Beneath Still Waters - Multistage Aquatic
Exploitation of Euryale ferox (Salisb.) during
Naama Goren-Inbar1, Yoel Melamed2, Irit Zohar3 , Kumar
Akhilesh4 and Shanti Pappu4,5
Cite this as: Goren-Inbar, N., Melamed, Y., Zohar, I., Akhilesh, K. and Pappu, S. (2014). Beneath
Still Waters - Multistage Aquatic Exploitation of Euryale ferox (Salisb.) during the Acheulian.
'Human Exploitation of Aquatic Landscapes' special issue (ed. Ricardo Fernandes and John
Meadows), Internet Archaeology. doi:10.11141/ia.37.1
1. Institute of Archaeology, The Hebrew University of Jerusalem, Mt Scopus, Jerusalem 91905,
Israel. Email: email@example.com
2. The Mina and Everard Goodman Faculty of Life Sciences, Bar-Ilan University, Ramat-Gan
3. National Natural History Collections, Berman Building, Edmond J. Safra Campus, The Hebrew
University of Jerusalem, Givat Ram, Jerusalem 91904, Israel.
4. Sharma Centre for Heritage Education, 28, I Main Road, C.I.T. Colony, Mylapore, Chennai
600004, Tamil Nadu, India. Email:firstname.lastname@example.org / email@example.com
5. Department of Archaeology, Deccan College Postgraduate and Research Centre, Pune 411006,
Keywords: Acheulian; Gesher Benot Ya'aqov; India; Aquatic exploitation; E. ferox
This issue has been funded by the Graduate School "Human Development in Landscapes",
University of Kiel with additional funding from the Institute for Ecosystem Research, University of
Kiel and theCentre for Baltic and Scandinavian Archaeology, Schloss Gottorf.
© Author(s). Except where otherwise noted, content from this work may be used under the terms
of the Creative Commons Attribution 3.0 Unported licence, which permits unrestricted use,
distribution, and reproduction in any medium, provided that attribution to the author(s), the title
of the work, the Internet Archaeology journal and the relevant URL/DOI is given.
Remains of the highly nutritious aquatic plant Fox nut – Euryale ferox Salisb.
(Nymphaeaceae) – were found at the Acheulian site of Gesher Benot Ya'aqov,
Israel. Here, we present new evidence for complex cognitive strategies of
hominins as seen in their exploitation of E. ferox nuts. We draw on excavated
data and on parallels observed in traditional collecting and processing practices
from Bihar, India. We suggest that during the early Middle Pleistocene, hominins
implemented multistage procedures comprising underwater gathering and
subsequent processing (drying, roasting and popping) of E. ferox nuts.
Hierarchical processing strategies are observed in the Acheulian lithic reduction
sequences and butchering of game at this and other sites, but are poorly
understood as regards the exploitation of aquatic plant resources. We highlight
the ability of Acheulian hominins to resolve issues related to underwater
gathering of E. ferox nuts during the plant's life cycle and to adopt strategies to
enhance their nutritive value.
List of Figures
Figure 1: Fox nut (E. ferox) from Gesher Benot Ya'aqov Layer II-6 Level 1,
complete seed with its characteristic germination aperture and attachment scar
(hilum). (Image credit: authors)
Figure 2: Map showing: (A) General view of the location of the regions studied in
Israel and India; (B) Location of the Acheulian site of Gesher Benot Ya'aqov,
Israel (33°00'30''N, 35°37'30''E) and; (C) Location of water bodies in Madhubani
District, Bihar, India where traditional methods of gathering and processing E.
ferox are practiced. The water bodies studied are located within a radius of
10km south and east of the town of Madhubani (26°22' 0'' N, 86°5'0'' E).
(Image credit: authors)
Figure 3: E. ferox seed remains from GBY: (A) Fox nut (E. ferox, GBY Layer II-6
Level 1), seed coat fragments easily identified by their characteristic attachment
scar (hilum) close to the germination aperture (appears in the right fragment)
(SEM); (B) Fox nut (E. ferox, GBY Layer II-6 Level 1), seed coat fragments,
eight showing the convex outer side and three the concave inner side; (C) Fox
nut (E. ferox, GBY Layer III-7), complete and compressed seeds. (Image credit:
Figure 4: Flow charts highlighting the main events in the gathering and
processing of E. ferox (locally called Makhana) based on our ethnographic
observations in Madhubani District, Bihar and summarised from published
literature (Jha et al. 1991; 2003; Mishra et al. 2003; Mandal et al. 2010).
(Image credit: authors)
Figure 5: Procedures involved in the collection of E. ferox nuts, Madhubani
District, Bihar, India; (A) General view of men of the fishing community involved
in the gathering of E. ferox (Makhana) nuts; note one type of collecting basket in
the background; (B) Close-up of E. ferox, showing the flower and characteristic
leaves; (C) View of a diver manipulating a bamboo pole that aids in demarcating
areas selected for underwater gathering of E. ferox nuts; (D) Diver surfacing
after gathering E. ferox nuts using a special type of basket (Gaanja), with an
aluminum pot also used for collection; (E) Adolescent boy collecting stray nuts
that rise to the water surface from the pond bed during collection; (F) Divers
(underwater) manipulating a large basket (Auka) in which nuts are collected.
Note the two hands at the edge of the basket. (Image credit: authors)
Figure 6: Procedures involved in the collection of E. ferox nuts, Madhubani
District, Bihar, India: (A) Diver with the basket that is rotated within water to
cleanse the nuts; (B) Divers bring the basket to the shore to complete the
process of cleansing the nuts; (C) Close-up of the collected nuts; (D) View of
the E. ferox nuts with associated molluscs; (E) Trampling helps remove the pulp;
F) A mound of nuts piled up near the water body ready for transport to the
village. (Image credit: authors)
Figure 7: Procedures involved in processing of E. ferox nuts, Madhubani District,
Bihar, India: (A) E. feroxnuts are spread out to dry in the sun in the village; (B)
The nuts are sorted into differing size ranges using sieves of different
dimensions; (C) The nuts are roasted and stirred using bamboo sticks; (D)
Popping of theE. ferox nuts immediately after roasting; note the wooden anvil
and hammer; (E) View of the popped E. ferox nuts ready to be eaten. (Image
Figure 8: Hammer and anvil used for popping the dried and roasted E. ferox nuts
in traditional practices in Madhubani District, Bihar, India: (A) General view of a
wooden hammer (Thaapi) (photograph: Gabi Laron); (B) A wooden anvil
(Aphara); (C) Measurements of a typical hammer; (D) Close-up of an anvil used
for popping E. ferox nuts. (Image credit: authors)
Figure 9: Debris of E. ferox nuts after popping: (A) A handful of nuts (eight in
number) generally popped together (N=50 fragments varying from 1mm to 1cm,
the rest being <1mm). At the top right of the image, note a complete roasted
nut of the same size group (diameter=11.49mm) as well as the final
poppedMakhana; (B) Close-up of nut fragments showing inner concave side; (C)
One roasted Makhana nut (N=5 fragments) after popping. (Image credit:
Figure 10: Anvils at the site of Gesher Benot Ya'aqov: A) Pitted anvils; B) Thin
anvil. (Image credit: authors)
List of Tables
Table 1: Nuts and shell fragments of E. ferox and T. natans at GBY (updated
from previous work). Apart from layers/levels noted below, pitted stones
(without botanical remains) were also found in Layer V-4: N=1; Layer V-5: N=4;
Jordan Bank (Layers V-5 and V-6 excavated together under water): N=7+2
anvils; Layer II-6 Level 4b: N=26+2 anvils.
