Saturday, October 27, 2018

Additional evidence for early modern human morphological diversity in Southeast Asia at Tam Pa Ling, Laos

Laura Shackelford
Fabrice Demeter
Kira Westaway
Philippe Duringer
Jean-Luc Ponche
Thongsa Sayavongkhamdy
Jian-Xin Zhao
Lani Barnes
Marc Boyon
Phonephanh Sichanthongtip
Frank Sénégas
Elise Patole-Edoumba
Yves Coppens
Jean Dumoncel
Anne-Marie Bacon
Quaternary International
Volume 466, Part A,
February 2018


Despite its geographic primacy as the intersection of dispersal paths to Australasia, mainland Southeast Asia has played little role in scenarios of early human migrations. Tam Pa Ling (TPL), a cave site in northern Laos, is the source of early modern human fossils – a partial cranium (TPL1) and a complete mandible (TPL2) – that represent the earliest anatomically modern humans in continental Southeast Asia and introduce new migration routes into the region during Marine Isotope Stage 3. In the current analysis, a new partial mandible from the site, TPL 3, is introduced, described, and evaluated using geometric morphometrics in the context of Pleistocene archaic, early modern and Holocene humans. In addition, the sedimentary context of TPL3 is described and refined through further quartz single-grain optically stimulated luminescence (SG-OSL) measurements and additional feldspar post-infrared infra-red stimulated luminescence (pIR-IRSL) techniques.

The TPL3 mandible has discrete traits similar to eastern Asian early modern humans, including a well-developed chin and a lack of lateral corporal robusticity. It does, however, retain a relatively broad anterior mandibular arch that is more commonly associated with archaic populations. In this way, it is similar to the TPL2 mandible as well as other similarly-aged fossils from the region that show a mixture of archaic and derived traits. The combined quartz SG-OSL and feldspar pIR-IRSL techniques provide an age range of 70 ± 8–48 ± 5 ka for the depositional age of the layer containing the TPL3 mandible. This upper age estimate is ∼20 ka older than the depositional ages for the TPL1 and 2 fossils, extending the upper limit for the currently excavated sedimentary and associated fossil evidence.

Friday, October 26, 2018

'Are'are Music

Preview for the Hugo Zemp 1979 documentary:
'Are'are Music and Shaping Bamboo

Are'are People of the island of Malaita

Further Reference:

Hugo Zemp, 'Are'are Classification of Musical Types and Instruments,
Vol. 22, No. 1 (Jan., 1978), pp. 37-67 

Wednesday, October 24, 2018

Far from empowering young women, the internet silences their voices

Jane Dudman
The Guardian
October 23, 2018

Young EU citizens have almost equal digital skills and online access: 93% of boys and 92% of girls aged 16-24 use technology every day. But there is a marked difference in their digital confidence, with 73% of boys aged 15-16 saying they would be comfortable using less familiar digital devices, compared to 49% of girls the same age.

Young women and men also behave very differently online, with men taking a higher part in public activities such as debates. They are also more likely to post their opinions on social and political issues or take part in online voting. There is increasing evidence, too, that young women censor themselves to avoid being judged harshly online.

“I’m too frightened to express my opinions online,” says one 15-year-old Swedish girl in the EIGE survey. “I keep them to myself because I know that there will always be someone out there who won’t like what I post.” Another 15-year-old girl expressed a similar view: “The things you get hated for are your opinions and sexual things, if you’re a girl.”

Tuesday, October 23, 2018

A framework for enhancing ethical genomic research with Indigenous communities

Katrina G. Claw,
Matthew Z. Anderson,
Rene L. Begay,
Krystal S. Tsosie,
Keolu Fox,
Nanibaa’ A. Garrison &
Summer internship for Indigenous peoples in Genomics (SING) Consortium

Nature Communications,
volume 9,
Article number: 2957 (2018)
(Link) open access


Integration of genomic technology into healthcare settings establishes new capabilities to predict disease susceptibility and optimize treatment regimes. Yet, Indigenous peoples remain starkly underrepresented in genetic and clinical health research and are unlikely to benefit from such efforts. To foster collaboration with Indigenous communities, we propose six principles for ethical engagement in genomic research: understand existing regulations, foster collaboration, build cultural competency, improve research transparency, support capacity building, and disseminate research findings. Inclusion of underrepresented communities in genomic research has the potential to expand our understanding of genomic influences on health and improve clinical approaches for all populations.

