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Tracing Domestication and Cultivation of Bananas from Phytoliths: An update from Papua New Guinea

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There is now good evidence from current banana distributions and genetic analysis that Papua New Guinea and nearby regions have played a key role in the domestication of edible Eumusa and Australimusa bananas. Strong support for this also comes from phytoliths in the rchaeobotanical record. Seeds have diagnostic phytoliths which can be used to discriminate between the two main sections of edible bananas, the giant banana, Musa ingens, and Ensete. Therefore, the presence of seed phytoliths and their subsequent disappearance from archaeological assemblages can be used to trace processes of domestication leading to parthenocarpy and sterility. Following loss of viable seeds, banana presence can still be documented from phytolith morphotypes from other plant parts, particularly the volcaniform morphotypes from leaves. Nevertheless, according to several pioneer studies, these are more difficult to differentiate unless they occur in regions where certain species or varieties of bananas are not endemic. This paper reviews results from morphometric and morphotypic analyses of Musaceae phytoliths and briefly introduces the 'New Guinea Banana Project' which builds upon previous analyses. The morphometric database, combined with a comprehensive set of images, facilitates banana phytolith identification and is another step forward in solving the issues surrounding banana dispersal, cultivation and domestication, especially in the Pacific/New Guinea region.
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Correspondence
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
Ethnobotany Research & Applications 7:247-270 (2009)
Carol J. Lentfer, School of Social Science, University of Queen-
sland, Queensland, AUSTRALIA.
clentfer20@hotmail.com
(bell), seeds and owers can also be eaten; the leaves
are used for cooking, wrapping, serving food and for shel-
ter from the sun and rain; bers from the stem and peti-
oles are used for making string, rope and other cordage
for weaving; the sap is used for dye; and, the seeds are
used as beads or money (Burkill 1935, Lentfer 2003a,
b). Given this multitude of uses, even in their wild state
and prior to the development of eshy, seedless (or near-
ly seedless) fruit (Lentfer 2003a), it is likely that banana
plants would have been recognized as a prized resource
and exploited in the past in ways similar to the present.
Indeed, this is the picture that is gradually emerging from
a host of archaeobotanical, biogeographical, biomolecu-
lar and genetic evidence (e.g., Carreel et al. 2002, De
Langue & de Maret 1999, Denham et al. 2003, Kennedy
2008, Mbida et al. 2001, Perrier et al. 2009, Vrydaghs
et al. 2009). Complex origins and multiple dispersals for
banana cultivars are indicated, but perhaps more strik-
ing is the signicant role that the New Guinea region has
played in the development of the domesticated banana.
After a brief review of the evidence for the origins and
spread of cultivated bananas, this paper outlines a cur-
rent research project which expands current banana phy-
Tracing Domestication and
Cultivation of Bananas
from Phytoliths: An update
from Papua New Guinea
Carol J. Lentfer
Research
Abstract
There is now good evidence from current banana distribu-
tions and genetic analysis that Papua New Guinea and
nearby regions have played a key role in the domestica-
tion of edible Eumusa and Australimusa bananas. Strong
support for this also comes from phytoliths in the archaeo-
botanical record. Seeds have diagnostic phytoliths which
can be used to discriminate between the two main sec-
tions of edible bananas, the giant banana, Musa ingens,
and Ensete. Therefore, the presence of seed phytoliths
and their subsequent disappearance from archaeological
assemblages can be used to trace processes of domes-
tication leading to parthenocarpy and sterility. Following
loss of viable seeds, banana presence can still be docu-
mented from phytolith morphotypes from other plant parts,
particularly the volcaniform morphotypes from leaves.
Nevertheless, according to several pioneer studies, these
are more difcult to differentiate unless they occur in re-
gions where certain species or varieties of bananas are
not endemic.