Table 2: Common aquatic taxa from the Upper Jordan Valley (Lake Hula)
(Danin 2004), the Acheulian site of GBY and Bihar (India) (Melamed 2003;
Melamed et al. 2011). Data on fish from Goren and Ortal 1999; Zohar and
Biton 2011; Montana et al. 2011. In Bihar, 35 species have a high commercial
value, among which Cyprinidae and Clariidae predominate, as well as
Balitoridae. The richness of native fish species here is very high (>260 species),
thus differing from that of Lake Hula, similarities being at the family level
(Cyprinidae predominate in both Bihar and GBY).
Table 3: Edible species reported from GBY, including details of parts that can be
Studies of the evolution of hominin cognitive abilities and the origins of
intelligence and language focus primarily on stone tool manufacture and on the
exploitation of medium-sized to large terrestrial mammals. Here, we examine
additional aspects of these cognitive abilities as reflected in a little-known
example of skilled behaviour patterns: the exploitation of aquatic flora and fauna
in the wetland habitats of paleo-Lake Hula. Although wetlands play an important
role in supplementing human diet and enhancing its nutritional balance
(Joordens et al. 2009; Wrangham et al. 2009; Cunnane and Steward 2010), few
studies have explored the nutritional and/or medicinal properties of wetlands
plants in the archaeological context (Stewart 1994; 2010; Colonese et al. 2011;
Cortés-Sánchez et al. 2011; Hardy and Moncel 2011; Verhaegen and
Figure 1: Fox nut (E. ferox) from Gesher Benot Ya'aqov Layer II-6 Level 1, complete seed with its characteristic
germination aperture and attachment scar (hilum). (Image credit: authors)
Nuts of the aquatic plant Euryale ferox Salisb. (Nymphaeaceae) (common
names: Fox nut, Gorgon nut, Prickly water lily and Makhana in Bihar, India)
(Figure 1) were identified in Early to Middle Pleistocene deposits (Marine Isotope
Stages 18–20) at the site of Gesher Benot Ya'aqov (GBY), Israel (Goren-Inbar et
al. 2000; Feibel 2004; Sharon et al. 2011) (Figure 2). Along with Trapa
natans (Water chestnut), they formed part of the botanically rich aquatic habitat
of paleo-Lake Hula, comprising over 24 species of water plants (Table 1). Both
species are currently extinct in the Levant (Melamed 2003; Melamed et
al. 2011). E. ferox and T. natans are floating annual aquatic plants that grow in
low-energy or still-water bodies generally around 1.5m deep, occurring within a
wetlands ecosystem that was exploited by the GBY Acheulian hominins
(Melamed 2003; Ashkenazi et al. 2005; Ashkenazi et al. 2009; Spiro et al. 2009;
Mienis and Ashkenazi 2011; Zohar and Biton 2011). The prickly nature of E.
ferox renders gathering and processing its nuts far more difficult in comparison
with those of T. natans, thus providing us with an opportunity to explore the
ways in which this species was exploited at GBY.
Here, we present novel evidence for advanced cognitive abilities of Acheulian
hominins at GBY as attested by their adoption of complex multistage procedures
for collecting and processing E. ferox nuts. E. ferox is widely prevalent in tropical
and subtropical regions in ecological contexts similar to those of the paleo-Lake
Hula environment. In many such places, it is collected and processed using
traditional methods by predominantly freshwater fishing communities (Jha et
al. 1991; 2003). The range of these strategies, particularly evident in the water
bodies of northern Bihar (Madhubani District, India) (Figure 2), is of immense
relevance when examining the archaeological context of E. ferox nut remains at
The Acheulian site of GBY, situated within the Benot Ya'akov Formation
(Belitzky 2002), is located on the shores of paleo-Lake Hula in the Upper Jordan
Valley, Dead Sea Rift (Goren-Inbar et al. 2000). This Early to Middle Pleistocene
sedimentary sequence documents an oscillating freshwater lake and represents
~100,000 years of hominin occupation (Marine Isotope Stages 18–20) beginning
earlier than 790,000 years ago (Feibel 2001; 2004). Studies of the 15 excavated
archaeological horizons indicate that Acheulian hominins repeatedly occupied
lake margins, produced stone tools, systematically butchered and exploited
animals, gathered plant food, and controlled fire (Goren-Inbar et
al. 1994; 2002a; 2002b; 2004; Rabinovichet al. 2008; Alperson-Afil et al. 2009).
Figure 2: Map showing: (A) General view of the location of the regions studied in Israel and India; (B) Location
of the Acheulian site of Gesher Benot Ya'aqov, Israel (33°00'30''N, 35°37'30''E) and; (C) Location of water
bodies in Madhubani District, Bihar, India where traditional methods of gathering and processing E. ferox are
practiced. The water bodies studied are located within a radius of 10km south and east of the town of
Madhubani (26°22' 0'' N, 86°5'0'' E). (Image credit: authors)
2. Euryale ferox (Salisb.)
Remains of Euryale ferox nuts (seeds in botanical terminology) at GBY were
identified by their characteristic features such as the prominent longitudinal
ridge (raphe), shape and location of the operculum and hilum, as well as the
structure of the seed surfaces (Figures 1, 3). These characteristics
distinguish Euryale feroxfrom related fossil genera such
as Palaeoeuryale and Pseudoeuryale (Miki 1960).
Figure 3: E. ferox seed remains from GBY: (A) Fox nut (E. ferox, GBY Layer II-6 Level 1), seed coat fragments
easily identified by their characteristic attachment scar (hilum) close to the germination aperture (appears in
the right fragment) (SEM); (B) Fox nut (E. ferox, GBY Layer II-6 Level 1), seed coat fragments, eight showing the
convex outer side and three the concave inner side; (C) Fox nut (E. ferox, GBY Layer III-7), complete and
compressed seeds. (Image credit: authors)
The prickly water-lily is an annual or perennial plant with long-petiole leaves
whose large rounded blades (normally up to 1.3 and occasionally 2.4m in
diameter) float on the water surface. The long petiole and veins that protrude
from the bottom of the blade are densely covered with sharp prickles. The
rhizome is sunk deep in the ground with the help of groups of thick and fleshy
roots. The plant develops approximately 15–20 spongy fruits, each of which
contains 30–40 nuts. When the fruit is ripe it dehisces and releases the nuts,
which are covered by a mucilaginous arillus (Jha et al. 1991).
The plant grows in shallow stagnant water generally 0.3–1.5m deep and at a
neutral pH. In the study region in Madhubani District, Bihar, water depths reach
a maximum depth of around 3.5m. Flowering occurs in April–May and the fruits
ripen and dehisce between June and August, when spherical nuts are released.
The nuts have a mucilaginous arillus that holds them above the water surface for
several days, after which they sink to the bottom of the water body. The plant
germinates in early winter and grows with surprising speed, the biomass
doubling each month from January to July. The maximal biomass found in a
pond in India was 1.7kg/m² fresh weight in July. Temperature has a profound
effect on the rate of biomass production (Jha et al. 1991).
As regards their nutritional value, E. ferox nuts contain 12.8% moisture, 9.7%
protein, 0.1% fat, 0.5% minerals, 76.9% carbohydrates, 0.9% phosphorus,
0.02% calcium and 1.4 mg/100g carotene. The calorific value is 362 kcal/100g
for raw E. ferox and 328 kcal/100g for popped nuts (see below for description of
the processing techniques of these nuts). Popped nuts are comparable with
staple food such as wheat and rice. The essential amino acid indices (EAAI) in
the raw and popped parts of edible E. ferox nuts are 93% and 89%,
respectively. These are higher than the values for rice (83%), wheat (65%),
Bengal grain (81.55%), soya bean (85.6%), amaranth (57.5%), human milk
(81.55%), cow's milk (88.8%), fish (89.2%) and mutton (87.24%) (Jha et
al. 1991; Jha and Barat 2003; Ghosh and Santra 2003). E. ferox nuts are
superior to dry fruits such as almonds, walnuts, coconuts and cashew nuts in
terms of sugar, protein, ascorbic acid and phenol content (Jha et al. 1991; Jha
and Barat 2003; Ghosh and Santra 2003).