Wednesday, October 17, 2018

Cranio-morphometric and aDNA corroboration of the Austronesian dispersal model in ancient Island Southeast Asia: Support from Gua Harimau, Indonesia

Hirofumi Matsumura, 
Ken-ichi Shinoda, 
Truman Shimanjuntak, 
Adhi Agus Oktaviana, 
Sofwan Noerwidi, 
Harry Octavianus Sofian, 
Dyah Prastiningtyas, 
Lan Cuong Nguyen, 
Tsuneo Kakuda, 
Hideaki Kanzawa-Kiriyama, 
Noboru Adachi, 
Hsiao-chun Hung, 
Xuechun Fan, 
Xiujie Wu, 
Anna Willis, 
Marc F. Oxenham 
June 22, 2018


The Austronesian language is spread from Madagascar in the west, Island Southeast Asia (ISEA) in the east (e.g. the Philippines and Indonesian archipelagoes) and throughout the Pacific, as far east as Easter Island. While it seems clear that the remote ancestors of Austronesian speakers originated in Southern China, and migrated to Taiwan with the development of rice farming by c. 5500 BP and onto the northern Philippines by c. 4000 BP (the Austronesian Dispersal Hypothesis or ADH), we know very little about the origins and emergence of Austronesian speakers in the Indonesian Archipelago. Using a combination of cranial morphometric and ancient mtDNA analyses on a new dataset from Gua Hairmau, that spans the pre-Neolithic through to Metal Period (5712—5591 cal BP to 1864—1719 cal BP), we rigorously test the validity of the ADH in ISEA. A morphometric analysis of 23 adult male crania, using 16 of Martin’s standard measurements, was carried out with results compared to an East and Southeast Asian dataset of 30 sample populations spanning the Late Pleistocene through to Metal Period, in addition to 39 modern samples from East and Southeast Asia, near Oceania and Australia. Further, 20 samples were analyzed for ancient mtDNA and assigned to identified haplogroups. We demonstrate that the archaeological human remains from Gua Harimau cave, Sumatra, Indonesia provide clear evidence for at least two (cranio-morphometrically defined) and perhaps even three (in the context of the ancient mtDNA results) distinct populations from two separate time periods. The results of these analyses provide substantive support for the ADH model in explaining the origins and population history of ISEA peoples.

Fig 4. A neighbour net splits tree generated from a Q-mode correlation coefficients matrix, based on the craniometric data, comparing the archaeological and modern sample populations.
The Late (pre-Neolithic) and Early Gua Harimau samples are boxed for ease of identification.

Sunday, October 14, 2018

The Hoabinhian of Southeast Asia and its Relationship to Regional Pleistocene Lithic Technologies

Ben Marwick
Chapter 4, peer-reviewed and published in:
Robinson, Erick, Sellet, Frederic (Eds.) 2018. Lithic Technological Organization Paleoenvironmental Change Global and Diachronic Perspectives. Springer International Publishing.


The Hoabinhian is a distinctive Pleistocene stone artefact technology of mainland and island Southeast Asia. Its relationships to key patterns of technological change both at a global scale and in adjacent regions such as East Asia, South Asia and Australia are currently poorly understood. These key patterns are important indicators of evolutionary and demographic change in human prehistory so our understanding of the Hoabinhian may be substantially enhanced by examining these relationships. In this paper I present new evidence of ancient Hoabinhian technology from Northwest Thailand and examine connections between Hoabinhian technology and the innovation of other important Pleistocene technological processes such as radial core geometry. I present some claims about the evolutionary significance of the Hoabinhian and recommend future research priorities. 


