Showing posts with label Phytoliths. Show all posts
Showing posts with label Phytoliths. Show all posts

Sunday, 1 January 2023

Examining stone tools for evidence of Rice harvesting in the Neolithic of the Lower Yangtze River Basin.

Rice cultivation is believed to have originated with the Neolithic Shangshan (上山) Culture in the Early Holocene of the Lower Yangtze Valley Basin, between about 10 000 and 8200 years before the present. The Neolithic Rice-farming tradition of this region persisted for several millennia, culminating in the Liangzhu (良渚) Culture of about 5300-4400 years ago. Archaeologists have tried to study this process in a variety of ways, although most investigations have concentrated on changes to Rice Plants over this time, or else changes in cultivation practices and their impact on the landscape.

In a paper published in the journal PLoS one on 7 December 2022, Jiajing Wang of the Department of Anthropology at Dartmouth College, Jiangping Zhu of the Pujiang Museum, Dongrong Lei of the Longyou Museum, and Leping Jiang of the Zhejiang Provincial Institute of Cultural Relics and Archaeology, and the Department of History of Science and Scientific Archaeology at the University of Science and Technology of China, present the results of a study of early Rice cultivation in the Lower Yangtze Valley that concentrated on the stone tools used by these early farmers to harvest Rice.

Harvesting techniques in themselves apply selective pressure to a cultivated crop. Most Cereal crops have lost the ability to shatter their own seed cases, effectively making them dependent on Human intervention. This development in early cereal crops has been widely linked to the use of sickles as a harvesting tool, as this selects for seeds with a tougher rachis (section of the stem to which the seed is attached), something only found in a small proportion of wild Cereal Plants. This fits well with the development of Cereal crops in the Fertile Crescent, where the onset of Cereal cropping is marked by the appearance of hundreds of flint sickles. However, these tools are absent at the Neolithic sites of the Lower Yangtze where Rice cultivation originated. This has raised the possibility that the initial cultivation of Rice was marked by a different form of harvesting, such as see picking or plucking and beating. Evidence to support such theories, however, is lacking.

Excavations of Shangshen Culture (10 000-8200 years before present) and Kuahuqiao (跨湖桥) Culture (8000-7000 years before present) culture sites in the Lower Yangtze River Basin have produced hundreds of crudely made flaked stone tools in the core-and-flake tradition widely found in South China during this interval. These have been recorded variously as scrapers, burins, or drills, based upon their shape, but little thought has been applied to their purpose. Wang et al. observe that many of these blades have sharp edges which would be suitable for harvesting crops, and, in order to test this hypothesis, analysed use-wear marks upon 52 of these blades, as well as examining them for phytolith residues. 

Phytoliths are produced by plants as a way of handling silica absorbed with water from the soil. Most plants produce phytoliths to some extent, but Monocotyledons, such as Grasses and Palms, which utilise phytoliths both as structural support and a defence against herbivores (silica phytoliths quickly wear down the teeth of animals which lack specific adaptations to deal with them), produce phytoliths which can often be used to identify the maker to genus or even species level. Rice Plants produce three highly distinctive types of phytolith, double-peak husk cells, Oryza-type bulliform leaf cells, and scooped parallel bilobates, which can make tools which have been used to work Rice easy to identify.

Cereal harvesting produces both distinctive wear traces upon a tool, and leaves a residue of plant tissues. Examination of stone tools that have been used to cut Poaceaen Plants (Grasses, Reeds, etc.) reveal that this activity produces a particularly distinct pattern of striations and polished areas, and that this remains constant across a variety of stone types that would have been available to Neolithic farmers in East Asia. 

The Shangshan Culture are thought to have been the first people to have cultivated Rice, and to have built permanent settlements in the Lower Yangtze River Valley. To date, Archaeologists have uncovered 19 settlements associated with the Shangshan Culture. The Kuahuqiao Culture dates from slightly later in the same area, and used many of the same technologies, but also engaged in more intense landscape modification with specialised Earth-working tools.