E. ferox was present in Europe in the geological past, becoming extinct during
the Quaternary (Simpson1936; Miki 1960; Soboleweska 1970; Jha et al. 1991;
Ghosh and Santra 2003). Fossil nuts of this species have been reported from the
Pleistocene in Poland (Soboleweska 1970) and England (Gibbard et al. 1996),
and from the Oligocene in Scotland (Simpson 1936). Evidence of E. ferox is also
noted in Tertiary deposits in Kolkata, India (Jha et al. 1991; Ghosh and
Santra 2003). Nuts of several extinct fossil Euryale species, including E.
europaea Weber, E. lissa Reid and E. nodulosa Reid, were found in geological
layers in Europe and Japan (Miki 1960). At present, E. ferox is the only surviving
species of the genus, which is the only recent genus of the
3. Materials and Methods
The methodology for calculating the number of Euryale ferox and Trapa
natans nuts in the archaeological record at GBY was as follows:
E. ferox – calculation of nut numbers: The number of complete nuts (Table
1) was calculated for each excavated unit by estimating the number of
fragments that could be conjoined to form a single nut coat. When this estimate
suggested the presence of more than one nut, the number was rounded up to
two, and so forth. However, some units yielded fragments including
characteristic structures such as the attachment scar (hilum) or germination
aperture of the nuts. In such cases, their number was compared to the
estimated value derived from the conjoined fragments and the higher value was
considered to represent the minimum number of complete nuts in the
T. natans – calculation of nut numbers: The nuts of T. natans (Table 1)
consist of a single nut enclosed within a bony calyx. The calyx bears four lateral
spines, a typical depression at the base and an aperture surrounded by an
extended rim at the tip. Most of the T. natans remains from GBY are fragments
of the nut wall, while others are spines, apertures or bases. Hence, the number
of nuts in each layer/level was calculated by estimating the number of fragments
that could be conjoined to form a single nut wall. In addition, for layers/levels
that also contained spines, apertures and bases, each structure was counted
separately and the number of spines was divided by four. In such cases, as
for E. ferox, their number was compared to the estimated value derived from the
conjoined fragments and the higher value was considered to represent the
minimum number of complete nuts in the assemblage.
Table 1: Nuts and shell fragments of E. ferox and T. natans at GBY (updated from previous work). Apart from layers/levels
noted below, pitted stones (without botanical remains) were also found in Layer V-4: N=1; Layer V-5: N=4; Jordan Bank
(Layers V-5 and V-6 excavated together under water): N=7+2 anvils; Layer II-6 Level 4b: N=26+2 anvils
4. The Context of E. ferox Remains at GBY
The identification of complete and fragmentary remains of E. ferox within
waterlogged archaeological horizons at GBY necessitated study of the site's
taphonomy, with particular attention to the degree to which hominins were the
agents responsible for the accumulation, spatial patterning and physical
condition of the remains.
Examination of site formation processes at GBY indicates that the archaeological
horizons were sealed rapidly, demonstrating a high integrity of preservation and
an excellent context for studies of spatial patterning (Alperson-Afil et al. 2009).
Observations from several archaeological horizons at the site may be
summarised as follows:
a. Intact embryos and faecal pellets of two extinct freshwater
molluscs, Viviparus apameae galileae and Bellamya sp. (Ashkenazi et
al. 2009) were present, together with packstone of the molluscs (Sharon
and Goren-Inbar 1999; Goren-Inbar and Sharon 2006; Rabinovich et
al. 2012), in a single layer at GBY in association with thousands of stone
artefacts and fossil mammal bones. Trampling and other post-depositional
agents would not have left the embryos and faecal pellets intact.
b. The record of organic material comprises wood, bark, seeds and fruits
(Goren-Inbar et al. 2002b; Melamed 2003), signifying minimal exposure
to atmospheric conditions and hence minimal presence of bacterial activity
that would have destroyed the anatomical structure and resulted in
c. The preservation of medium-sized and large mammal bones at the site is
excellent and hominin-induced markings (cut marks, percussion marks
and hack marks) are discernible on them (Rabinovich et al. 2008; 2012).
Experimental studies of faunal remains support these observations
(Gaudzinski-Windheuser et al. 2010; Rabinovich et al. 2012). Further
evidence includes the presence of conjoinable bones in some levels and
layers, indicating that taphonomically the bones underwent minimal
reorganisation (Goren-Inbar et al. 1994).
d. Fish remains at the site demonstrated differential preservation, being
deposited either as a natural death assemblage (Zohar and Biton 2011) or
in association with remains of hominin activities (Alperson-Afil et
e. The presence and spatial clustering of flint, basalt and limestone
microartefacts is a clear indication that taphonomic processes at GBY had
a minimal effect on archaeological horizons. The unique record of the
burned flint component (microartefacts) viewed as 'phantom hearths'
further confirms the excellent preservation of archaeological and
associated materials/features. In these cases, spatial patterning reflects
the time of abandonment. The lack of sedimentary obstacles, linear
deposition or winnowing, and the spatial association of finds (Alperson-Afil
and Goren-Inbar 2010), emphasise the absence of discernible taphonomic
Table 1 presents the occurrence and frequencies of the two types of water nuts
at GBY. Both E. ferox and T. natans nuts are present in both archaeological
horizons and geological layers, the latter being devoid of lithic artefacts. The
marked difference between archaeological and geological layers in quantities
of E. ferox and T. natans arises primarily from different sampling strategies.
Geological layers were minimally sampled along the walls of the trenches
(Trenches II–VI) as compared to the extensive studies conducted on
archaeological horizons. Within geological layers, there are differences in the
occurrences of the two types of nuts. In Layer III-7, both nuts are well
represented with E. ferox predominating. In other layers, T. natans is the
dominant seed, with its highest occurrence being in Layer II-9, a gray mud
sediment characteristically deposited under the highest water column and which
represents the deepest part of the paleo-lake. Sedimentologically, a group of
geological layers (Layers II-9, II-10, II-11, III-7) with a high occurrence of T.
natans represent coquinas. The absence of lithics in these layers does not
necessarily mean that they are archaeologically sterile. Although this may
appear to be a contradiction, our experience at the site leads us to suggest that
layers that are apparently archaeologically sterile may bear artefacts further
along the strike, beyond the boundaries of excavated areas. In some cases (e.g.
Layer II-12 in Goren-Inbar et al. 2002b, 81), an isolated microartefact indicated
the presence of an as yet unknown and unexcavated archaeological horizon.
Thus, the presence of seeds in archaeologically sterile layers may be explained
in two ways; as a natural occurrence or as a result of human behaviour. In the
absence of further excavations, neither possibility can be confirmed.
Remains of E. ferox are extremely abundant in the rich archaeological horizons
(Table 1, rows marked by *). They occur in varying frequencies in these layers,
as well as within the archaeological complex of Layer II-6. Hominin activities
varied significantly between the 15 rich archaeological horizons, as reflected by
differences in the presence and frequencies of finds (stone artefacts, fossil bones
and organic remains). Here, we propose the hypothesis that the spatial
patterning, physical condition and concentration of E. ferox nut fragments at the
site is the result of intentional hominin gathering and processing rather than of
natural factors. The key evidence for this is the co-occurrence of nuts and their
fragments with phantom hearths and pitted stones (Goren-Inbar et al. 2002a),
which occur in 10 of the 15 archaeological strata containing E. ferox remains
(Figure 3). One possible use of the pitted stones was as 'anvils' for popping nuts.