This survey of key late Pleistocene lithic assemblages in three regions of the eastern hemisphere allows us to engage with a few global debates on technological change. The data presented here lend little support to Mellars’ (2006) claim of a single hominin dispersal responsible for the appearance of microliths out of Africa. The proliferation of microliths in China can be traced to a set of much earlier local technologies that pre-date Mellars’ dispersal event. Microlithic technology in South Asia is most parsimoniously explained as an in situ development from earlier technologies in response to a reduction in summer rainfall and temperature, producing semi-glacial mosaic environments and desert expansion (Hiscock et al., 2011). The timing of the proliferation of microliths in both South and East Asia around 35,000 cal. BP is also not consistent with Mellars’ claim for 60-50,000 cal. BP for the expansion of microlith-bearing modern humans. Similarly, anatomically modern humans appear in both South and East Asia before the proliferation of microliths. These details point to microliths as autochthonous developments rather than part of a migration package.
The absence of microliths from assemblages in Southeast Asia further challenges the model of microlith-bearing modern humans. The assemblages at Tham Lod and Jerimalai are most parsimoniously explained as products of modern humans and yet little of the distinctive archaeological evidence that Mellars’ claims to be characteristic of modern humans is present. A similar situation has been noted in Australian Pleistocene lithic assemblages by Brumm and Moore (2005). In brief, they argue that the absence of classical indicators of modern humans is not evidence of their absence, but of a need to reconsider how modernity is defined. Drawing further on the Australian Pleistocene, Langley et al. (2011) conclude that the search for singular ‘packages’ of behavioral modernity is misguided and the diversity of specific manifestations of early modern human populations was more likely a result of differences in population size and density, interaction and historical contingency. The absence of typically modern human artefacts such as microlithic blades and backed artefacts in Southeast Asian assemblages such as Jerimalai and Tham Lod reinforces the importance of these kinds of artefacts as adaptions to specific environmental and demographic conditions (Clarkson et al., 2009; Petraglia et al., 2009a).
The data discussed here are also relevant to the question of hominin taxonomic affiliations of lithic assemblages. The similarities between the pre and post YTT event have been interpreted by Clarkson et al. (2012) as indicating that modern humans produced both assemblages, although this claim has been challenged by (Mellars et al., 2013) using genetic evidence. The many points of similarity between Jerimalai, Liang Bua and Mata Menge suggest that as many as three species of hominin were producing very similar assemblages. The overall picture is that these assemblages give little support to the claims of Foley and Lahr (2003) that different hominid species are associated with different stone artefact technologies. The implications are that either the hominins active during the Pleistocene in Southeast Asia were much more similar in their behavioral and cognitive expressions than their taxonomic affiliations would imply, or that stone artefact assemblages such as those found in this region either lack sufficient detail (or current approaches to describing the assemblages sufficient detail) to discriminate between the technological behaviors of the three hominins.
An interesting common element in the assemblages here are radial cores, which are often associated with Levallois flake products. Even though Tham Lod has radial cores overlapping with the Hoabinhian, it is tempting to identify the centripetal flaking that characterizes sumatraliths as a variant of radial core technology. The Hoabinhian may thus be a regional expression, as a divergent descendant, of a very widespread radial method of core reduction. This invites the question of whether radial cores and analogous forms are a result of technological behaviors that are homologous, having persisted from when radial cores first proliferated in Africa around 300 Ka (Ambrose, 2001). The alternative possibility is that radial cores developed as homoplasy or convergent technological evolution (Brumm et al., 2009) as local adaptations. Answering this question is important for accurately contextualizing the relationships between the three major regions discussed here. Clarkson et al. (2012) have had some success using stone artifact data to differentiate modern human core technology from Neanderthals and late Acheulean populations. Future work with their methods on early Hoabinhian assemblages may help to resolve these questions. 
The possibility of an ambiguous trace of the Levallois technology in the Tham Lod and other Southeast Asian assembles raises the question of why there is no obvious evidence of Levallois technology in Southeast Asia. The frequent association of bifacial pieces in Acheulean assemblages preceding Levallois technology elsewhere in the world (Lycett, 2007; Schick, 2002) is also problematic for Southeast Asia, where Pleistocene bifaces are less frequent (Moncel et al., 2017). Two current hypotheses are relevant to these questions. First, the hypothesis that complex stone artifacts were not produced because of the abundance and ease of using bamboo to make complex tools (Pope, 1989). West and Louys (2007) have shown experimentally that it is possible to distinguish cuts marks on bone made by bamboo from marks made by stone, but these methods are yet to be applied to an archaeological assemblage. Similarly, Bar-Yosef et al. (2012) have replicated complex forms such as bifacial hand axes and preferential Levallois cores and flakes on raw materials common in Pleistocene Southeast Asian assemblages such as metaquarzite and indurated sandstone. They further experimented with bamboo tools for cutting pig meat and hide and found that while bamboo knives were effective at meat-cutting, they were ineffective for cutting the skin.
These two experimental studies do not directly confirm the bamboo hypothesis, but indicate that an important priority in future research should be the analysis of microscopic cut-marks, use-wear and residues on archaeological fauna and stone artifacts to determine to role of bamboo tools during the Pleistocene. Of relevance to the Hoabinhian is the observation by Bar-Yosef et al. (2012) that unifacially flaked choppers were more efficient than bifaces at felling bamboo. This lends support to the long-standing claim that Hoabinhian technology was optimized for bamboo harvesting (Gorman, 1972). We might speculate that the technology perhaps became abundant during biogeographical events related to the expansion of the temperate and paleotropical woody bamboo clades (cf. Kelchner, 2013) during cooler and dryer episodes in Southeast Asia. Current phylogenetic work on the timing of major biogeographical changes in Southeast Asian bamboo populations during the late Pleistocene will help test this hypothesis.
             A second hypothesis for the low frequency of Levallois and bifaces in Southeast Asia is that the Pleistocene hunter-gatherer populations were too small for widespread adoption and persistence of shared complex technologies (Lycett, 2007). Simulations by Shennan (2001) and numerical analysis by Heinrich (2004) demonstrate that larger populations result in more chances of artifact makers copying from more skilled makers. Higher degrees of social interconnectedness in larger populations similarly result in a higher likelihood of encountering a given craft skill and the greater regularity of such encounters. In smaller populations with lower degrees of interconnectedness encounters of highly skilled craftspeople are rarer, opportunities from copying are fewer and the probability of technological loss is higher. Lycett (2007) argues that the overall low density of artifacts and site densities in East Asia indicate lower demographic conditions which were the primary constraint leading to small numbers of bifaces and assemblages with Levallois technologies. While this is an intriguing hypothesis, it is currently rests on a problematic ‘absence of evidence’ argument, where the absence of Levallois is taken to indicate the presence of a constraint, rather than simply a smaller sample size. It is also difficult to persuasively test with the currently available archaeological data, given uncertainties involved in inferring demography from distributions of small numbers of sites and dates (ie. <1000, cf. Buchanan et al., 2008). The most persuasive test of this hypothesis is likely to come from genetic material recovered from ancient human remains. These are an especially rare type of find in any region, but recent developments in high-throughput sequencing of highly fragmented and morphologically indistinct bone may improve recovery of human aDNA from archaeological assemblages (Murray et al., 2013). Genetic evidence currently available from South Asia poses a challenge to this hypothesis, suggesting relatively high populations during the late Pleistocene (Atkinson et al., 2008).