This paper reviews results from morphometric and mor-
photypic analyses of Musaceae phytoliths and briey in-
troduces the ‘New Guinea Banana Project’ which builds
upon previous analyses. The morphometric database,
combined with a comprehensive set of images, facilitates
banana phytolith identication and is another step forward
in solving the issues surrounding banana dispersal, cul-
tivation and domestication, especially in the Pacic/New
Guinea region.
Introduction
Musa bananas (including plantains) constitute major food
staples in the tropics and in terms of gross value of food
production they are by far the most important world fruit
crop. Furthermore, the banana plant is valued for more
than just its fruit (Kennedy 2009): the stem, corm, bract
Published: July 30, 2009
Ethnobotany Research & Applications248
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tolith databases for further assessment of phytolith varia-
tion within and between wild and domesticated Musa ba-
nanas and Ensete in Papua New Guinea.
Origins and distributions of banana cultivars
There are several wild species of Musa bananas and
closely related Ensete species ranging from Africa (Ensete
only) to India, Southeast Asia and as far east as New
Guinea and the Solomon Islands (Ensete and Musa). Ad-
ditionally, there are hundreds of diploid, triploid and poly-
ploid cultivars derived from a few wild species from two
sections of bananas (Eumusa and Australimusa) that are
cultivated in tropical and subtropical regions worldwide
(Argent 1976, Arnaud & Horry 1997, Daniells et al. 2001,
De Langhe et al. 2009, Kennedy 2008, Lentfer 2003a,
Pollefeys et al. 2004, Sharrock 1990, Simmonds 1959,
1962, Valmayor 2001, Wong et al. 2001). Phytogeogra-
phy and genetic evidence shows that the domesticated
Australimusa Fe`i banana almost certainly originated in
the New Guinea region (Jarret et al. 1992, MacDaniels
1947, Simmonds 1959), but the origins of the more com-
monly known and widely marketed Eumusa cultivars have
been more difcult to determine. Based on the distribution
of diploid and triploid Eumusa bananas that contain an A
genome derived from either Musa acuminata Colla sub-
species banksii (F. Muell.) Simmonds or errans (Blanco)
R.V. Valmayor or both (see summary in Table 6, Kennedy
2008:85), it appears that the primary center for the do-
mestication of edible Eumusa section bananas was the
Philippines/New Guinea region.
Signicantly, a long history of banana manipulation by hu-
mans, particularly in the New Guinea region, is indicated
by the presence of the banksii A genome in AAB plan-
tains as far aeld as Africa and the Pacic. Coupled with
the proliferation of diploid AA cultivars in New Guinea, this
points to the likelihood of an early dispersal from the New
Guinea region in two directions, eastwards into the Pa-
cic region and westwards through Island Southeast Asia,
Malaysia and across to Africa (Kennedy 2008:85-86). The
large number of diploid AA cultivars with both banksii and
errans genomes, as well as the absence of the errans ge-
nome in the African AAB plantains but its presence togeth-
er with the banksii genome in the Pacic AAB Maia Maoli
plantains are interesting and imply complex species and
subspecies interactions within the Philippines/New Guin-
ea regions. This would have involved human diffusion of
bananas, probably concurrent with the earliest transfer of
the banksii A genome westwards into Island Southeast
Asia, and then to mainland Asia and eventually Africa, and
also subsequently over an extended period as people ex-
panded eastwards into the Pacic (Kennedy 2008, Perrier
et al. 2009).
Archaeobotanical records for bananas
The archaeobotanical record for bananas is very sparse
and is mostly derived from microfossil evidence, partic-
ularly phytoliths (reviewed in Denham & Donohue 2009
and Donohue & Denham 2009). Evidence from starch
shows good promise of adding to this (Lentfer 2009), but
currently there is only one conrmed record with a positive
identication of banana starch associated with an archae-
ological deposit from Santa Cruz in the Solomon Islands
(Crowther 2009). The only other record comes from the
Yuku rock shelter site in the Western Highlands of Papua
New Guinea, but this has not been conrmed as banana
(Horrocks et al. 2008). Most phytolith records to date have
been based on the identication of distinctive volcaniform
morphotypes from Musaceae leaves. However, identica-
tion beyond the family level has not been successfully at-
tempted in most studies because of the difculties in dis-
criminating between volcaniform morphotypes from differ-
ent Musaceae genera, sections and species.