Wang et al. obtained stone tools from two sites in Zhejiang Province; Shangshan, the type location of the Shangshan Culture, and Hehuashan, which has both Shangshan and Kuahuqiao layers. The Shangshan settlement covered an area of about 30 000 m², and includes dwellings, storage and midden pits, and possible burials. It has produced a diverse stone tool assemblage, including flaked stone tools (82% of the material), grinding stones (15%) and polished stone tools (3%). Hehuashan is located about 80 km to the southwest of Shangshan, on a small hill, and is divided into two areas, the East and West. East Hehuashan has an (excavated) area of about 600 m², and includes houses, pits and Shangshan-style stone tools and ceramics, while West Hehuashan has an excavated area of about 150 m², including houses, pits, ditches, and Kuahuqiao-style stone tools and ceramics.

Archaeological background. (a) Location of Shangshan and Hehuashan. (b) Cultural history and rice domestication process in the Lower Yangtze River region. Wang et al. (2022).

At both sites, chipped flake tools form a significant part of the total lithic assemblage (82% at Shangshan and 57% at Hehuashan). At both sites these tools seem to have been made when needed, and discarded after use, with attempts at retouching being rare. None of these blades resemble the sickle blades developed in the Neolithic of the Fertile Crescent, or the denticulate sickles of the Huai River Valley of China (developed by the Peiligang (裴李崗) Culture of Henan Province in central China between 9000 and 7000 years before present, and thought to have been used to harvest Grasses and Rushes). These tools are made from locally available rock, including vitric tuff, river pebble, porphyry, and sandstone, with the blades made by the two cultures essentially identical in form.

The tools analysed in the tests came from the collections of the Pujiang and Longyou museums. Twenty tools were selected from the early phase of the Shangshan Culture (10 000 to 9000 years before present), eighteen from the late phase of the Shangshan Culture (9000 to 8200 years before present), and fourteen from the Kuahuqiao Culture (8000 to 7000 years before present). These averaged 42 mm in length and 42.8 mm in width, suggesting that they were hand held tools.

A selection of stone flakes analyzed in the study. (a)-(h) Flakes from the Shangshan culture; (i)-(l) Flakes from the Kuahuqiao culture. Red dots delineate working edges. Wang et al. (2022).

Use wear analysis of these tools suggested that they had been used for five different tasks; harvesting Siliceous Plants (i.e, Grasses or Reeds) (30 tools), cutting Animal tissue (7 tools), processing hard materials (10 tools), scraping woody material (6 tools), and an unidentified function (13 tools). 

Tools used for cutting Siliceous Plants generally have an uneven finish, with fine and uneven striations, and and extensive patches of polishing, on both raised and scored areas. Of the 30 tools in this category, 22 have striations perpendicular or diagonal to the cutting edge of the tool, suggested as being indicative of cutting or scraping, while 14 have striations parallel to the cutting edge, probably caused by slicing, and six have both.

Use-wear traces from cutting Grass. Wang et al. (2022).

Tools used to cut the soft tissues of Animals tend to develop patches of highly polished surface, but lack the striations produced by cutting Plant matter or hard material.

Use-wear traces from cutting Animal tissues. Wang et al. (2022).

Stone tools used to scrape woody material develop a rough polish on sinuous domed areas, with occasional deep, tapering striations perpendicular to the cutting edge.

Use-wear traces from scraping wood. Wang et al. (2022).

Scraping hard materials such as Animal bones produces a level topography and rough polish on a stone tool, with deep, even striations. The polished areas are concentrated on the highest parts of the tool, with reticulated areas between.

Use-wear traces from scraping Animal bones. Wang et al. (2022).

Interestingly, while the tools had all been identified as simple, single edged, single function tools, analysis of the striations identified secondary cutting edges on thirteen of them. These typically had a principle cutting edge with a low-angled concave surface, and a more steeply convex secondary cutting edge. In these tools the concave surface showed signs of having used to process Grasses, while the convex surface appeared to have been used to cut Animal tissues.