Saturday, October 13, 2018

Ethnographic and Archaeological Correlates for a Mainland Southeast Asia (MSEA) Linguistic Area

Roger Blench
Conference ‘Beyond the Sanskrit Cosmopolis’. ISEAS, Singapore,
November 2013(Link)

"One of the most characteristic instruments of the MSEA area is the free-reed mouth-organ (Blench in press b). Using the same principle as the European harmonica, free-reeds are found in horns and single tube flute-like instruments. The principle of the free-reed was confined to a specific geographical area in Southeast Asia, before its worldwide diffusion in the last two centuries. Free-reed instruments are widely distributed and morphologically highly diverse, pointing to several millennia of evolution, as confirmed by archaeological evidence. However, most commonly they are found in the free-reed mouth-organ which consists of groups of at least five stopped pipes.  The arrangement of the pipes allows the player to sound block chords, which form the underlying metrical frame of large Chinese ensembles. Free-reed mouth-organs are played almost everywhere in the region, and the oldest types have a spherical gourd resonator."

". . . there is a deep-level lithic culture which corresponds extremely well with the boundaries of MSEA, i.e. the so-called ‘Sumatraliths’ that characterise the Hoabinhian technocomplex. Sumatraliths are a type of stone tool, often made from river pebbles, and very roughly shaped. The precise use of Sumatraliths is still under discussion, but there is a growing consensus that their main use was to process bamboos, rattans and other wood-like plants (Blench 2013a)."

The oldest Hoabinhian technocomplex in Asia (43.5 ka) at Xiaodong rockshelter, Yunnan Province, southwest China

Xueping Ji, Kathleen Kuman, R.J. Clark, Hubert Forestier, Yinghua Li, Juan Ma, Kaiwei Qiu, Hao Li, Yun Wu
Quaternary International
Volume 400, 2 May 2016, Pages 166-174


The Hoabinhian is the most representative technocomplex in Southeast Asian prehistory for the later hunter–gatherer period. As a mainland technology based exclusively on seasonal tropical environments, this core-tool culture was previously defined in northern Vietnam in 1932 and characterized originally by its large, flat and long, largely unifacial cobble tools associated with tropical forest fauna. The recent discoveries and dates obtained at Xiaodong rockshelter in Yunnan Province (southwest China) allow us to discuss the origin and the homeland of this singular Asian technocomplex which spread to Southeast Asia during the end of the Late Upper Pleistocene. Here we present the first Chinese Hoabinhian lithic implements in their stratigraphic and chronological context within a rockshelter site, and we address the question of the dispersal of modern humans from South China to Southeast Asia.

from the Conclusion:

Although most of the tools belong to a selected collection from disturbed and surface deposits and just a few pieces are from the test excavations, the type most diagnostic of the Hoabinhian technocomplex-the core-axe/adze-is present, including in the bottom layer of our test trench.  The high-backed, plano-convex shape of these heavy duty tools is distinctive, with cortex often forming the flat surface.  The function of such tools is widely considered to be for woodworking, with the working of bamboo in particular a most likely possibility (Gorman, 1971; Thaw, 1971).  In this respect, the type (although unrelated culturally and technologically) has some similarities with the Sangoan industry core-axes produced during the late Middle Pleistocene in the more wooded habitats of central Africa (Clark, 1970).  The best description of such tools, also argued to be used in woodworking, was published by Clark (2000) for the Kalambo Falls site in Zambia.  For the Hoabinhian, the adze-like nature of the working ends of these high-backed tools is particularly diagnostic of woodworking.  The blunted edges and tapering bases of the typical core-axe/adze could assist use as either a hand-held tool or a tool fixed into some form of haft.  The wood-working function of the site is also suggested by the associated heavy-duty core scrapers with steep working edges, as well as the denticulated and notched scrapers and choppers.