In his pioneering work at Kuk in the Western Highlands
of Papua New Guinea, Wilson (1985) used morphometric
analyses to discriminate between three sections of banan-
as and he also identied some Musaceae morphotypes in
sediments dated to c. 10,000 cal B.P. as Australimusa.
However, these identications were problematic and in-
conclusive, partly due to the limited set of comparative ref-
erence material analyzed, but also due to the assumption
that Eumusa section bananas were introduced into New
Guinea from Southeast Asia during the mid-to-late Holo-
cene (e.g., Spriggs 1996). More recently, Lentfer (2003a)
found that the seeds from Australimusa, Eumusa, Ingenti-
musa and Ensete have diagnostic phytolith morphotypes
(Figure 1), and was able to conrm the presence of Musa
ingens Simmonds - the giant cold-tolerant banana be-
longing to Section Ingentimusa, Ensete glaucum (Roxb.)
Cheesman, and Eumusa section bananas from seed phy-
tolith morphotypes in a similar archaeological context at
the Kuk Swamp site (Denham et al. 2003, Lentfer 2003b).
The earliest records for Eumusa and Ingentimusa seed
phytoliths recovered from palaeochannel lls in this analy-
sis were dated at c. 10,000 cal B.P. Eumusa persisted to
the top of the archaeological sequence dated at c. 2500
cal B.P. The earliest date for Ensete seed phytoliths re-
covered from palaeosurface feature lls was c. 7000-6500
cal B.P. Similar to Eumusa, these persisted to the top of
the sequence. Volcaniform leaf phytoliths were also pres-
ent but were not identied to any particular section or spe-
cies.
In addition to the Kuk site, Lentfer also conrmed the
presence of Eumusa section bananas from diagnostic
seed phytoliths at the coastal Lapita site of SAC on Wa-
tom Island, East New Britain, Papua New Guinea (Lentfer
& Green 2004), as well as Ensete, Eumusa and Australi-
musa section bananas at the archaeological site of FIF/4
at the Yombon airstrip in South West New Britain (Lentfer
et al. 2008). As with Kuk, volcaniform leaf phytoliths were
present in both assemblages but no attempt was made to
identify them beyond the family level in the initial analy-
ses.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
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249
Figure 1. Diagnostic seed morphotypes of wild Musa bananas and Ensete from Papua New Guinea. A-B. Ensete
glaucum (Accession No. QH28807); C-D. Musa ingens, Section Ingentimusa (Accession No. WH1); E-F. Musa peekelii,
Section Australimusa (Accession No. WNB488).
C D
FE
BA
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Figure 1 cont. Diagnostic seed morphotypes of wild Musa bananas and Ensete from Papua New Guinea. G-H. Musa
maclayi, Section Australimusa (Accession No. MB6); I-J. Musa acuminata ssp.banksii, Section Eumusa (Accession No.
QH067962); K-L. Musa schizocarpa, Section Eumusa (Accession No. NB489).
H
G
I J
L
K
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
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251
Apart from phytolith studies in New Guinea there are few
other published accounts of banana identication beyond
the family level with the exception of two African studies
in Cameroon and Uganda. Volcaniform morphotypes re-
covered from refuse pits at an agricultural village site,
Nkang in southern Cameroon dating to c. 2500 cal B.P.
were identied to genus Musa (Mbida et al. 2001, 2004).