Fifty of the fifty two tools analysed yielded phytoliths, the vast majority of which came from types of Grass. These included silate or sinuate elongates from Grass leaves, stems, and inflorescences. More specific material included ouble-peak husk cells, Oryza-type bulliform leaf cells, and scooped parallel bilobates, all diagnostic of Rice, as well as Reed-type bulliforms, Sedge achenes, and articulated quadrilobates, which are associated with Panicoid Grasses (the group which includes Sugarcaine, Maize, Sorghum, and Switchgrass). This suggests strongly that a high proportion of the tools were used in the harvesting of Grassy crops, including Rice.

Phytolith morphotypes recovered from flakes from Shangshan and Hehuashan flakes. (a) Double-peak (Oryza, Rice); (b) Oryza-type bulliform (Oryza, Rice); (c) Scooped parallel bilobate (Oryzeae); (d) Reed-type bulliform (Phragmites); (e) Sedge achene (Cyperaceae); (f) Articulated quadrilobate (Panicoideae); (g) Rondel (Poaceae); (h) Saddle (Poaceae); (i) Scutiform, elongate sinuate, and rectangular (Poaceae). Wang et al. (2022).

The composition of the phytolith assemblage does, however, change significantly between the Shangshan and Kuahuqiao tools. In the Shangshan material, double-peak Rice husk phytoliths are found on 71% of the tool assemblage, while in the Kuahuqiao material this falls to 7%. The reverse trend for Rice leaf/stem phytoliths, which were recovered from 34% of the Shangshan tools, but 57% of the Kuahuqiao tools. The phytoliths of Reeds and Sedges also increased, with both being recovered from 8% of the Shangshan tools, while Reed phytoliths were found on 43% of the Kuahuqiao tools and Segde phytoliths on 29%. These changes probably reflect increased landscape management and a change in harvesting technique.

Wang et al. found a significant correlation between the abundance of phytoliths and wear patterns on blades, with blades showing wear patterns associated with Grass processing yielding a significantly higher proportion of Grass phytoliths. Furthermore, of the 30 blades interpreted as having been used in harvesting activities, 28 yielded Rice phytoliths. Comparing the phytolith and wear-use data suggests that the blades were used in two ways, as finger knives used to reap the panicles (seed clusters) at the top of the Plant, and as sickles used to cut the stems at their base.

Schematic representation of the use-wear traces and phytoliths from Rice-harvesting finger knives and sickles. Wang et al. (2022).

The best example of a blade used to cut Rice using the finger knife technique is specimen SS-10, a flaked blade made from a river pebble, with a slightly convex working edge 58 mm in length. This tool has numerous fine striations perpendicular to the blade edge, interpreted as indicative of cutting or scraping Grass. The blade edge of this tool yielded a large number of phytoliths of the Rice glume double-peak type, suggesting that it has been in direct contact with Rice seeds. In combination, this is interpreted as evidence of being used to cut the panicles at the top of the Rice Plant during harvesting.

In contrast, specimen HHS 38 is interpreted as having been used in sickle harvesting. The blade of this tool has a convex surface about 92 mm in length, with many fine striations parallel to its cutting edge, thought to be indicative of slicing or sawing through plant matter. This tool yielded no phytoliths associated with Rice husks, but numerous phytoliths with morphotypes associated with Grass leaves and stems, including Oryza-type bulliforms and scooped parallel bilobates from Rice leaves. This combination of sawing-type striations and leaf and stem phytoliths is considered to be indicative of cutting Rice stems close to the ground with a sickle-action.

Use-wear traces from rice harvesting flakes. (a) and (b) Use-wear traces from Artifact SS-10 exhibit fine striations perpendicular or diagonal to the cutting edge, suggesting a transverse motion. (c) and (d) Use-wear traces from Artifact HHS-38 are dominated by striations parallel to the cutting edge, suggesting a slicing motion. Wang et al. (2022).

Wang et al.'s findings strongly support the hypothesis that these flaked stone tools were used in the harvesting of Rice. Furthermore, they demonstrate a shift in the way in which Rice was harvested which had not previously been suspected. In the Shangshan Culture Rice appears to have predominantly been harvested using a finger-knife technique to remove seeds from the top of the Plant, while the use of sickles to cut the stems was rare, while in the Kuahuqiao Culture the use of sickles appears to have become the predominant method of harvesting Rice.