Thursday, October 11, 2018

Bamboo Diversity and Traditional Uses in Yunnan, China

Yang Yuming, Wang Kanglin, Wang Kanglin, Hao Jiming
Mountain Research and Development
24(2): 157-165
May 2004


Bamboo is a giant grass that takes on tree-like functions in forest ecosystems. Around 75 genera and 1250 species of bamboo are known to exist throughout the world. Five hundred species in 40 genera are recorded in China, mostly in the monsoon areas of south and southwest China. Of these, 250 species in 29 genera grow naturally in the mountainous province of Yunnan, in the Chinese Himalayan region. Bamboo has a long history of being used for multiple purposes by various mountain communities in China. Among others, bamboo has served-and still serves-as construction material, fiber, food, material for agricultural tools, utensils, and music instruments, as well as ornamental plants. Yunnan as a landlocked mountain province in southwest China holds a great number of species in its natural bamboo forests. This article presents the diversity of bamboo species and of their utilization in Yunnan Province, China.

Tuesday, October 9, 2018

Ecological Contingency Accounts for Earliest Seagoing in the Western Pacific Ocean

Atholl Anderson
The Journal of Island and Coastal Archaeology
0:1-11, 2017


Seagoing at 1 mya to Flores, and sea gaps of >50 km crossed by 47 kya to Sahul, are evidence of earlier maritime migration in the western Pacific than anywhere else. Current opinion attributes the latter to the influence of anatomically modern human cultural complexity on seagoing technology and practice, together with the impetus of serial resource depression. It is argued here that seagoing was unusually advantaged in the western Pacific by a fortuitous conjunction of the warmest seas with a ready availability of large-diameter bamboo that occurred as natural rafts, and which could also be constructed into rafts large enough to transport viable colonizing groups from island to island across Wallacea to Sahul. The geography of Wallacea allowed migration solely by drifting, and exploratory landscape learning might have produced landfalls on Sahul sooner than is implied by subsistence forcing of mobility. Seagoing by drifting raft was much harder from Sahul to the east because of the virtual absence of large-diameter bamboo and longer distance to fewer or small islands; colonization occurred much later.

Bamboo Rafts

Bamboo Raft, Laguna, Philippines

Thailand Bamboo Raft

Bangladesh Bamboo Raft,_Bangladesh.jpg

Bamboo Rafts, Parambikulam Tiger Reserve, Kerala

Bamboo Raft, Periyar, Kerala, India

The Manufacture and Terminology of Temiar Bamboo Rafts

G. W. H. Davison
Journal of the Malaysian Branch of the Royal Asiatic Society
Vol. 62, No. 1 (256) (1989), pp. 97-104

"A raft was constructed from 32 culms of the giant bamboo Gigantochloa scortechinii . . ."

Related post on this blog:

Towards a complete generic-level plastid phylogeny of the paleotropical woody bamboos (Poaceae: Bambusoideae)

Monday, October 8, 2018

Towards a complete generic-level plastid phylogeny of the paleotropical woody bamboos (Poaceae: Bambusoideae)

Meng-Yuan Zhou, Yu-Xiao Zhang, Thomas Haevermans, De-Zhu Li
TAXON 66 (3)
June 2017: 539–553


Paleotropical woody bamboos (PWB) are phylogenetically and taxonomically intractable. Because previous studies included deficient samples or lacked informative characters for tree construction, phylogenetic relationships within the PWB remain incompletely resolved. This study presents the most extensively sampled phylogeny of the PWB with 18 plastid regions and a sample of 144 (35%) ingroup species representing 40 (85%) genera and 8 outgroup species. Results confirmed Melocanninae as the earliest diverging lineage from the rest of the group, and Hickeliinae (including Nastus s.str.) and Racemobambosinae are separately placed within the PWB. Bambusinae is phylogenetically heterogeneous and consists of the Dinochloa-Greslania-Mullerochloa-Neololeba-Sphaeroambos (DGMNS) assemblage, Temburongia simplex, and the core Bambusinae. The core Bambusinae may be redefined to include a basal grade, which contains Kinabaluchloa, Holttumochloa, Bonia, Neomicrocalamus, Temochloa, Soejatmia and an unidentified taxon, and the Bambusa-Dendrocalamus-Gigantochloa (BDG) complex. The BDG complex is extremely diverse in morphology and is subdivided into six subclades. Within the Melocanninae, Davidsea, Neohouzeaua and Ochlandra are closely related to Schizostachyum. Phylogenetic relationships are mostly supported by morphological and geographical evidence. In addition, novel interpretations are provided in the redelimitation of some taxa.