On the basis of ethnographic parallels discussed below, we believe that the
association of many E. ferox fragments with pitted stones and phantom hearths
is an indication of intentional roasting followed by popping to extract the nut
from its hard shell and exploit its maximum nutritive value (Jha and
Barat 2003). A new type of anvil, a component of the percussive tool
assemblage, was recently identified at GBY (Goren-Inbar et al. in prep). These
passive (dormant) percussive tools (de Beaune 2000) consist of thin natural
basalt slabs with two flat surfaces that are sometimes broken, probably as a
result of use. This is the first report of such thin anvils from an Acheulian site.
They are characterised by the presence of pits and other abrasive damage marks
on one or both flat surfaces and sometimes on the thin edges. Such anvils are
not found at the lake's edge but were selected for their particular morphology
and specific characteristics (e.g., hardness and non-vesicular nature) and
brought from exposures of basalt flows that are currently unknown in the vicinity
of the site. The total number of these anvils at GBY is 36, 26 of which are pitted.
Along with pitted stones (the latter primarily of basalt but also of limestone),
these anvils occur diachronically throughout the cultural sequence of the site
(Table 1). The same is true for pitted stones, although their availability was most
probably more extensive since their morphology is more varied (Goren-Inbar et
al.2002a). Together, these percussive tools form the bulk of the tools that we
propose were used to pop theE. ferox seeds and crack other types of nuts.
Figure 4: Flow charts highlighting the main events in the gathering and processing of E. ferox (locally called Makhana) based on our ethnographic observations in
Madhubani District, Bihar and summarised from published literature (Jha et al. 1991; 2003; Mishra et al.2003; Mandal et al. 2010). (Image credit: authors)
5. Traditional Modes of Exploitation of E.
ferox in India
Here, we draw on ethnographic analogies citing traditional methods of E.
ferox exploitation in the water bodies of northern Bihar, India, where planned
sequential procedures and decision-making strategies are employed by local
communities in collecting and processing E. ferox nuts (Jha et al. 1991; 2003;
Jha and Barat 2003; Ahmad and Singh 2003; Goswami 2003; Khan and
Halim 2003; Mishra et al. 2003; Singh 2003; Jain et al. 2010; Jain et al. 2011;
Mandal et al. 2010; Singh and Singh 2011). The procedures adopted in Bihar
imply an excellent knowledge of the environment and seasonality in relation to
the plants' life cycle (Jha et al. 1991) (Figure 4). Important to note here is the
fact that the prickles/spines on mature fruits make them very difficult to harvest
with bare hands. Once the mature fruits burst, the seeds float near the leaves
and then sink to the base of the pond, from where they are collected. Thus, a
structured sequential process has been devised for gathering and processing the
nuts, which has been discussed in the literature (Jha et al. 1991; 2003; Ahmad
and Singh 2003; Mishra et al. 2003; Singh 2003; Mandal et al. 2010) and
documented as part of this study (see Figures 5–8).
Based on these sources of information, we note that gathering is carried out by
adult males assisted by a few adolescent boys, with a division of activities that is
related to age and/or experience. The process can be summarised as follows: 1)
the equipment required (bamboo collection baskets of various types) is
organised; 2) bamboo poles fixed to the base of the pond serve as guides to
demarcate spaces selected for underwater gathering of the nuts and are shifted
as collection proceeds across the water body; 3) adults repeatedly dive
underwater to collect nuts that have sunk to the pond bed, at the base of the
plant; 4) the nuts are scooped into bamboo baskets (sieves); 5) in larger ponds
the nuts are scooped into a large cane basket and given a preliminary cleaning
underwater by repeated rotation; 6) an adolescent (inexperienced in diving)
floats on the water surface with the aid of pitchers or jerry cans and employs a
sieve to collect stray nuts that float to the surface; 7) the nuts brought to the
shore are cleaned by trampling to remove roots, plant matter and associated
molluscs (Jha et al. 1991; 2003; Ahmad and Singh 2003; Mishra et al. 2003;
Singh 2003; Mandal et al. 2010). Children actively participate in gathering
molluscs, crabs and other plants that are associated with the E. ferox roots and
are washed up on the shore during the gathering and cleaning procedure.
Figure 5: Procedures involved in the collection of E. ferox nuts, Madhubani District, Bihar, India; (A) General
view of men of the fishing community involved in the gathering of E. ferox (Makhana) nuts; note one type of
collecting basket in the background; (B) Close-up of E. ferox, showing the flower and characteristic leaves; (C)
View of a diver manipulating a bamboo pole that aids in demarcating areas selected for underwater gathering
of E. ferox nuts; (D) Diver surfacing after gathering E. ferox nuts using a special type of basket (Gaanja), with an
aluminum pot also used for collection; (E) Adolescent boy collecting stray nuts that rise to the water surface
from the pond bed during collection; (F) Divers (underwater) manipulating a large basket (Auka) in which nuts
are collected. Note the two hands at the edge of the basket. (Image credit: authors)
Figure 6: Procedures involved in the collection of E. ferox nuts, Madhubani District, Bihar, India: (A) Diver with
the basket that is rotated within water to cleanse the nuts; (B) Divers bring the basket to the shore to
complete the process of cleansing the nuts; (C) Close-up of the collected nuts; (D) View of the E. ferox nuts
with associated molluscs; (E) Trampling helps remove the pulp; F) A mound of nuts piled up near the water
body ready for transport to the village. (Image credit: authors)
Figure 7: Procedures involved in processing of E. ferox nuts, Madhubani District, Bihar, India: (A) E. ferox nuts
are spread out to dry in the sun in the village; (B) The nuts are sorted into differing size ranges using sieves of
different dimensions; (C) The nuts are roasted and stirred using bamboo sticks; (D) Popping of the E. ferox nuts
immediately after roasting; note the wooden anvil and hammer; (E) View of the popped E. ferox nuts ready to
be eaten. (Image credit: authors)
Figure 8: Hammer and anvil used for popping the dried and roasted E. ferox nuts in traditional practices in
Madhubani District, Bihar, India: (A) General view of a wooden hammer (Thaapi) (photograph: Gabi Laron); (B)
A wooden anvil (Aphara); (C) Measurements of a typical hammer; (D) Close-up of an anvil used for popping E.
ferox nuts. (Image credit: authors)
Figure 9: Debris of E. ferox nuts after popping: (A) A handful of nuts (eight in number) generally popped
together (N=50 fragments varying from 1mm to 1cm, the rest being <1mm). At the top right of the image, note
a complete roasted nut of the same size group (diameter=11.49mm) as well as the final popped Makhana; (B)
Close-up of nut fragments showing inner concave side; (C) One roasted Makhana nut (N=5 fragments) after
popping. (Image credit: authors)
Processing the nuts in the village comprises size sorting the nuts using a set of
sieves, several stages of sun-drying and roasting on a wood fire and lastly
popping, which results in the final nutritious product (Jhaet al. 1991; 2003;
Figures 7–9). The procedure of popping E. ferox seeds is widely used in Bihar
and involves drying followed by at least two cycles of roasting the nuts. The nuts
are then placed on a wooden anvil (around 60–70cm in diameter) and struck
with a wooden mallet-like hammer (around 17–27cm in diameter) (Figures 7–8).
These tools are purpose-built and are carved from the heartwood of Dalbergia
sisoo, Acacia lenticularis and Shorea robusta (Jha et al. 2003) (Figures 7–8).
Popping is a process whereby superheated vapour is created within the
conditioned nut by heating the moisture contained within, following which the
pressure is suddenly released, resulting in expansion of the volume of the
kernel. Popping the E. ferox seeds by traditional methods increases the
concentration of micronutrients per unit weight.
A distinct gender-based division of labour is observed. Activities within the
pond/lake are male dominated, while those involving drying, cleaning, roasting
and popping the seeds are largely the preserve of women, aided by children and
It has been mentioned that the prickles/spines on the outer surface of mature
fruits makes them difficult to harvest with bare hands, a problem resolved in the
study region by collection after the fruits burst and sink to the bed of the pond.