Although the materials from which tools are made have changed, both finger-knife and sickle harvesting of Rice have persisted into modern times. The finger-knife technique allows the farmer to repeatedly revisit the same Plant, which seems unduly arduous, but can be useful under conditions where the seeds of the Plant do not all ripen at the same time. This technique reduces the number of panickles (seed husks) that shatter during harvesting, thereby maximising the amount of Rice recovered from each Plant. Sickle harvesting enables the farmer to quickly cut through bunches of stems, gathering the entire harvest rapidly, with the seeds subsequently separated from the stems and leaves, which can be used for fuel or Animal feed. The choice of tool can be driven by spiritual beliefs as well as practical considerations, with many Southeast Asian cultures having resisted the introduction of the sickle for a long time, as sickles were perceived as harming the soul of the Rice, and therefore being less pleasing to the Rice Goddess.

Little is known (or is likely to be known) about the spiritual beliefs of the earliest Rice farmers of the Lower Yangtze Basin, but it is possible to make a comparison between the change in Rice harvesting technique and the process of domestication. The wild progenitor of modern East Asian Rice, Oryza rufipogen, grows in swampy wetlands, where its seeds ripen unevenly, and are shed into the (often deep muddy water. Trying to harvest this Plant with sickles has been shown to be very difficult experimentally, while a better yield can be achieved with finger knives. This would presumably also have been the case in the Early Holocene, when the process of Rice domestication began. Samples of Rice spikelets from Huxi, a late Shangshan Culture site, dated to between 9000 and 8400 years before the present, included only 8.7% non-shattering husks, which are predominant in modern Domestic Rice. This presumably indicates that the Rice fields of the Shangshan people were dominated by a type of Rice with unevern ripening and self-shattering husks, better harvested with the more selective finger knife technique. 

The succeeding Kuahuqiao people had a much more developed farming culture, originating about 8000 years ago, in which Rice was cultivated in managed Grass-Reed swamps, where water flow was controlled by irrigation systems and periodic burning was used. This system is thought to have been used to cultivate multiple wetland and forest Plants together, and to have promoted more synchronous Rice growth, making the harvesting of bundles of stems with a sickle a more viable option. At the Kuahuqiao type site (which gives its name to the culture), one of the key finds was a bunch of Rice stalks cut squarely at the base, something most likely to have been achieved with a sickle technique.

This adoption of sickle harvesting appears to have been roughly synchronous across several cultures in Ancient China about 8000 years ago. At the Peiligang Culture Jiahu Site in Henan Province, denticulate sickles begin to appear in large numbers during Phase III, which has been dated to between 8000 and 7500 years before the present. Wear use analyses carried out on these blades has suggested that they were used to harvest Grasses, and phytolith analysis of deposits from the site's residential areas suggests that Rice stalks were being brought into the settlement.

While finger knife harvesting requires more time within the Rice field, the product it produces is relatively 'clean' and requires little post-harvest processing. Sickle harvesting on the other hand, is likely to introduce more 'weeds' into the yield, which need to be removed later. The Kuahuqiao blades yield significantly more Sedge phytoliths than the Shangshan blades, which is probably evidence of a common weed being harvested alongside the Rice. There is also an increase in the number of Reed phytoliths at this time, but this may reflect the use of the new sickle technology to harvest an Reed stems as an additional crop. Items made from Reed stems have not, to date been discovered at any Kuahuqiao sites, but the use of Reeds in the Chinese neolithic is well documented at later Hemudu (河姆渡) Culture sites such as Tianluoshan, dated to about 7000 years ago, and the Hemudu type-site, dated to about 6000 years ago.

Both finger knife and sickle harvesting would produce selective pressure for non-shattering husks in Rice. With finger-knife harvesting, the Plants would be revisited numerous times, with the last crop obtained each year presumably contributing most to the seed crop saved for the following year. Since the final crop will include the highest proportion of non-shattering genotypes (which ripen slower), this would lead to a higher proportion these genotypes in each crop. With the introduction of sickle-harvesting, seeds with self-shattering husks would be more likely to be lost between the harvesting site and the processing site, effectively removing themselves from the domestic Rice genepool, applying another selective pressure in favour of non-shattering husks. 