Sunday, October 7, 2018

How many genera of vascular plants are endemic to New Caledonia? A critical review based on phylogenetic evidence

Yohan Pillon, Laure Barrabé, Sven Buerki 
Botanical Journal of the Linnean Society, 
Volume 183, Issue 2, 1 February 2017, Pages 177–198
(Link) free


New Caledonia is a biodiversity hotspot located in the south-western Pacific, well known for its rich, unique and endangered flora. The island flora has a high level of endemism not only at the species level (75%), but also at the generic and family (three endemic) levels. We review here the taxonomic validity of the c. 100 endemic New Caledonian genera of vascular plants (13%) by using the monophyly criterion based on the available phylogenetic data. As observed in other island floras, some of these genera were recovered nested in larger genera and are consequently likely to lose their rank. After a critical review, we concluded that the New Caledonian plant vascular flora contains between 62 and 91 endemic genera. This large variation in the number of endemic genera is mainly caused by a lack of DNA sequences (eight genera) and limited phylogenetic evidence. This work highlights gaps of knowledge that will have to be addressed to stabilize the taxonomy of the New Caledonian flora. Although this study shows that several genera are not monophyletic, New Caledonia still harbours more endemic genera than any other islands in the Pacific Ocean. Preliminary results indicate that the high level of endemism at higher taxonomic levels could be explained by an accumulation of relictual lineages, rather than adaptive radiations. Hypotheses explaining this phenomenon are provided in this study.

From the section on Poaceae:

"The dwarf bamboo Greslania Balansa (2/3) has been recovered with strong support as sister to an Australasian clade composed of Cyrtochloa S.Dransf., Dinochloa Buse, Mullerochloa K.M.Wong, Neololeba Widjaja, Parabambusa Widjaja, Pinga Widjaja and Sphaerobambos S.Dransf. (Chokthaweepanich, 2014)."

Phylogenetics and Evolution of the Paleotropical Woody Bamboos (Poaceae: Bambusoideae: Bambuseae)

Hathairat Chokthaweepanich
PhD Thesis
Iowa State University


The paleotropical woody bamboos (PWB) are members of Bambuseae (tropical woody bamboos) in subfamily Bambusoideae of the grass family (Poaceae). Although the PWB are important in ecological and economic functions, the phylogenetic relationships of the PWB remain equivocal. All four currently recognized subtribes of the PWB have never been completely examined for their evolutionary relationships and no morphological synapomorphy of the PWB has ever been revealed in previously published work. Six chloroplast sequences with coding (ndhF3' and matK) and non-coding (rpl16 intron, rps16 intron, trnD-trnT spacer, and trnT-trnL spacer) regions and three low-copy nuclear sequences (GPA1, Pabp1, and PvCel1) are analyzed to clarify the systematics and evolution of the PWB based on maximum parsimony, maximum likelihood and Bayesian inference. Forty morphological and anatomical characters are analyzed and mapped on the respective plastid and nuclear consensus trees. The objectives of this study are to: (1) test the monophyly of the PWB and all subtribes using chloroplast and low-copy nuclear sequences; (2) examine the phylogenetic relationships of all subtribes in the PWB clade based on molecular analyses; and (3) evaluate morphological evolution of the PWB.

The PWB form a monophyletic group with six clades [Bambusinae, Madagascan Hickeliinae, Melocanninae, Racemobambosinae, CDMNPS (Cyrtochloa-Dinochloa-Mullerochloa-Neololeba-Parabambusa-Pinga-Sphaerobambos) + Greslania, and Temburongia] and are sister to the Neotropical woody bamboo (NWB) clade based on plastid and nuclear markers. The relationships among subtribes of the PWB are different between two molecular topologies. The plastid analysis supports the monophyly of Melocanninae and its position as sister to the remaining PWB. Within PWB and exclusive of Melocanninae, Madagascan Hickeliinae form a monophyletic group sister to the rest of the PWB. Non-Madagascan Nastus are clustered with Racemobambosinae to form a clade. The core Bambusinae basically consists of a polytomy and the CDMNPS + Greslania clade is removed from subtribe Bambusinae and resolved as sister to the Racemobambosinae. In the nuclear analysis, the six lineages of the PWB from a polytomy, and six diverse genomic components are representative of Bambusoideae: one genome for the herbaceous bamboos (H), two genomes each for the temperate woody bamboos (genomes A and B) and the NWB (genomes C and D), and three genomes for the PWB (genomes C, D, and E). However, some taxa of the PWB show variation of genomic components from one to three genomes.