In Bangladesh, however, this species is harvested using boats and long-handled
curved knives in order to cut the pedicle underwater (Khan and Halim 2003) and
the pulpy aril and seeds are eaten, while the endosperm of the seeds is
consumed raw or roasted (Khan and Halim 2003). In parts of north-east India,
such as Assam, the edible seeds are eaten raw or in rare cases processed further
(Goswami 2003, 34). In Manipur, the leaf petioles and seeds are eaten raw or
boiled (Singh 2003, 10–11). Elsewhere in this state, the tender leaves, seed aril
and fruit skin are also consumed after removal of the prickles by fire or
otherwise, as is the mature leaf petiole after removal of the spines (Singh 2003,
11). It is also noted that the seeds are eaten raw after roasting in the sand and
dehusking (as quoted in Singh 2003, 11). In several regions here, ripe seeds
may also be eaten raw (Jain et al. 2010, 64; Jain et al. 2011) or used in a range
of traditional food preparations for dietary or medicinal purposes. Others (Singh
and Singh2011) note that the immature fruits are consumed after boiling while
the ripe ones are eaten fresh. In all cases, attention is paid to seasonality in
harvesting parts of the plant (Jain et al. 2010, 66).
6. Significance in Terms of E.
ferox Exploitation at GBY
The Indian record of E. ferox seed consumption indicates that there are several
ways to harvest and process E. ferox nuts. Clearly, the major difficulty to be
overcome is the presence of prickles/spines that make processing the fruit
difficult. Further difficulties involve collection of seeds after the fruit bursts. The
Bihar method described here overcomes both of these problems through
adoption of underwater collection procedures. Key points emerging from
investigation of traditional methods from Bihar in the context of theE.
ferox remains from GBY are summarised as follows: a) gathering of E.
ferox seeds takes place after they ripen and sink to the bed of the water body;
b) gathering by diving is a necessity, as the plants grow in still waters and seeds
are not washed to the edges of the water body; c) the work necessitates
observation of the lifecycle of the plant and of the prime time for gathering
seeds; d) drying and popping seeds was done at a distance from the water body,
where fire and dry land facilitated later stages of processing; e) roasting and
popping are both procedures requiring the technology of fire and that of anvils
and hammers; and f) a well-established division of labour was associated with
each stage of gathering and processing.
Different methods of E. ferox consumption are cited here, showing that the
seeds are a highly valuable component in the diet of wetland communities. It is
clear from literature and our own data that common processing methods of E.
ferox seeds in India are those that include the use of fire. Clearly, the prickly
nature of the plant parts is an important trait that local communities have to
consider and overcome. We believe that the traditional ethnobotanical
procedures involved in the gathering and multistage processing of E. ferox in
Bihar are instructive parallels for the interpretation of the GBY archaeological
data. Thus, the archaeological record of GBY, which includes the use of fire and
the presence of pitted stones, anvils and hammerstones in association with E.
ferox seeds, strongly supports the use of analogies with traditional modes of
gathering and processing, such as that practiced by communities in Bihar.
Studies of the GBY archaeological record provides information on the co-
occurrence of a range of finds that may be compared with the ethnographic
data. 1) In each of the archaeologically rich horizons there were spatial
concentrations of burned flint microartefacts. Analysis of these concentrations
suggests the presence of phantom hearths, the earliest evidence for the control
and continual use of fire in western Eurasia (Alperson-Afil and Goren-
Inbar 2010). High-resolution data from excavations enables estimation of the
size of these hearths, which were around 0.49m long and 0.35m wide (Alperson-
Afil and Goren-Inbar 2010, 74, table 4.1). 2) Pitted stones and hammerstones,
as well as the newly identified thin basalt anvils, were also found in each of
these horizons (Figure 10). 3) In all archaeological horizons, remains of T.
natans and E. ferox were discovered as well. The pristine taphonomic context of
the archaeological horizons at GBY, along with the significant patterns of
association noted between various find categories discussed above, provide a
background for our discussions of the spatial patterning of past activities. Spatial
analysis of these associations and analyses of Layer II-6 Levels 2 and 6 provide
further insight into the proximity of hearths and pitted stones (on both blanks
and blocks) (Table 1; Alperson-Afil and Goren-Inbar 2010, 91, figs 4.8, 4.9;
Alperson-Afil et al. 2009, 1678, fig. 2). This correlation of nuts, phantom hearths
and pitted stones at GBY leads us to suggest that some key aspects of the
methods of collecting and processing noted in Bihar, which includes roasting and
subsequent popping of the seeds, may be of greater relevance for the GBY data
than those described from elsewhere in India. Greater precision in spatial
associations between the nuts and other features in the vicinity of the paleo-lake
is not possible owing to the light weight of the seeds. However, common aquatic
taxa in the Upper Jordan Valley (Lake Hula), the Acheulian site of GBY and Bihar
(India) (Table 2) reflect the extent of ecological similarity, despite their great
biogeographical distance. The habitat and surrounding environment of paleo-
Lake Hula was a rich and diverse Mediterranean one, as evidenced by the
identification of an array of 60 edible taxa recorded at GBY (Melamed 2003,
table 3), as well as a wealth of fish, crustaceans, birds and mammals.
Figure 10: Anvils at the site of Gesher Benot Ya'aqov: A) Pitted anvils; B) Thin anvil. (Image credit: authors)
E. ferox grows in water bodies with water depths ranging from around 0.3m to
1.5m, up to a maximum of around 2.5m (Jha et al. 1991; Mishra et al. 2003;
Mandal et al. 2010) or up to around 3.5m as noted in the study region. Although
Acheulian hominins may have consumed seeds raw, this would have entailed
considerable effort in harsh conditions owing to the prickly nature of the plants.
With the technology enabling them to process nuts using fire, anvils and
percussive tools, hominins could avoid the difficulties posed by exploitation of
The fluctuating water level of paleo-Lake Hula would have been an obstacle to
adopting simpler methods for gathering nuts, in view of the plant's life cycle and
the water depths and geochemistry required for its growth and survival. A
desiccation scenario of fluctuating lake levels would have resulted in the death of
plants, unable to regenerate as germination occurs under water. Exposure to
atmospheric conditions would have resulted in the complete decomposition of
the macrobotanical remains found at the site. The entire issue of organic
preservation is based on anaerobic conditions (and hence inappropriate
conditions for bacteria responsible for the decomposition of organic material).
Irrespective of the depth of the water, hominins would have had to collect nuts
from beneath the lake surface, entailing some amount of time spent under
It is important to note that we do not suggest that Acheulian hominins followed
modes of collecting or processing that were identical to those practiced today,
particularly in the case of elements dictated by modern economic conditions (use
of bamboo poles to demarcate underwater areas for collecting nuts, sieves for
sorting nuts for sale or even gender-based division of labour).
Such cognitive procedural abilities of planning and performance in aquatic
habitats, particularly when combined with exploitation of fish (Zohar and
Biton 2011) have not previously been reported for Acheulian hominins.
Ethnographic analogies demonstrate that exploitation of E. ferox nuts is
performed by communities of fishermen in water bodies that are also used for
fishing (Jha et al. 1991). The most abundant fish species currently exploited in
habitats associated with E. ferox include three families of air-breathing fish:
Cyprinidae (carps), Clariidae (catfish) and Bagridae (catfish) (Table 2). At GBY,
remains of Cyprinidae and Clariidae were recovered, predominantly Cyprinidae
(mainly the large Barbus sp. and Barbus longiceps). Interestingly, the cyprinids
remains were recovered in association with living floors excavated in Area B
(Table 2) (Alperson-Afil et al. 2009; Zohar and Biton 2011). The archaeological
association between E. ferox nuts, large quantities of cyprinid remains and other
cultural activities documented at GBY presents novel evidence for intensive
exploitation of the aquatic fauna and flora of paleo-Lake Hula.