Given these pressures, it is somewhat surprising that the domestication process for Rice took as long as it did (about 5000 years). This probably relates to the knowledge level of the farmers themselves, who would not have considered they were involved in a multi-generation domestication process, but rather have been concentrating on each years crop. Such farmers would not have intentionally selected non-shattering husked Rice as a seed crop, and may not always have selected their seed crop from the last harvest of the year. Furthermore, they are highly unlikely to have cleared a virgin plot to farm each year, so that self-seeded Rice, with self-shattering husks, would have been present in each year's crop.

For the Shangshan and Kuahuqiao cultures, Rice was probably only a small proportion of the diet, supplementing wild-gathered foods such as Acorns. This process of domestication proceeded through the following Hemudu Culture, with Rice probably not becoming fully domestic until the onset of the terminal-Neolithic Liangzhu Culture, considered to have been the first culture in southern China to form centralised states, about 5300 years ago.

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Wednesday, 22 August 2018

Identifying plant materials used in an Epipalaeolihic hut from Kharaneh IV.

During the Epipalaeolithic (between 23 000 and 11 500 years ago) hunter-gatherer populations around the Eastern Mediterranean began to build the oldest dwelling structures that we are aware off, beginning with plant-based huts that were probably only temporary structures to begin with, and developing to permanent stone-structures by the final Natufian phase of the period (14 500-11 500 years ago). As the earliest known purpose-built Human structures, the early Epipalaeolithic huts have been studied by archaeologists for some decades, with comparisons being made to those made by more recent hunter-gatherer populations in the African Kalahari Desert and Central Australian Desert, areas which are climatically similar to the modern day environments in Israel and Jordan, where these remains are found. However, the huts were constructed during the Late Pleistocene, when the climate was much cooler than it was today, with the environment where the Epipalaeolithic hut-builders lived likely to  have had a cold temperate climate, similar to northern Europe or North America today, which requires some thinking of the nature and purpose of these huts.

In a paper published in the Journal of Archaeological Anthropology on 30 March 2018, Monica Ramsey of the McDonald Institute for Archaeological Research at the University of Cambridge, Lisa Maher of the Department of Anthropology at the University of California, Berkely, Danielle Macdonald of the Department of Anthropology at the University of Tulsa, Dani Nadel of the Zinman Institute of Archaeology at the University of Haifa, and Arlene Rosen of the Department of Anthropology at rhe University of Texas at Austin, present an analysis of phytoliths (silica fragments produced by plants) recovered from  Kharaneh IV an Early Epipalaeolithic site in the Azraq Basin in Jordan.

Phytoliths are produced by plants as a way of handling silica absorbed with water from the soil. Most plants produce phytoliths to some extent, but Monocotyledons, such as Grasses and Palms, which utilise phytoliths both as structural support and a defence against herbivores (silica phytoliths quickly wear down the teeth of animals which lack specific adaptations to deal with them), produce phytoliths which can often be used to identify the maker to genus or even species level.

The Kharaneh IV site was also discovered in the 1980s, prior to which it was covered by wetlands, that were lost due to heavy water withdrawal to supply the growing city of Amman. The site is large, covering about 21 000 m², and about 1.5 m deep, and is thought to have been first occupied about 20 000 years ago, with continuous or intermittent occupation continuing for several thousand years. As well as the hut structures the site has yielded an underfloor burial, numerous ground stone tools, bone artifacts, and marine shell beads. Three hut structures (Structures 1, 2, and 3) examined at this site have been attributed to the earliest Epipalaeolithic. These have yielded a range of tools, bones and shells, which appear to have been left on the ground then covered over by a new floor.

Location map of Kharaneh IV and the location of other Epipaleolithic sites in the Azraq Basin. Ramsey et al. (2018).