The incongruence between plastid and nuclear analyses is most strongly indicated at the tribal rank. The chloroplast analysis strongly supports the tropical woody bamboos as sister to the herbaceous bamboos, forming a tropical bamboo clade, with this clade sister to the temperate woody bamboos, whereas the low-copy nuclear analysis strongly supports the woody bamboo clade (tropical + temperate woody bamboos) as sister to the herbaceous bamboos.

Morphological investigation suggests that the presence of six stamens is a potential synapomorphy of the PWB. Melocanninae exhibit a narrow and glabrous ovary with an elongated and hollow style. The presence of extravaginal or infravaginal branching with a dipping nodal line is a unique character combination of Hickeliinae. Climbing bamboos with a wrinkled and prominent girdle diagnose the CDMNPS clade. Racemobambosinae + non-Madagascan Nastus possess a cap on the top of ovary apex with short style, whereas the Temburongia clade exhibits a blunt ovary apex without cap and moderate style length

Friday, October 5, 2018

Taxonomic interpretations of Australian native bamboos (Poaceae: Bambuseae) and their biogeographic implications

Donald C. Franklin
Telopea 12(2) 179–191
November, 2008
(Link) pdf


Australia’s three native bamboo species – Bambusa arnhemica, Mullerochloa moreheadiana and Neololeba atra – are restricted to northern Australia. The most parsimonious explanation for the occurrence of bamboo in Australia is that there have been at least three founder events from Asia, but other possibilities exist including an Australian radiation involving Neololeba and Mullerochloa. Bambusa arnhemica may be allied to the Asian B. blumeana which occurs as close as Timor. I summarise historical evidence and biogeographic patterns for B. arnhemica, and describe the flowering wave phenomenon in the species, evidence which collectively suggests that B. arnhemica is neither very ancient nor very recent in origin. Current studies of the population genetics of B. arnhemica, and of the relationship between Australian and Asian bamboos, may yield fascinating further biogeographic insights.

Related posts on this blog:

A 250 plastome phylogeny of the grass family (Poaceae): topological support under different data partition

Folk Classification and Conservation of Bamboo in Xishuangbanna, Yunnan, Southwest China

Australian Bamboo Uses and Manufacture

Were bamboo tools made in prehistoric Southeast Asia: An Experimental view from South China

Monday, October 1, 2018

Revising the hypodigm of Homo heidelbergensis: A view from the Eastern Mediterranean

Mirjana Roksandic, Predrag Radovic, Joshua Lindal
Quaternary International
November, 2017


The hominin mandible BH-1 from the Middle Pleistocene cave of Mala Balanica suggested the possibility that human populations in this part of the continent were not subject to the process of Neanderthalization observed in the west. We review the paleoanthropological evidence from the Central Balkans in the context of the Eastern Mediterranean geographic entity. The current hominin fossil record of the early Middle Pleistocene in the region suggests that Europe was inhabited by two different populations: a population in the west of the continent with derived Neanderthal morphology; and a more variable population in the east characterized by a combination of plesiomorphous and synapomorphous traits. We suggest that - in order to continue using the nomenclature of Homo heidelbergensis - the current hypodigm needs to be revised to include only the specimens from the latter group.

Page 3:

Central Europe

"The Bilzingsleben material consists of 28 skull fragments, one mandible and eight very fragmented teeth with worn edges, dated to 350-400 ka (Vlcek et al., 2000; Mania and Mai, 2001) or 250-200 ka (Eissmann, 1997). The ensemble is characterized by a preponderance of primitive (Erectine) traits, especially in the shape of the occipital and the supraorbital torus (Schwartz and Tattersall, 2002). The robust individuals I and II are represented by a series of skull fragments, and individual III is represented by a gracile mandible (Vlcek et al., 2000). According to Vlcek and colleagues, individuals I and II from Bilzingsleben are most similar to OH-9 from Olduvai Gorge, although similarities were also seen with Asian H. erectus specimens Zhoukoudian III and Sangiran III. The mandible is most similar to Zhoukoudian mandibles H1 and B1, but also has likeness to Arago II and XIII. However, a flat articular eminence on the temporal bone should be noted as a Neanderthal trait."

