we infer the frequency of hidden borrowing among 2346 cognates (etymologically related words) of basic vocabulary distributed across 84 Indo-European languages. The dataset includes 124 (5%) known borrowings. Applying the uniformitarian principle to inventory dynamics in past and present basic vocabularies, we find that 1373 (61%) of the cognates have been affected by borrowing during their history. Our approach correctly identified 117 (94%) known borrowings. Reconstructed phylogenetic networks that capture both vertical and horizontal components of evolutionary history reveal that, on average, eight per cent of the words of basic vocabulary in each Indo-European language were involved in borrowing during evolution. Basic vocabulary is often assumed to be relatively resistant to borrowing. Our results indicate that the impact of borrowing is far more widespread than previously thought.     

Language trees ≠ gene trees

Darwin saw similarities between the evolution of species and the evolution of languages, and it is now widely accepted that similarities between related languages can often be interpreted in terms of a bifurcating descent history (‘phylogenesis’). Such interpretations are supported when the distributions of shared and unshared traits (for example, in terms of etymological roots for elements of basic vocabulary) are analysed using tree-building techniques and found to be well-explained by a phylogenetic model. In this article, we question the demographic assumption which is sometimes made when a tree-building approach has been taken to a set of cultures or languages, namely that the resulting tree is also representative of a bifurcating population history. Using historical census data relating to Gaelic- and English-speaking inhabitants of Sutherland (Highland Scotland), we have explored the dynamics of language death due to language shift, representing the extreme case of lack of congruence between the genetic and the culture–historical processes. Such cases highlight the important role of selective cultural migration (or shifting between branches) in determining the extinction rates of different languages on such trees.     

A phylogenetic approach to cultural evolution

There has been a rapid increase in the use of phylogenetic methods to study the evolution of languages and culture. Languages fit a tree model of evolution well, at least in their basic vocabulary, challenging the view that blending, or admixture among neighbouring groups, was predominant in cultural history. Here, we argue that we can use language trees to test hypotheses about not only cultural history and diversification, but also bio-cultural adaptation. Phylogenetic comparative methods take account of the non-independence of cultures (Galton's problem), which can cause spurious statistical associations in comparative analyses. Advances in phylogenetic methods offer new possibilities for the analysis of cultural evolution, including estimating the rate of evolution and the direction of coevolutionary change of traits on the tree. They also enable phylogenetic uncertainty to be incorporated into the analyses, so that one does not have to treat phylogenetic trees as if they were known without error.     

The shape and tempo of language evolution

There are approximately 7000 languages spoken in the world today. This diversity reflects the legacy of thousands of years of cultural evolution. How far back we can trace this history depends largely on the rate at which the different components of language evolve. Rates of lexical evolution are widely thought to impose an upper limit of 6000–10 000 years on reliably identifying language relationships. In contrast, it has been argued that certain structural elements of language are much more stable. Just as biologists use highly conserved genes to uncover the deepest branches in the tree of life, highly stable linguistic features hold the promise of identifying deep relationships between the world''s languages. Here, we present the first global network of languages based on this typological information. We evaluate the relative evolutionary rates of both typological and lexical features in the Austronesian and Indo-European language families. The first indications are that typological features evolve at similar rates to basic vocabulary but their evolution is substantially less tree-like. Our results suggest that, while rates of vocabulary change are correlated between the two language families, the rates of evolution of typological features and structural subtypes show no consistent relationship across families.     

Oceanic barriers promote language diversification in the Japanese Islands

Good barriers make good languages. Scholars have long speculated that geographical barriers impede linguistic contact between speech communities and promote language diversification in a manner similar to the process of allopatric speciation. This hypothesis, however, has seldom been tested systematically and quantitatively. Here, we adopt methods from evolutionary biology and attempt to quantify the influence of oceanic barriers on the degree of lexical diversity in the Japanese Islands. Measuring the degree of beta diversity from basic vocabularies, we find that geographical proximity and, more importantly, isolation by surrounding ocean, independently explains a significant proportion of lexical variation across Japonic languages. Further analyses indicate that our results are neither a by‐product of using a distance matrix derived from a Bayesian language phylogeny nor an epiphenomenon of accelerated evolutionary rates in languages spoken by small communities. Moreover, we find that the effect of oceanic barriers is reproducible with the Ainu languages, indicating that our analytic approach as well as the results can be generalized beyond Japonic language family. The findings we report here are the first quantitative evidence that physical barriers formed by ocean can influence language diversification and points to an intriguing common mechanism between linguistic and biological evolution.     