Ramsey et al. took sediment samples from several locations from  Structure 1 at Kharaneh IV and analysed them for phytolths. These were then identified under a light microscope. They found that a high density of Grass and Sedge phytoliths in areas of the site associated with both the floor of the hut and its superstructure, though the floor deposits had a higher density of Sedges, which they suggest is an indication that the floor of the hut was covered by Sedge material, either as a loose covering or as woven mats. Such a floor covering would make sense in the presumed cool climate of the Late Pleistocene, and is known to have been used by more recent hunter-gatherer groups living in similar climates, such as the Nunamiut people of northern Alaska. It would also help to explain the high number of cultural objects (tools, beads etc.) found at the site, as these are more easily lost where there is a loose floor covering.

Phytolith microfossils from Kharaneh IV, Structure 1. (a) Sedge cones; (b) Grass husk; (c) stacked keystone bulliforms (cf. Reed leaves); (d) Sedge cones; and (e) Sedge cones (side view). Ramsey et al. (2018).

The sediments associated with the superstructure of the hut were found to contain a high proportion of phytoliths derived from Reeds (Phragmites sp.), which Ramsey et al. take as an indication that reeds were used in the construction of the hut. They suggest that, rather than resembling a brush hut from Southern Africa or the Australian interior, the hut may have more closely resembled the structures made by the peoples of the Great Basin of North America (most of Nevada, Utah and Oregon, plus parts of California, Idaho and Wyoming), in which a structure was made from erect wooden poles, then pulled together at the top with a series of woven Willow rings, leaving a smoke hole at the top. This basic superstructure could then be covered with a range of secondary materials, such as bundles of Grasses tied directly onto the structure (thatch) or a variety of woven mats (often several layers of matting, with a waterproof outer layer and insulating layers inside).

 Northern Paiute cattail house. Photograph by Mary Freeman, Stillwater, Nevada, ca. 1900. Libraries Special Collections/University of Nevada, Reno in Ramsey et al. (2018).

See also...

http://sciencythoughts.blogspot.com/2018/08/bread-from-pleistocene-of-jordan.htmlhttp://sciencythoughts.blogspot.com/2017/11/determining-origin-of-yabroud.html
http://sciencythoughts.blogspot.com/2017/04/gobekli-tepe-does-ancient-anatolian.htmlhttp://sciencythoughts.blogspot.com/2015/08/evidence-of-cereal-cultivation-by-sea.html
http://sciencythoughts.blogspot.com/2014/06/does-catalhoyuk-mural-depict-volcanic.htmlhttp://sciencythoughts.blogspot.com/2012/02/20-000-year-old-stone-huts-from.html
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Thursday, 24 May 2018

Determining the diet of Miocene Gomphotheriid Proboscidean using phytoliphs preserved in dental calculus.

The first Proboscideans (Elephants), appeared in the Palaeocene, and like other herbivorous Mammals of this time are presumed to have been browsers (leaf and fruit eaters). Modern Elephants, in contrast, are primarily grazers (grass eaters). Grasses first appeared in the Cretaceous, but extensive grasslands did not become a distinct ecosystem until the Miocene around 23 million years ago. Early Proboscidians had low-crowned teeth, with few lophids (ridges), consistent with a browsing diet, while modern Elephants have high-crowned teeth with numerous lophids, which offers some protection agianst the abbrasive nature of Grasses. The switch to Grasses as a food possibly occured in the Gomphotheres, which appeared in the Middle Miocene and are thought to have been ancestral to True Elephants. The earliest members of the group were trilophadont (had three ridges on their teeth), while later forms, particularly those thought to be ancestral to True Elephants, were tetralophadont (had four ridges), suggestive of a switch towards a more Grass-based diet.