Further to the east, at Munsa, Uganda, both Musa and
Ensete phytoliths were identied from swamp sediment
cores (Lejju et al. 2006). The oldest dates for these were
c. 5200 cal B.P. (cf. Neumann & Hildebrand 2009). In con-
trast to New Guinea, no seed phytoliths were recorded
from either African site, probably because Musa bananas
were already seedless by the time they had been intro-
duced into Africa.
The question of cultivation
Banana plants in their natural state are light-demanding
pioneer species of tropical environments. Growing nat-
urally from seed and via suckers, bananas are adapted
to opportunistic colonization of mostly well-drained open
sites such as forest margins, forest gaps resulting from
tree fall, and scree slopes associated with landslides and
erosion (e.g., Argent 1976, MacDaniels 1947). Human
selection, which eventually led to female sterility, loss of
seeds and parthenocarpy, has produced hundreds of dif-
ferent land races and hybrids of Eumusa, Australimusa
and Eumusa x Australimusa section bananas occurring in
the Indo-Pacic region and Africa (see De Langhe et al.
2009) and implies a long history of somatic mutation and
human manipulation involving cultivation.
Nevertheless, given the sparse record of prehistoric ba-
nana distribution, tracking evidence for cultivation and dis-
persal of cultivars is difcult. The record is derived most-
ly from two broad categories of evidence: ecological and
geographical. This is mostly reliant on archaeobotanical
nds showing presence of bananas: outside their natural
range; in contexts with archaeological, sedimentary and
ecofactual features indicative of cultivation; or, in associa-
tion with other known domesticates and associated spe-
cies, plants and/or animals. At Kuk, for instance, the alti-
tude is exceptionally high (>1560 metres above sea lev-
el) for wild Eumusa section bananas and Ensete to occur
naturally (see Argent 1976), but even if the early Holocene
environment was warmer than it is today, the presence
of stake holes, post holes and mounds, coupled with the
relatively high proportions of Musaceae phytoliths in the
phytolith assemblages, particularly following erosion and
burning episodes, are strongly supportive of human inu-
ences and cultivation at least by about 7000-6500 cal B.P.
(Denham et al. 2003).
Evidence for cultivation at the Watom site is equally strong.
Banana phytoliths are found in black, humic rich soils typi-
cal of gardens. They are in association with phytoliths and
macrobotanical remains from other cultivars including co-
conut, Canarium, Job’s tears, possibly sugar cane, phyto-
liths derived from pioneer tree species and grasses that
colonise gardens, and pig and chicken bones (Lentfer &
Green 2004). All other evidence for cultivation primarily
relies on bananas being outside their natural range of dis-
tribution. For instance, bananas are outside their natural
range in the Pacic east of the Solomon Islands. There-
fore, all records for bananas east of the Solomons are
indicative of human translocation and cultivation includ-
ing wild M. acuminata ssp. banksii found in Samoa (De
Langhe 2009, MacDaniels 1947) and the Musa found as-
sociated with Lapita deposits in Vanuatu (Horrocks et al.
in press). The same applies in Africa where only Ensete
species are indigenous. Musa banana cultivation can be
inferred at the Nkang site in Cameroon by 2500 cal B.P.
from the presence of Musa phytoliths (Mbida et al. 2001,
2004) and pending the accuracy of dating and the mor-
photypic discrimination between Musa and Ensete volca-
niform phytoliths (Neumann & Hildebrand 2009), it is pos-
sible that banana cultivation occurred in Uganda as early
as c. 5200 cal B.P. (Lejju et al. 2006).
Identifying banana cultivation in areas where wild banan-
as grow naturally is perhaps the most difcult, especially
in the absence of supportive archaeological and/or palae-
obotanical evidence. At the Yombon airstrip locale, south
West New Britain, Papua New Guinea, for instance, ba-
nanas appear in the early to middle Holocene. Howev-
er, although this coincides with a major burning episode,
there is no other evidence supportive of cultivation other
than the presence of a few potential cultivars including
Saccharum sp. and Coix lachryma-jobi L. In cases such
as this, cultivation could only be conrmed if it could be
proven that bananas were indeed seedless, and therefore
cultivars.