Cultural phylogeography of the Bantu Languages of sub-Saharan Africa

There is disagreement about the routes taken by populations speaking Bantu languages as they expanded to cover much of sub-Saharan Africa. Here, we build phylogenetic trees of Bantu languages and map them onto geographical space in order to assess the likely pathway of expansion and test between dispersal scenarios. The results clearly support a scenario in which groups first moved south through the rainforest from a homeland somewhere near the Nigeria–Cameroon border. Emerging on the south side of the rainforest, one branch moved south and west. Another branch moved towards the Great Lakes, eventually giving rise to the monophyletic clade of East Bantu languages that inhabit East and Southeastern Africa. These phylogenies also reveal information about more general processes involved in the diversification of human populations into distinct ethnolinguistic groups. Our study reveals that Bantu languages show a latitudinal gradient in covering greater areas with increasing distance from the equator. Analyses suggest that this pattern reflects a true ecological relationship rather than merely being an artefact of shared history. The study shows how a phylogeographic approach can address questions relating to the specific histories of certain groups, as well as general cultural evolutionary processes.     

Bringing together linguistic and genetic evidence to test the Bantu expansion

The expansion of Bantu languages represents one of the most momentous events in the history of Africa. While it is well accepted that Bantu languages spread from their homeland (Cameroon/Nigeria) approximately 5000 years ago (ya), there is no consensus about the timing and geographical routes underlying this expansion. Two main models of Bantu expansion have been suggested: The 'early-split' model claims that the most recent ancestor of Eastern languages expanded north of the rainforest towards the Great Lakes region approximately 4000 ya, while the 'late-split' model proposes that Eastern languages diversified from Western languages south of the rainforest approximately 2000 ya. Furthermore, it is unclear whether the language dispersal was coupled with the movement of people, raising the question of language shift versus demic diffusion. We use a novel approach taking into account both the spatial and temporal predictions of the two models and formally test these predictions with linguistic and genetic data. Our results show evidence for a demic diffusion in the genetic data, which is confirmed by the correlations between genetic and linguistic distances. While there is little support for the early-split model, the late-split model shows a relatively good fit to the data. Our analyses demonstrate that subsequent contact among languages/populations strongly affected the signal of the initial migration via isolation by distance.     

Does Lateral Transmission Obscure Inheritance in Hunter-Gatherer Languages?

In recent years, linguists have begun to increasingly rely on quantitative phylogenetic approaches to examine language evolution. Some linguists have questioned the suitability of phylogenetic approaches on the grounds that linguistic evolution is largely reticulate due to extensive lateral transmission, or borrowing, among languages. The problem may be particularly pronounced in hunter-gatherer languages, where the conventional wisdom among many linguists is that lexical borrowing rates are so high that tree building approaches cannot provide meaningful insights into evolutionary processes. However, this claim has never been systematically evaluated, in large part because suitable data were unavailable. In addition, little is known about the subsistence, demographic, ecological, and social factors that might mediate variation in rates of borrowing among languages. Here, we evaluate these claims with a large sample of hunter-gatherer languages from three regions around the world. In this study, a list of 204 basic vocabulary items was collected for 122 hunter-gatherer and small-scale cultivator languages from three ecologically diverse case study areas: northern Australia, northwest Amazonia, and California and the Great Basin. Words were rigorously coded for etymological (inheritance) status, and loan rates were calculated. Loan rate variability was examined with respect to language area, subsistence mode, and population size, density, and mobility; these results were then compared to the sample of 41 primarily agriculturalist languages. Though loan levels varied both within and among regions, they were generally low in all regions (mean 5.06%, median 2.49%, and SD 7.56), despite substantial demographic, ecological, and social variation. Amazonian levels were uniformly very low, with no language exhibiting more than 4%. Rates were low but more variable in the other two study regions, in part because of several outlier languages where rates of borrowing were especially high. High mobility, prestige asymmetries, and language shift may contribute to the high rates in these outliers. No support was found for claims that hunter-gatherer languages borrow more than agriculturalist languages. These results debunk the myth of high borrowing in hunter-gatherer languages and suggest that the evolution of these languages is governed by the same type of rules as those operating in large-scale agriculturalist speech communities. The results also show that local factors are likely to be more critical than general processes in determining high (or low) loan rates.     