Table 2: Common aquatic taxa from the Upper Jordan Valley (Lake Hula) (Danin 2004), the Acheulian site of GBY and Bihar
(India) (Melamed 2003; Melamed et al. 2011). Data on fish from Goren and Ortal 1999; Zohar and Biton 2011; Montana et
al. 2011. In Bihar, 35 species have a high commercial value, among which Cyprinidae and Clariidae predominate, as well as
Balitoridae. The richness of native fish species here is very high (>260 species), thus differing from that of Lake Hula,
similarities being at the family level (Cyprinidae predominate in both Bihar and GBY)
* = endemic species of fish
Euryale ferox Salisb.
Lemna minor L.
Potamogeton pectinatus L.
P. pectinatus L.
P. nodosus Poir.
Potamogeton lucens L.
P. lucens L.
P. perfoliatus L.
Trapa natans L.
Trapa natans L.
Trapa natans L.
Utricularia australis R.Br.
Aldrovanda vesiculosa L.
Utricularia gibba L.
lissneri (Tortonese 1952)
lissneri (Tortonese 1952)
Cyprinids - 19
Garra rufa (Heckel 1843)
Garra rufa (Heckel 1843)
hulensis (Goren et al.
Mirogrex hulensis (Goren et al.
kervillei (Pellegrin 1911)
kervillei (Pellegrin 1911)
(Banarescu and Nalbant
panthera (Heckel 1843)
Nun galilaeus* (Günter
Clarias gariepinus (Burchell
Aphanius mento (Heckel
galilaeus (Artedi 1757)
Sarotherodon galilaeus (Artedi
Tilapia zillii (Gervais
Tilapia zillii (Gervais 1848)
intermedia* (Steinitz and
intermedia* (Steinitz and Ben-
S= number of species
The advanced and sophisticated cognitive abilities described above are
supported by a series of additional observations drawn from various
multidisciplinary studies of the GBY Acheulian record. These include aspects of
planning and communication as derived from stone tool production sequences
(Sharon et al.2011), spatial cognition of the landscape and intra-site spatial
organisation (Goren-Inbar and Sharon 2006; Rabinovich et al. 2006;
Rabinovich et al. 2008), procedural cognition, technical and procedural know-
how and specialisation (Madsen and Goren-Inbar 2004; Alperson-Afil et
al. 2009; Goren-Inbar 2011; Goren-Inbar et al. 2011; Rabinovich and
Biton 2011), as well as social cognition (Goren-Inbar et al. 2002b; Goren-
Inbar2011). These cognitive abilities are expressed in the multiphase process of
realisation of the plan for achieving a particular goal. This is seen especially in
the chaîne opératoire of basalt bifaces (handaxes and cleavers), documenting
cognitive abilities in the structure of the long-term processes involved in biface
In addition to the above cultural observations based on lithic assemblages and
their reduction sequences, there is also evidence derived from the faunal record
at the site. This is characterised by both richness and diversity of species,
contributing substantially to the reconstruction of hominin knowledge of the
environment in exploitation of both terrestrial wildlife (Rabinovich et al. 2012;
Rabinovich and Biton 2011), such as modern-like processing of Dama sp.
(Rabinovich et al. 2008), and aquatic resources such as turtles (Hartman 2004)
and fish (Alperson-Afil et al. 2009; Zohar and Biton 2011).
Table 3: Edible species reported from GBY, including details of parts that can be consumed
Ceratophyllum demersum L.
Euryale ferox Salisb.
Nuphar lutea (L.) Sm.
leaf, petiole, rhizome
Lemna minor L.
Potamogeton pectinatus L.
leaf and stem
Trapa natans L.
Utricularia australis R. Br.
Palaeobotanical evidence contributes to our understanding of the multiple facets
of the environmental knowledge of Acheulian hominins and their ability to
structure modes of exploitation of diverse resources. At GBY, this exploitation of
multiple resources includes that of seven fruit-bearing species with edible nuts
belonging to both extant Mediterranean vegetation and locally extinct species
(Goren-Inbar et al. 2002a; Melamed 2003; Melamed et al. 2011). Two of the
locally extinct species (in the Mediterranean biome) are edible aquatic nuts that
flourished in paleo-Lake Hula and were found in quantities at the site: E.
ferox and T. natans (Goren-Inbar et al. 2000; Goren-Inbar et al. 2002a;
Rabinovich et al. 2012) (Table 3). The pitted stones and anvils mentioned above
support the view that these remains were components of the paleo-diet (Goren-
Inbar et al. 2002a). These were found in close spatial proximity with burned flint
microartefacts, the latter indicating the location of Acheulian hearths (Goren-
Inbar et al. 2002a; Goren-Inbar et al. 2004). Evidence of fire is seen throughout
the sequence of Acheulian occupation at GBY (around 50,000 years at the site;
Sharon et al. 2011) and attests to use and control of this component of culture.
Continual fire making (Goren-Inbar et al. 2004) and the transmission of
particular modes of technological tool production indicate evolved
communication within the group, interpreted as language (Alperson-Afil and
Goren-Inbar2010; Sharon et al. 2011). Clearly, long-term memory was already
a component of the evolutionary realm of GBY hominins. The exploitation of E.
ferox at GBY, supplemented by data from ethnographic parallels, indicates that
Acheulian hominins implemented complex strategies to extract maximum
nutritive value from plant species, despite the opportunity of consuming them
fresh. It also suggests delayed gratification implied by the time gap between
appearance and collecting of the nuts. We do not claim that Acheulian hominin
cognitive abilities were similar to those of modern humans, but do suggest that
some aspects of complex cognition possibly overlapped in these hominins.
Ethnographic analogies, when considered with archaeological evidence of nuts,
pitted anvils and charred organic material, among other features, point to the
possibility of a complex sequence of exploitation of an aquatic nut that included
gathering by diving, underwater processing, drying, roasting and possibly
popping. This process adds to a plethora of evidence of Acheulian hominin
activities and diverse associated cognitive abilities, all of which emerge from the
analyses of early Middle Pleistocene Acheulian finds from the Levantine Corridor.
We thank Professor Vidyanath Jha, who shared with us his expert knowledge
of E. ferox. Mr Ranjay Kumar Jha organised the field visit to Madhubani District,
Bihar, India, and along with Mr Bhogi Mukhiya helped us to document
procedures. We are grateful to G. Hivroni for the graphic design (Figures 1 and
4), Gabi Laron for photography of Figure 10a, and to Dr Y. Langsham for the
SEM photography. K.A. and S.P. thank the Sharma Centre for Heritage Education
for logistic aid. Botanical remains are stored at the Mina and Everard Goodman
Faculty of Life Sciences, Bar-Ilan University, Ramat-Gan 5290000, Israel. At the
faculty, the macrobotanical remains are stored at the National Natural History
Collection (seeds). We are grateful to the two reviewers: Professor Lyn Wadley
and an anonymous reviewer who helped us improve the article.
The study of the fish remains was supported by the Israel Science Foundation
(Grant No. 300/06) to the Center of Excellence Project Title: 'The Effect of
Climate Change on the Environment and Hominins of the Upper Jordan Valley
between ca. 800Ka and 700Ka ago as a Basis for Prediction of Future Scenarios'
and the Irene Levi Sala CARE Archeological Foundation.
Ahmad, S.H. and Singh, A.K. 2003 'Scope of integration of Makhana with fish
culture' in R.K. Mishra, V. Jha and P.V. Dehadrai (eds) Makhana, New Delhi:
Directorate of Information and Publications of Agriculture, Indian Council of
Agricultural Research. 85–97.
Alperson-Afil, N., Sharon, G., Zohar, I., Biton, R., Melamed, Y., Kislev, M.E.,
Ashkenazi, S., Rabinovich, R., Werker, E., Hartman, G. and Goren-Inbar, N.