Identication of cultivars:
morphometric and morphotypic
analyses of phytoliths
Tracing the history of banana cultivation, domestication
and dispersal could be greatly facilitated if phytoliths can
be readily differentiated in archaeobotanical assemblag-
es. However, the variation of phytoliths within and be-
tween Musaceae taxa and within and between plant parts
(with the exception of banana seed phytoliths) has, until
recently, been poorly understood. To address this shortfall
and to expand previous morphotypic and morphometric
analyses undertaken by Wilson (1985) and Mbida et al.
(2001), phytolith researchers have commenced a series
of rigorous morphometric analyses of banana phytoliths to
determine their diagnostic value (e.g., Ball et al. 2006).
Lentfer (2003a) has undertaken preliminary studies to in-
vestigate the variation of phytoliths within and between
species and also within and between plant parts. The rst
set of exploratory analyses examined seventeen acces-
Ethnobotany Research & Applications252
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Figure 2. A. Sheet of polygonal and globular seed
phytoliths from Musa acuminata ssp. banksii. These
morphotypes have craters and were included in the
analysis. The plant material was obtained from the
Queensland Herbarium (Accession No. QH067962). B-C.
Examples of volcaniform and globular leaf morphotypes
from Musa maclayi (Accession No. NB487) examined in
the analysis.
Table 1. Banana accessions analysed by Lentfer (2003a,
b).
Section Wild species Accession code
(plant parts*)
Eumusa Musa acuminata
ssp. banksii
QH325354
(lf,sd,sk,mrb,ped);
QH541190 (sd,br)
Musa schizocarpa 489 (lf,sd,sk);
QH356650 (lf)
Australimusa Musa peekelii QH067966
(lf,sd); 488 (lf;sd/
fr); 489 (fr/sd);
QH067968 (sk)
Musa maclayi QH537000 (lf,br);
NB487 (lf,lfbs/st);
QH356648(mrb)
N/A Ensete glaucum QH28807 (sd,sk);
482 (lf,sk,sd);
QH356652 (lf)
Cultivars
Eumusa Musa acuminata QH438477 (lf)
Musa paradisiaca? QH4000037 (lf)
Australimusa TT(Fe`i) QH067969 (lf)
*lf=leaf; sd=seed; fr=fruit; sk=skin; br=bract;
mrb=mid
rib; st=stem; lfbs=leaf base; ped=peduncle Note Fe`i is
referred to as M. fei F. Muell. in gures.
B
A
sions consisting of Ensete and wild and cultivated Australi-
musa and Eumusa Section bananas (Table 1). Twenty-ve
phytoliths from a number of different plant parts includ-
ing leaf blades, leaf mid-ribs, leaf bases, fruit and seed,
skin, pseudostems, bracts and peduncles were analyzed
separately. Only phytoliths with craters (i.e., spherical to
sub-spherical to blocky morphotypes but not necessarily
volcaniform morphotypes, see Figure 2) were included in
the analysis. It did not include any of the clearly diagnos-
tic seed morphotypes referred to previously and shown in
Figure 1.
Analysis 1
a) Differentiation of phytolith seed morphotypes from
other plant parts: Long dimensions of phytolith bodies
and craters were measured (see Figure 1, Vrydaghs et
al., 2009) and ratios of body length to crater width were
calculated. Results of the analysis using pooled data
show that the ratio of mean body length to mean crater
width is signicantly different at α = 0.05, differentiating
between seed/fruit pulp morphotypes and morphotypes
from other plant parts (Figure 3). Notably, body length
and crater width scores by themselves were less helpful
in this regard.