Ethno-cultural heritage of the peoples of west Pamir

The paper presents a study on the West Pamir communities (Badakhshan, Tajikistan) based on language data of the Pamir languages of East-Iranian origin. At present, the ethnic groups of the West Pamir: Shughni, Rushani, Bartangi, Roshorvi, Khufi, Ishkashimi and Wakhi are communities of the Ismaili Shia Islam faith. The underlying concept of the paper is that ethnic and cultural key-concepts take place in the origin of the traditional beliefs and rituals and can be reliably reconstructed from the vocabulary of a particular community. Pamir languages vocabulary of the Pamir ethnic communities that uses East and West Iranian culturally significant terms may serve as a basis for understanding the genesis of Pamir Ismailism, its particular way of absorbing external cultures, and its present specific form. The purely linguistic criteria, like temporal dimension (linguistic chronology), and geographic dispersion and mobility, are believed to be critical for linking etymology to broader socio-cultural development context of the Pamir communities socio-historical and cultural life. Contemporary social and linguistic dynamics in the relations between nation, religion and language in Tajikistan needs systematic research, and reasonable interpretation.     

Bayesian phylogeography of the Arawak expansion in lowland South America

Phylogenetic inference based on language is a vital tool for tracing the dynamics of human population expansions. The timescale of agriculture-based expansions around the world provides an informative amount of linguistic change ideal for reconstructing phylogeographies. Here we investigate the expansion of Arawak, one of the most widely dispersed language families in the Americas, scattered from the Antilles to Argentina. It has been suggested that Northwest Amazonia is the Arawak homeland based on the large number of diverse languages in the region. We generate language trees by coding cognates of basic vocabulary words for 60 Arawak languages and dialects to estimate the phylogenetic relationships among Arawak societies, while simultaneously implementing a relaxed random walk model to infer phylogeographic history. Estimates of the Arawak homeland exclude Northwest Amazonia and are bi-modal, with one potential homeland on the Atlantic seaboard and another more likely origin in Western Amazonia. Bayesian phylogeography better supports a Western Amazonian origin, and consequent dispersal to the Caribbean and across the lowlands. Importantly, the Arawak expansion carried with it not only language but also a number of cultural traits that contrast Arawak societies with other lowland cultures.     

Distinctive Phyllosphere Bacterial Communities in Tropical Trees

Recent work has suggested that in temperate and subtropical trees, leaf surface bacterial communities are distinctive to each individual tree species and dominated by Alpha- and Gammaproteobacteria. In order to understand how general this pattern is, we studied the phyllosphere bacterial community on leaves of six species of tropical trees at a rainforest arboretum in Malaysia. This represents the first detailed study of ‘true’ tropical lowland tree phyllosphere communities. Leaf surface DNA was extracted and pyrosequenced targeting the V1–V3 region of 16S rRNA gene. As was previously found in temperate and subtropical trees, each tree species had a distinctive bacterial community on its leaves, clustering separately from other tree species in an ordination analysis. Bacterial communities in the phyllosphere were unique to plant leaves in that very few operational taxonomic units (0.5%) co-occurred in the surrounding soil environment. A novel and distinctive aspect of tropical phyllosphere communities is that Acidobacteria were one of the most abundant phyla across all samples (on average, 17%), a pattern not previously recognized. Sequences belonging to Acidobacteria were classified into subgroups 1–6 among known 24 subdivisions, and subgroup 1 (84%) was the most abundant group, followed by subgroup 3 (15%). The high abundance of Acidobacteria on leaves of tropical trees indicates that there is a strong relationship between host plants and Acidobacteria in tropical rain forest, which needs to be investigated further. The similarity of phyllosphere bacterial communities amongst the tree species sampled shows a significant tendency to follow host plant phylogeny, with more similar communities on more closely related hosts.     

How do we use language? Shared patterns in the frequency of word use across 17 world languages

We present data from 17 languages on the frequency with which a common set of words is used in everyday language. The languages are drawn from six language families representing 65 per cent of the world's 7000 languages. Our data were collected from linguistic corpora that record frequencies of use for the 200 meanings in the widely used Swadesh fundamental vocabulary. Our interest is to assess evidence for shared patterns of language use around the world, and for the relationship of language use to rates of lexical replacement, defined as the replacement of a word by a new unrelated or non-cognate word. Frequencies of use for words in the Swadesh list range from just a few per million words of speech to 191 000 or more. The average inter-correlation among languages in the frequency of use across the 200 words is 0.73 (p < 0.0001). The first principal component of these data accounts for 70 per cent of the variance in frequency of use. Elsewhere, we have shown that frequently used words in the Indo-European languages tend to be more conserved, and that this relationship holds separately for different parts of speech. A regression model combining the principal factor loadings derived from the worldwide sample along with their part of speech predicts 46 per cent of the variance in the rates of lexical replacement in the Indo-European languages. This suggests that Indo-European lexical replacement rates might be broadly representative of worldwide rates of change. Evidence for this speculation comes from using the same factor loadings and part-of-speech categories to predict a word's position in a list of 110 words ranked from slowest to most rapidly evolving among 14 of the world's language families. This regression model accounts for 30 per cent of the variance. Our results point to a remarkable regularity in the way that human speakers use language, and hint that the words for a shared set of meanings have been slowly evolving and others more rapidly evolving throughout human history.     