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The Apennine Peninsula

"Except for the well-preserved Ceprano calvarium, all of the early MP hominin specimens from Italy are highly fragmentary. At Castel di Guido, dated to 442 ±7 ka to 250-170 ka ( Michel et al., 2001), the occipital and postcranial elements show a combination of primitive traits that set it apart from Neanderthals, and some traits that resemble the Arago specimens. In particular, the temporal bone (Mallegni and Radmilli, 1988) shows a flat articular eminence - a trait considered to be distinctly Neanderthal (Martínez and Arsuaga, 1997; Martínez et al., 2006). According to Mallegni et al. (1983: 273), of the two femoral fragments, CdG-1 is comparable to the massive specimens from East Africa such as KNM-ER 999, ER 803, ER 737; they note, however, that the marked cortical thickness is a feature of all Homo material. According to the same authors, the proximal fragment CdG-2 resembles the Arago material. The rounded cross-section of the femoral diaphysis (Cesnola and Mallegni, 1996) is noted as Neanderthal-like (Chevalier et al., 2015). A relatively small fragment of the right superior region of the occipital bone (CdG-3) preserves a true continuous occipital torus with a well-defined supratoral sulcus that runs parallel to the torus. This sulcus does not extend to the suprainiac region but starts about 3 cm laterally, which together with the absence of the sulcus below the torus differentiate this specimen from Neanderthals and align it with Bilzingsleben, Petralona, and Vertesszoelloes (Mallegni et al., 1983: 273). The right maxilla (CdG-4), including the area from the intermaxillary suture to the mesial walls of the M1 buccal and M2 lingual alveoli, does not preserve many diagnostic elements. The reconstructed arcade is gently parabolic, with a high and narrow palate, a large distance between the incisive foramen and the anterior alveolar margin, and the presence of alveolar prognathism. The reconstructed width of the nasal aperture falls within the range of Chinese and Javanese Homo erectus. The overall puffing of the maxilla and a large maxillary sinus are listed as in line with Arago and Petralona. CdG-5 is a posteroinferior fragment of the right parietal, of remarkable thickness, with a well-defined angular torus and coarse indentations of the lambdoid sutural margin comparable to the Arago specimen. A similar angular torus is described for Petralona, and this feature, as well as other endocranial features including a parietooccipital sulcus, large branches of the meningeal vessels concentrated in the posterior part of the bone, and the absence of the groove for the transverse sinus at the mastoid angle, are all primitive traits found in Chinese Homo erectus and early European specimens. According to Mallegni et al. (1983) the specimens have a mixture of primitive erectus-like and derived Neanderthal-like features and show affinities with Arago, Petralona, Bilzingsleben and Vertesszoelloes."

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"Further to the east, The Kocabas site in Turkey is older than the period we are examining and the material from Karain is a bit younger; nevertheless, they are interesting for the present discussion. A partial calotte from Kocabas was a fortuitous find from the Uppert Travertine layer in a marble quarry situated in the vicinity of Kocabas village near Denizli in Southwestern Turkey. Based on a number of dating techniques, including paleomagnetism, cosmogenic nuclide concentrations on travertine sediments, and the burial age of pebbles above the travertine, Lebatard et al. (2014)proposed that the calotte is most likely between 1.1 and 1.3 Ma old, which is in agreement with a 1.2-1.6 Ma date for the Upper Travertine of Kocabas proposed by Khatib et al. (2014). A preliminary analysis by Kappelman et al. (2008) attributed Kocabas to Homo erectus s.l., a designation which has been supported by other researchers (Vialet et al., 2012, 2014; Aytek and Harvati, 2016). Vialet et al. (2014) suggested that the morphology of the specimen is closer to the African than to the Asian sample, while Aytek and Harvati (2016) find it to be most similar to MP Eurasian specimens including Zhoukoudian III, Ceprano, and Arago."


"Nadaouiyeh Aïn Askar near El Kowm in Syria (NAD-1) is an almost complete left parietal identified as H. erectus, dated to older than 500 ka (Schmid et al., 1997). This very thick, subrectangular bone, gently curved in the sagittal and coronal planes, with the temporal line relatively close to the sagittal suture, shows no derived traits. It is 12 mm thick in the region close to the parietal bossa and 9 mm near the coronal suture, with important thickening of the laminae and reduction of the diploe; 3 cm below bregma there is a thickening comparable to that in Chinese and Javanese material - accompanied by a parasagittal depression. Schmid (2015) superimposed the fragment onto Weidenreich's reconstruction of a H. erectus skull in order to demonstrate that the size and morphology of the Nadaouiyeh fragment corresponded exactly to the cast of skull II from Zhoukoudian."