2009 'Spatial organization of hominin activities at Gesher Benot Ya'aqov,
Israel', Science 326, 1677–80. http://dx.doi.org/10.1126/science.1180695
Alperson-Afil, N. and Goren-Inbar, N. 2010 The Acheulian Site of Gesher Benot
Ya'aqov: Ancient Flames and Controlled Use of Fire, Dordrecht:
Ashkenazi, S., Klass, K., Mienis, H.K., Spiro, B. and Abel, R. 2009 'Fossil
embryos and adult Viviparidae from the early–middle Pleistocene of Gesher
Benot Ya'aqov, Israel: ecology, longevity and fecundity', Lathaia 43, 116–
Ashkenazi, S., Motro, U., Goren-Inbar, N., Bitton, R., and Rabinovich, R. 2005
'New morphometric parameters for assessment of body size and population
structure in freshwater fossil crab assemblage from the Pleistocene site of
Gesher Benot Ya'aqov (GBY), Israel', Journal of Archaeological Science 32, 675–
Beaune, S.A. de 2000 Pour Une Archéologie du Geste, Paris: CNRS Editions.
Belitzky, S. 2002 'The structure and morphotectonics of the Gesher Benot
Ya'aqov area, northern Dead Sea rift, Israel', Quaternary Research 58, 372–80.
Colonese, A.C., Mannino, M. A., Mayer, D.E.B.-Y., Fa, D.A., Finlayson, J.C.,
Lubell, D. and Stiner, M.C. 2011 'Marine mollusc exploitation in Mediterranean
prehistory: an overview', Quaternary International 239, 86–
Cortés-Sánchez, M., Morales-Muñiz, A., Simón-Vallejo, M.D., Lozano-Francisco,
M.., Vera-Peláez, J.L., Finlayson, C., Rodríguez-Vidal, J., Delgado-Huertas, A.,
Jiménez-Espejo, F.J., Martínez-Ruiz, F., Martínez-Aguirre, M.A., Pascual-
Granged, M.A.J., Bergadá-Zapata, M.M., Gibaja-Bao, J.F., Riquelme-Cantal, J.A.,
López-Sáez, J.A., Rodrigo-Gámiz, M., Sakai, S., Sugisaki, S., Finlayson, G., Fa,
D.A. and Bicho, N.F. 2011 'Earliest known use of marine resources by
Neanderthals', PLoSONE 6(9): e24026.
Cunnane, S. and Stewart, K.M. (eds) 2010 Human Brain Evolution, New York:
John Wiley and Son.
Danin, A. 2004 Distribution Atlas of Plants in the Flora Palaestina Area,
Jerusalem: Israel Academy of Sciences and Humanities.
Feibel, S.C. 2001 'Archaeological sediments in lake margin environments' in J.K.
Stein and W.R. Farrand (eds) Sediments in Archaeological Contexts, Salt Lake
City: University of Utah Press. 127–48.
Feibel, C.S. 2004 'Quaternary lake margins of the Levant Rift Valley' in N.
Goren-Inbar and J.D. Speth (eds) Human Paleoecology in the Levantine
Corridor, Oxford: Oxbow Books. 21–36.
Gaudzinski-Windheuser, S., Kindler, L., Rabinovich, R. and Goren-Inbar, N. 2010
'Testing heterogeneity in faunal assemblages from archaeological sites. Tumbling
and trampling experiments at the early middle Pleistocene site of Gesher Benot
Ya'aqov (Israel)', Journal of Archaeological Science 37, 3170–90.
Ghosh, S.K and Santra, S.C. 2003 'Past and present distributional records of
Makhana and future prospects of its cultivation in West Bengal' in R.K. Mishra, V.
Jha and P.V. Dehadrai (eds) Makhana, New Delhi: Directorate of Information and
Publications of Agriculture, Indian Council of Agricultural Research. 3–7.
Gibbard, P.L., Aalto, M.M., Coope, G.R., Currant, A.P., McGlade, J.M., Peglar,
S.M., Preece, R.C., Turner, C., Whiteman, C.A. and Wrayton, R.C. 1996 'Early
Middle Pleistocene fossiliferous sediments in the Kesgrave Formation at
Broomfield, Essex, England' in C. Turner (ed) The Early Middle Pleistocene in
Europe, Rotterdam: Balkema. 83–119.
Goren-Inbar, N. 2011 'Culture and cognition in the Acheulian industry – a case
study from Gesher Benot Ya'aqov', Philosophical Transactions of the Royal
Society of London Series B 366, 1038–49.
Goren, M. and Ortal, R. 1999 'Biogeography, diversity and conservation of the
inland water fish communities in Israel', Biological Conservation 89(1), 1–
Goren-Inbar, N. and Sharon, G. 2006 'Invisible handaxes and visible Acheulian
biface technology at Gesher Benot Ya'aqov, Israel' in N. Goren-Inbar and G.
Sharon (eds) Axe Age: Acheulian Tool-making from Quarry to Discard, London:
Goren-Inbar, N., Lister, A., Werker, E. and Chech, M. 1994 'A butchered
elephant skull and associated artifacts from the Acheulian site of Gesher Benot
Ya'aqov, Israel', Paléorient 20(1), 99–112.
Goren-Inbar, N., Feibel, C.S., Verosub, K.L., Melamed, Y., Kislev, M.E.,
Tchernov, E. and Saragusti, I. 2000 'Pleistocene milestones on the Out-of-Africa
corridor at Gesher Benot Ya'aqov, Israel', Science 289, 944–74.
Goren-Inbar, N., Sharon, G., Melamed, Y. and Kislev, M. 2002a 'Nuts, nut
cracking, and pitted stones at Gesher Benot Ya'aqov, Israel', Proceedings of the
National Academy of Sciences USA 99(4), 2455–60.
Goren-Inbar, N., Werker, E. and Feibel, C.S. 2002b The Acheulian Site of Gesher
Benot Ya'aqov: The Wood Assemblage, Oxford: Oxbow Books.
Goren-Inbar, N., Alperson, N., Kislev, M.E., Simchoni, O., Melamed, Y., Ben-
Nun, A. and Werker, E. 2004 'Evidence of hominin control of fire at Gesher Benot
Ya'aqov, Israel', Science 304, 725–27.
Goren-Inbar, N., Grosman, L. and Sharon, G. 2011 'The technology and
significance of the Acheulian giant cores of Gesher Benot Ya'aqov,
Israel', Journal of Archaeological Science 38, 1901–17.
Goren-Inbar, N., Sharon, G., Herzlinger, G. and Alperson-Afil, N. (in prep) 'A
newly identified type of Acheulian basalt anvil from Gesher Benot Ya'aqov,
Goswami, M.M. 2003 'Ecology of Euryale ferox Salisb. in flood plain wetlands of
Assam' in R.K. Mishra, V. Jha and P.V. Dehadrai (eds) Makhana, New Delhi:
Directorate of Information and Publications of Agriculture, Indian Council of
Agricultural Research. 29–35.
Hardy, B.L. and Moncel, M.-H. 2011 'Neanderthal use of fish, mammals, birds,
starchy plants and wood 125–250,000 years ago', PLosONE 6(8).
Hartman, G. 2004 'Long-term continuity of a freshwater turtle (Mauremys
caspica rivulata) population in the Northern Jordan Valley and its
paleoenvironmental implications' in N. Goren-Inbar and J.D. Speth (eds)Human
Paleoecology in the Levantine Corridor, Oxford: Oxbow Books. 61–74.
Jain, A., Singh, H.B. and Kanjilal, P.B. 2010 'Economics of Foxnut (Euryale
ferox Salisb.) cultivation: a case study from Manipur in North eastern
India', Indian Journal of Natural Products and Resources 1(1), 63-67.