How Accurate and Robust Are the Phylogenetic Estimates of Austronesian Language Relationships?

We recently used computational phylogenetic methods on lexical data to test between two scenarios for the peopling of the Pacific. Our analyses of lexical data supported a pulse-pause scenario of Pacific settlement in which the Austronesian speakers originated in Taiwan around 5,200 years ago and rapidly spread through the Pacific in a series of expansion pulses and settlement pauses. We claimed that there was high congruence between traditional language subgroups and those observed in the language phylogenies, and that the estimated age of the Austronesian expansion at 5,200 years ago was consistent with the archaeological evidence. However, the congruence between the language phylogenies and the evidence from historical linguistics was not quantitatively assessed using tree comparison metrics. The robustness of the divergence time estimates to different calibration points was also not investigated exhaustively. Here we address these limitations by using a systematic tree comparison metric to calculate the similarity between the Bayesian phylogenetic trees and the subgroups proposed by historical linguistics, and by re-estimating the age of the Austronesian expansion using only the most robust calibrations. The results show that the Austronesian language phylogenies are highly congruent with the traditional subgroupings, and the date estimates are robust even when calculated using a restricted set of historical calibrations.     

Genetic variation in a tropical tree species influences the associated epiphytic plant and invertebrate communities in a complex forest ecosystem

Genetic differences among tree species, their hybrids and within tree species are known to influence associated ecological communities and ecosystem processes in areas of limited species diversity. The extent to which this same phenomenon occurs based on genetic variation within a single tree species, in a diverse complex ecosystem such as a tropical forest, is unknown. The level of biodiversity and complexity of the ecosystem may reduce the impact of a single tree species on associated communities. We assessed the influence of within-species genetic variation in the tree Brosimum alicastrum (Moraceae) on associated epiphytic and invertebrate communities in a neotropical rainforest. We found a significant positive association between genetic distance of trees and community difference of the epiphytic plants growing on the tree, the invertebrates living among the leaf litter around the base of the tree, and the invertebrates found on the tree trunk. This means that the more genetically similar trees are host to more similar epiphyte and invertebrate communities. Our work has implications for whole ecosystem conservation management, since maintaining sufficient genetic diversity at the primary producer level will enhance species diversity of other plants and animals.     

mtDNA variability in two Bantu‐speaking populations (Shona and Hutu) from Eastern Africa: Implications for peopling and migration patterns in sub‐Saharan Africa

In this study, we report novel data on mitochondrial DNA in two of the largest eastern Bantu‐speaking populations, the Shona from Zimbabwe and the Hutu from Rwanda. The goal is to evaluate the genetic relationships of these two ethnic groups with other Bantu‐speaking populations. Moreover, by comparing our data with those from other Niger‐Congo speaking populations, we aim to clarify some aspects of evolutionary and demographic processes accompanying the spread of Bantu languages in sub‐Saharan Africa and to test if patterns of genetic variation fit with models of population expansion based on linguistic and archeological data. The results indicate that the Shona and Hutu are closely related to the other Bantu‐speaking populations. However, there are some differences in haplogroup composition between the two populations, mainly due to different genetic contributions from neighboring populations. This result is confirmed by estimates of migration rates which show high levels of gene flow not only between pairs of Bantu‐speaking populations, but also between Bantu and non‐Bantu speakers. The observed pattern of genetic variability (high genetic homogeneity and high levels of gene flow) supports a linguistic model suggesting a gradual spread of Bantu‐speakers, with strong interactions between the different lines of Bantu‐speaker descent, and is also in agreement with recent archeological findings. In conclusion, our data emphasize the role that population admixture has played at different times and to varying degrees in the dispersal of Bantu languages. Am J Phys Anthropol, 2009. © 2009 Wiley‐Liss, Inc.     