b) Differentiation of Eumusa, Australimusa seed mor-
photypes and Ensete: Ensete glaucum has distinctive
C
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
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253
9
8
7
6
5
4
3
2
95% Cl crater dimension
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
3A
N-
Part
21
20
19
18
17
16
15
14
13
95% Cl long dimension
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
3B
N-
Part
9
8
7
6
5
4
3
2
95% Cl crater dimension
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
3A
N-
Part
A
B
21
20
19
18
17
16
15
14
13
95% Cl long dimension
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
3B
N-
Part
9
8
7
6
5
4
3
2
1
95% Cl R AT I O
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
N=
Part
3C
9
8
7
6
5
4
3
2
1
95% Cl R AT I O
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
N=
Part
3C
C
Figure 3. 95% condence intervals for mean crater widths A, mean long dimensions B and mean long dimension/
mean crater width ratios C of all phytoliths examined. Note that the leaf/stem sample consists of the base of leaf and
pseudostem samples and the seed/fruit samples consist of: A) the fruit pulp attached to seeds, and B) seeds. Fruit pulp
does not contain phytoliths and therefore phytoliths examined in the analysis are derived from seeds only. The ratio
plot (C) shows that seed and seed/fruit phytoliths have signicantly smaller craters compared to body length than leaf
phytoliths and can be discriminated at α = 0.05 regardless of species derivation.
Figure 4. 95% condence intervals for mean crater widths
of Musa spp. seed and Ensete glaucum leaf and fruit-skin
phytoliths based on full data set A and with outliers deleted
B (sd = seed; fr = fruit; lf = leaf; sk = skin). Musa acuminata
ssp. banksii is clearly differentiated at α = 0.05 when outliers
are removed. The Australimusa species M. peekelii is not
differentiated from Ensete glaucum.
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
A
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
21
20
19
18
17
16
15
14
13
95% Cl long dimension
332
leaf/bract
100
seed/fruit
25
leaf/stem
37
midrib
3B
N-
Part
B
22
16 22
625
7
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
Ensete sp.
(lf)
Ensete sp.
(sk)
M. peekelii
(sd)
4B
1
3
5
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
22
16 22
625
7
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
Ensete sp.
(lf)
Ensete sp.
(sk)
M. peekelii
(sd)
4B
1
3
5
B
Ethnobotany Research & Applications254
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Table 2. Categories and attributes used in analysis of volcaniform leaf phytoliths (attributes shown in Figure 6B are the
abbreviated forms shown in (…) and are equivalent terms from Madella et al. 2005).
Categories Attributes
Crater round [r] (orbicular), oval [o], square [sqt], irregular [irrt]
Morphology tabular [t], blocky [b], spherical [sph] (globose), platy [pl] (planar)
Base shape square [squb], rectangular [rb], quadrilateral [qb], triangular [tb], boat [bb] (oblong), round [cb]
(orbicular), other [ob]
Height short [fh] (h<1/3 length), medium [mh] (h=1/3 to 1/2 length), tall [th] (h≥1/2 length)
Texture psilate [stx], rough [rtx] verrucate, granulate [grx], dimpled [dtx]
Rim present [rp]/absent [ra], regular [regr]/irregular [irrr]
Ornamentation absent [no], short [sho], medium [mo], long [lo], lobed [lbo]
seed morphotypes (see Figure 1) that were not includ-
ed in this analysis. However, the leaf and fruit skin phy-
tolith morphotypes of Ensete have globular and polygo-
nal morphotypes similar to Musa seed morphotypes. An
analysis comparing mean body length and mean crater
width of M. acuminata ssp. banksii and Musa peekelii
Lauterb. seed morphotypes, and E. glaucum leaf and
fruit skin morphotypes showed that the width of craters
in M. acuminata ssp. banksii are signicantly smaller
than both M. peekelii and Ensete at α = 0.05 (Figure 4).