Jain, A, Sundriyal, M., Roshnibala, S., Kotoky, R., Kanjilal, P.B., Singh, H.B. and
Sundriyal, R.C. 2011 'Dietary use and conservation concern of edible wetland
plants at Indo-Burma hotspot: a case study from northeast India', Journal of
Ethnobiology and Ethnomedicine 7, 29. http://dx.doi.org/10.1186/1746-4269-7-
Jha, V. and Barat, G.K. 2003 'Nutritional and medicinal properties of Euryale
ferox Salisb.' in R.K. Mishra, V. Jha and P.V. Dehadrai (eds) Makhana, New
Delhi: Directorate of Information and Publications of Agriculture, Indian Council
of Agricultural Research. 230–38.
Jha, V., Kargupta, A.N., Dutta, R.N., Jha, U.N., Mishra, R.K. and Saraswati, K.C.
1991 'Utilization and conservation of Euryale ferox Salisbury in Mithila (North
Bihar), India', Aquatic Botany 39, 295–314.http://dx.doi.org/10.1016/0304-
Jha, V., Verma, A.M. and Jha, A.K. 2003 'Indigenous contrivances utilized
in Makhana cultivation in north and north-eastern India' in R.K. Mishra, V. Jha
and P.V. Dehadrai (eds) Makhana, New Delhi: Directorate of Information and
Publications of Agriculture, Indian Council of Agricultural Research. 241–48.
Joordens, J.C.A., Wesselingh, F.B., de Vos, J., Vonhof, H.B. and Kroon, D. 2009
'Relevance of aquatic environments for hominins: a case study from Trinil (Java,
Indonesia)', Human Evolution 57, 656–
Khan, M.S. and Halim, M. 2003 'Makhana, Euryale ferox Salisb. in Bangladesh' in
R.K. Mishra, V. Jha and P.V. Dehadrai (eds) Makhana, New Delhi: Directorate of
Information and Publications of Agriculture, Indian Council of Agricultural
Madsen, B. and Goren-Inbar, N. 2004 'Acheulian giant core technology and
beyond: an archaeological and experimental case study', Eurasian Prehistory 2,
Mandal, R.N., Saha G.S. and Sarangi, N. 2010 'Harvest and processing of
Makhana (Euryale ferox Salisb.) – a unique assemblage of traditional
knowledge', Indian Journal of Traditional Knowledge 9(4), 684–88.
Melamed, Y. 2003 Reconstruction of the Hula Valley vegetation and the hominid
vegetarian diet by the Lower Palaeolithic botanical remains from Gesher Benot
Ya'aqov, Unpublished PhD thesis, Bar-Ilan University (in Hebrew).
Melamed, Y., Kislev, M.E., Weiss, U. and Simchoni, O. 2011 'Extinction of water
plants in the Hula Valley: evidence for climate change', Journal of Human
Evolution 60, 320-27.http://dx.doi.org/10.1016/j.jhevol.2010.07.025
Mienis, H.K. and Ashkenazi, S. 2011 'Lentic basommatophora molluscs and
hygrophilous land snails as indicators of habitat and climate in the early-middle
Pleistocene (0.78 Ma) site of Gesher Benot Ya'aqov (GBY)', Journal of Human
Evolution 60, 328–40. http://dx.doi.org/10.1016/j.jhevol.2010.03.009
Miki, S. 1960 'Nymphaeaceae remains in Japan, with new fossil genus
Eoeuryale', Journal of the Institute of Polytechechnics 11(D), 63–78.
Mishra, R.K., Jha, V. and Dehadrai, P.V. (eds) 2003 Makhana, New Delhi:
Directorate of Information and Publications of Agriculture, Indian Council of
Montana, G., Choudhary, S.K., Dey, S. and Winemiller, K.O. 2011 'Compositional
trends of fisheries in the River Ganges, India', Fisheries Management and
Ecology 18, 282–96. http://dx.doi.org/10.1111/j.1365-2400.2010.00782.x
Rabinovich, R. and Biton, R. 2011 'The early-middle Pleistocene faunal
assemblages of Gesher Benot Ya'aqov – taphonomy and
paleoenvironment', Journal of Human Evolution 60, 357–
Rabinovich, R., S. Gaudzinski-Windheuser, and N. Goren-Inbar 2006 Site
Formation Processes - The Role of Hominin and Natural Agents in the Formation
of Striations and Cut Marks on Bones at the Acheulian site of Gesher Benot
Ya'aqov, Israel, Final Report, Jerusalem: German-Israeli Foundation for Scientific
Research and Development.
Rabinovich, R., Gaudzinski, S. and Goren-Inbar, N. 2008 'Systematic butchering
of Fallow deer (Dama) at the early middle Pleistocene Acheulian site of Gesher
Benot Ya'aqov, (Israel)', Journal of Human Evolution 54, 134–49.
Rabinovich, R., Gaudzinski-Windheuser, S., Kindler, L. and Goren-Inbar, N.
2012 The Acheulian Site of Gesher Benot Ya'aqov: Mammalian Taphonomy – The
Assemblages of Layers V-5 and V-6, Dordrecht:
Sharon, G. and Goren-Inbar, N. 1999 'Soft percussor use at the Gesher Benot
Ya'aqov Acheulian site?', Journal of the Israel Prehistoric Society 28, 55–79.
Sharon, G., Alperson-Afil, N. and Goren-Inbar, N. 2011 'Cultural conservatism
against variability in the continual Acheulian sequence of Gesher Benot Ya'aqov,
Israel', Journal of Human Evolution 60, 387–
Simpson, J.B. 1936 'Fossil pollen in Scottish Tertiary coal', Proceedings of the
Royal Society of Edinburgh 56, 90–108.
Singh, P.K. 2003 'Distribution and uses of Makhana in Manipur' in R.K. Mishra, V.
Jha and P.V. Dehadrai (eds) Makhana, New Delhi: Directorate of Information and
Publications of Agriculture, Indian Council of Agricultural Research. 8–13.
Singh, A.V. and Singh, P.K. 2011 'Nutritional, ethnotherapeutics and
socioeconomic relevance of Euryale Ferox Salisb. in Manipur, India', Life
Sciences Leaflets 22, 1104–15.
Soboleweska, U.S. 1970 'Euryale ferox Salisb. in the Pleistocene of Poland', Acta
Palaeobotanica 11, 13–20.
Spiro, B., Ashkenazi, S., Mienis, H.K., Melamed, Y., Feibel, C., Estacion, A.D. and
Starinsky, A. 2009 'Climate variability in the Upper Jordan Valley around 0.78
Ma, inferences from time-series stable isotopes of Viviparidae, supported by
mollusc and plant palaeoecology', Palaeogeography, Palaeoclimatology,
Palaeoecology 282, 32–44. http://dx.doi.org/10.1016/j.palaeo.2009.08.005
Stewart, K.M. 1994 'Early hominid utilisation of fish resources and implications
for seasonality and behaviour', Journal of Human Evolution 27, 229–
Stewart, L.M. 2010 'The case for exploitation of wetlands environments and
floods by pre-Sapiens hominins' in S.C. Cunnane and K.M. Stewart (eds) Human
Brain Evolution. The Influence of Freshwater and Marine Food Resources,
Hoboken, New Jersey: John Wiley and Sons. 137–
Verhaegen, M. and Munro, S. 2011 'Pachyosteosclerosis suggests archaic Homo
frequently collected sessile littoral foods', HOMO Journal of Comparative Human
Biology 62, 237–47.http://dx.doi.org/10.1016/j.jchb.2011.06.002
Wrangham, R., Cheney, D., Seyfarth, R. and Sarmiento, E. 2009 'Shallow-water
habitats as sources of fallback foods for hominins', American Journal of Physical
Anthropology 140, 630–42.http://dx.doi.org/10.1002/ajpa.21122
Zohar, I. and Biton, R. 2011 'Land, lake, and fish: investigation of fish remains
from Gesher Benot Ya'aqov (paleo-lake Hula)', Journal of Human Evolution 60,