Mitochondrial DNA diversity in two ethnic groups in Southeastern Kenya: Perspectives from the northeastern periphery of the bantu expansion

The Bantu languages are widely distributed throughout sub‐Saharan Africa. Genetic research supports linguists and historians who argue that migration played an important role in the spread of this language family, but the genetic data also indicates a more complex process involving substantial gene flow with resident populations. In order to understand the Bantu expansion process in east Africa, mtDNA hypervariable region I variation in 352 individuals from the Taita and Mijikenda ethnic groups was analyzed, and we evaluated the interactions that took place between the Bantu‐ and non‐Bantu‐speaking populations in east Africa. The Taita and Mijikenda are Bantu‐speaking agropastoralists from southeastern Kenya, at least some of whose ancestors probably migrated into the area as part of Bantu migrations that began around 3,000 BCE. Our analyses indicate that they show some distinctive differences that reflect their unique cultural histories. The Taita are genetically more diverse than the Mijikenda with larger estimates of genetic diversity. The Taita cluster with other east African groups, having high frequencies of haplogroups from that region, while the Mijikenda have high frequencies of central African haplogroups and cluster more closely with central African Bantu‐speaking groups. The non‐Bantu speakers who lived in southeastern Kenya before Bantu speaking groups arrived were at least partially incorporated into what are now Bantu‐speaking Taita groups. In contrast, gene flow from non‐Bantu speakers into the Mijikenda was more limited. These results suggest a more complex demographic history where the nature of Bantu and non‐Bantu interactions varied throughout the area. Am J Phys Anthropol 150:482–491, 2013. © 2013 Wiley Periodicals, Inc.     

Linking Trees on Farms with Biodiversity Conservation in Subsistence Farming Systems in Nepal

The Kingdom of Nepal extends 800 km east to west along the southern slopes of the Himalayas. Because of its biogeographical position, Nepal contains biological elements of both the Indo-Malayan and Palaearctic realms. Beside the trees in forest ecosystems, a large number of tree species are maintained on farms as part of subsistence farming systems. The role of these trees in ensuring the sustainability of agricultural production and the importance of traditional farming systems for the conservation of crop diversity have been well documented. However, the status of farm trees and their role in biodiversity conservation are poorly documented. This paper presents a case study of two villages in the western middle hills of Nepal. It highlights the role of traditional agroforestry practices for the conservation of tree diversity and argues that farms can be considered biodiversity reservoirs. Farm trees help to reduce pressure on community and government forests and create a favorable environment for many plant and animal species. Farm trees also provide social functions in that households with many farm trees no longer exercise their communal rights to extract grass products from community forests, which in turn benefits poor and disadvantaged households. The paper discusses possibilities to improve the role of farm trees in biodiversity conservation. It argues for the development of mechanisms such as tax exemptions and conservation credits that provide benefits to rural communities as compensation for their local and global environmental services including biodiversity conservation.     

Language as a multimodal phenomenon: implications for language learning,processing and evolution

Our understanding of the cognitive and neural underpinnings of language has traditionally been firmly based on spoken Indo-European languages and on language studied as speech or text. However, in face-to-face communication, language is multimodal: speech signals are invariably accompanied by visual information on the face and in manual gestures, and sign languages deploy multiple channels (hands, face and body) in utterance construction. Moreover, the narrow focus on spoken Indo-European languages has entrenched the assumption that language is comprised wholly by an arbitrary system of symbols and rules. However, iconicity (i.e. resemblance between aspects of communicative form and meaning) is also present: speakers use iconic gestures when they speak; many non-Indo-European spoken languages exhibit a substantial amount of iconicity in word forms and, finally, iconicity is the norm, rather than the exception in sign languages. This introduction provides the motivation for taking a multimodal approach to the study of language learning, processing and evolution, and discusses the broad implications of shifting our current dominant approaches and assumptions to encompass multimodal expression in both signed and spoken languages.     

How to become a “Kwa” noun

An important problem of comparative Niger-Congo morphology is understanding the processes that relate word structures in languages of the isolating ??Kwa?? type to those of the agglutinating ??Bantu?? type. A salient sub-problem of this larger morphological puzzle is charting the connection between the noun class systems of the Kwa-type languages which, at one extreme, can lack such classes entirely, against those of the Bantu type which, at the other extreme, are famously elaborated. This issue is examined by looking at a range of ways that Niger-Congo noun class systems have been observed to diverge from the canonical Bantu type. The main conclusion of this study is that Niger-Congo noun class systems are quite robust, in the sense that loss of one part of the system need not be correlated with loss of the other parts. This suggests that the level of noun class attrition found in Kwa languages was not a historically ??natural?? event and also has implications for models of agreement and inflectional morphology.