M. peekelii and E. glaucum could not be differentiated
according to crater width. However, mean body length
of M. peekelii was signicantly greater at α = 0.05 than
M. acuminata ssp. banksii and Ensete (Figure 5).
Figure 5. 95% condence intervals for mean long
dimensions of Musa spp. seed and Ensete leaf and fruit-
skin phytoliths based on full data set (sd = seed; fr = fruit;
lf = leaf; sk = skin). The Australimusa species, M. peekelii
is differentiated from Musa acuminata ssp. banksii and
Ensete glaucum.
Analysis 2
This analysis was based on morphotypic analysis of the
same set of leaf/bract volcaniform morphotypes used in
the rst analysis. Seven major categories of attributes
(Table 2) were examined. Attributes within each catego-
ry were given a score of 1 if present and 0 if absent and
statistically tested using principal components analysis.
Similar to the morphometric analysis, Ensete glaucum
morphotypes were clearly differentiated by body texture
and crater rim characteristics (Figure 6). Other taxa
could not be clearly differentiated.
Analysis 3
Additional morphometric analyses were undertaken to
determine if leaf phytolith morphotypes could be further
differentiated. Mean body length and mean crater width
of leaf/bract volcaniform morphotypes from different
taxa were compared (Figures 7 to 9). Crater width of
E. glaucum morphotypes was signicantly smaller at α
= 0.05 than all other Eumusa and Australimusa banan-
as with the exception of the wild Australimusa species,
Musa maclayi F. Muell. ex Mikl.-Maclay and the wild Eu-
musa species Musa schizocarpa Simmonds (Figure 7).
Body length was signicantly smaller than the two M.
acuminata cultivars (listed as M. acuminata and Musa
paradisiaca L.), M. maclayi and Fe`i (Figure 8). Crater
length and body length of morphotypes from other ba-
nanas were more similar and consequently these taxa
were found to be more difcult to differentiate (Figures 7
and 8). Crater width of the wild M. maclayi morphotypes
were signicantly different from the cultivated bananas,
M. acuminata and Fe`i and the other wild Australimusa
banana M. peekelii, but overlapped with the wild Eumu-
sa species (M. acuminata ssp. banksii and M. schizo-
carpa). Musa schizocarpa could only be differentiated
from the three cultivars (M. acuminata, M. paradisiaca
and Fe`i), and the wild M. acuminata ssp. banksii was
differentiated from only one of the Eumusa section cul-
tivars, M. acuminata, not M. paradisiaca. Body length
was a less helpful criterion for differentiating taxa than
crater width. Interestingly, the only signicant difference
at α = 0.05 was between Australimusa section banan-
22 16 22 25 6
40
30
20
10
0
N=
species
95% Cl long dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
Ensete sp.
(lf)
Ensete sp.
(sk)
M. peekelii
(sd)
5
22 16 22 25 6
40
30
20
10
0
N=
species
95% Cl long dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
Ensete sp.
(lf)
Ensete sp.
(sk)
M. peekelii
(sd)
5
22 16 22 25 6
40
30
20
10
0
N=
species
95% Cl long dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
Ensete sp.
(lf)
Ensete sp.
(sk)
M. peekelii
(sd)
5
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
8
6
4
2
0
N=
species
95% Cl crater dimension
M. banksii
(sd)
M. peekelii
(sd/fr)
E. glaucum
(lf)
E. glaucum
(sk)
M. peekelii
(sd)
4A
17
25
24
25 7
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
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255
Figure 6. Biplot of principal
components analysis of banana
leaf phytolith attribute data.
The sample plot A shows that
Ensete glaucum is differentiated
from other bananas. The main
attributes separating it from
other bananas are the irregular
rim and the rough texture
and to a lesser extent, short
ornamentation (SHO). See the
distribution of attributes in the
vector plot B. (see abbreviations
in Table 2)
2
3
1
0
-1
-2
-3
0 2-4 -2 864
PC 1
PC 2
6A
2
3
1
0
-1
-2
-3
0 2-4 -2 864
PC 1
PC 2
6A
Musa banksii F. Muell.