In a paper published in the journal Scientific Reports on 16 May 2018, Yan Wu of the Key Laboratory of Vertebrate Evolution and Human Origins at the Institute of Vertebrate Paleontology and Paleoanthropology of the Chinese Academy of Sciences, and the Center for Excellence in Life and Paleoenvironment, Tao Deng, also of the Key Laboratory of Vertebrate Evolution and Human Origins, and the Center for Excellence in Life and Paleoenvironmentm, and of the Center for Excellence in Tibetan Plateau Earth Sciences, Yaowu Hu and Jiao Ma, also of the Key Laboratory of Vertebrate Evolution and Human Origins, and of the Department of Archaeology and Anthropology at the University of Chinese Academy of Sciences, Xinying Zhou, again of the Key Laboratory of Vertebrate Evolution and Human Origins, and the Center for Excellence in Life and Paleoenvironment, Limi Mao of the Key Laboratory of Economic Stratigraphy and Palaeogeography, at the Nanjing Institute of Geology and Palaeontology, Hanwen Zhang of the School of Earth Sciences at the University of Bristol, and of the Earth Sciences Department at the Natural History Museum, Jie Ye, again of the Key Laboratory of Vertebrate Evolution and Human Origins, and Shi-Qi Wang, once again of the Key Laboratory of Vertebrate Evolution and Human Origins, and the Center for Excellence in Life and Paleoenvironmentm, and of the Center for Excellence in Tibetan Plateau Earth Sciences, examine the diets of two species of Middle Miocene trilophadont Gomphotheriid Proboscideans, by examining phytoliths (silica fragments produced by plants) preserved in dental calculus of six specimens from the Miocene Halamagai Formation in the northern Junggar Basin of Xinjiang Province, China.

The Middle Miocene deposits of the Junggar Basin produce a diverse range of Gomphotheriid specimens, accompanied by floral remains indicative of a largely forested environment. Late Miocene strata from the same area, in contrast, have a much less diverse Gomphotheriid fauna, dominated by a few tetralophadont forms, and a more arid, Grass-dominated environment, suggesting that this area may have played an important role in the switch between browsing and grazing behaviour in early Proboscideans.

Wu et al. examined calculus from four specimens of Gomphotherium connexum and two specimens of Gomphotherium steinheimense. Gomphotherium steinheimense is thought to have been closely related to the early tetralophadont Gomphotheriid Tetralophodon longirostris, while Gomphotherium connexum is a more distant relative.

Geography, geology, and phylogeny in relation to the study material. (A) The location of the study area (black star). The map was generated by GTOPO309 using Globalmapper (v10). (B) Stratigraphic column and polarity with palaeomagnetic age, also denoting the horizon of study material in the strata (in light yellow). (C) The 50% majority consensus tree from 29 maximum parsimonious trees showing the phylogenetic position of the Gomphotherium species and Tetralophodon longirostris, the number at each node representing the support value calculated by majority rules (percentages of supported MPTs in the total MPTs, which are always larger than 50%) and the orange frame indicating the sister-taxon relationship of Gomphotherium steinheimense and Tetralophodon longirostris. (D) Gomphotherium steinheimense, right m3. (E) Gomphotherium connexum, left M3. Wu et al. (2018).

Of the phytoliths obtained from the calculus of Gomphotherium connexum, between 40% and 50% were identified as having originated from Grasses, whereas between 28% and 34% could be identified as having come from broadleaved plants. This would at first seem to imply a diet with a high proportion of Grasses, but Grasses produce a far greater amount of phytoliths than broadleaved plants (hence their more abrasive nature), so this probably indicates a diet with a high proportion of broadleaved plants. In contrast about 85% of the phytolihs from the calculus of Gomphotherium steinheimense could be identified as having come from grasses, indicative of a much more Grass-based diet.

See also...


http://sciencythoughts.blogspot.co.uk/2017/11/image-of-elephant-human-conflict-wins.htmlhttp://sciencythoughts.blogspot.co.uk/2017/10/elephants-kill-four-rohingya-refugees.html
http://sciencythoughts.blogspot.co.uk/2017/02/elephas-cf-e-planifrons-new-elephant.htmlhttp://sciencythoughts.blogspot.co.uk/2017/02/loxodonta-cyclotis-african-forest.html
http://sciencythoughts.blogspot.co.uk/2016/03/partial-gomphothere-tooth-from-miocene.htmlhttp://sciencythoughts.blogspot.co.uk/2016/03/dating-pleistocene-stegodon-from-west.html
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