Musa schizocarpa Simmonds
Musa peekelii Lauterb.
Musa paradisiaca L.
Musa maclayi F . Muell. ex Mikl.-Maclay
Musa fei F. Muell.
Musa acuminata Colla
Ensete sp.
6Akey
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.8
0.2
-0.8 0.0-0.6 -0.2-0.4 0.2 0.4 0.6 0.8
RTX
IRRR
T
R
TH
SHO
RA
FH
GRX
QB
CB
RP
MH
LO
SQUB
MO
RB
TB
BB
DTX
OB
O
B
NO
IRRT
SPH
STX
REGR
PC 1
PC 2
SQT
Rough texture
Irregular rim
6B
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.8
0.2
-0.8 0.0-0.6 -0.2-0.4 0.2 0.4 0.6 0.8
RTX
IRRR
T
R
TH
SHO
RA
FH
GRX
QB
CB
RP
MH
LO
SQUB
MO
RB
TB
BB
DTX
OB
O
B
NO
IRRT
SPH
STX
REGR
PC 1
PC 2
SQT
Rough texture
Irregular rim
6B
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.8
0.2
-0.8 0.0-0.6 -0.2-0.4 0.2 0.4 0.6 0.8
RTX
IRRR
T
R
TH
SHO
RA
FH
GRX
QB
CB
RP
MH
LO
SQUB
MO
RB
TB
BB
DTX
OB
O
B
NO
IRRT
SPH
STX
REGR
PC 1
PC 2
SQT
Rough texture
Irregular rim
6B
A
the greatest variability and could not be differentiated
from any other taxon.
Lentfers ndings broadly concur with those of Ball et al.
(2006) and Vrydaghs et al. (2009), which have been con-
ned to the distinctive volcaniform leaf phytoliths from a
different set of Musaceae accessions including M. acumi-
nata, Musa balbisiana Colla and various cultivar groups.
A combination of morphometric (base length and crater
width) and morphotypic analyses (base shape, crater po-
sition and cone shape) (Ball et al. 2006:3) can help to dis-
criminate between certain taxa. Notably, wild diploid M.
balbisiana (BB) volcaniform morphotypes were found to
be signicantly larger than both wild and edible diploid
M. acuminata (AA) morphotypes (Ball et al. 2006:7), but
edible AA could not be differentiated from wild AA. Sub-
sequent studies analyzing AA, AAA, AAB and ABB have
found a very complex pattern of phytolith variation. Con-
tinuing analyses with additional samples are further inves-
tigating the variation in crater width, particularly the role of
banksii alleles in its expression (Vrydaghs et al. 2009).
Implications for future research
The presence of seeded bananas in archaeobotanical as-
semblages can be identied from seed phytolith morpho-
types. Distinctive morphotypes shown in Figure 1 are di-
agnostic at the section level. Other globular and polygonal
morphotypes can be differentiated from other plant parts
by body length/crater width ratios. Additionally, prelimi-
nary studies indicate that Eumusa bananas (M. acuminata
ssp. banksii) can be differentiated from Australimusa ba-
nanas (M. peekelii) on the basis of crater width and body
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.8
0.2
-0.8 0.0-0.6 -0.2-0.4 0.2 0.4 0.6 0.8
RTX
IRRR
T
R
TH
SHO
RA
FH
GRX
QB
CB
RP
MH
LO
SQUB
MO
RB
TB
BB
DTX
OB
O
B
NO
IRRT
SPH
STX
REGR
PC 1
PC 2
SQT
Rough texture
Irregular rim
6B
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.8
0.2
-0.8 0.0-0.6 -0.2-0.4 0.2 0.4 0.6 0.8
RTX
IRRR
T
R
TH
SHO
RA
FH
GRX
QB
CB
RP
MH
LO
SQUB
MO
RB
TB
BB
DTX
OB
O
B
NO
IRRT
SPH
STX
REGR
PC 1
PC 2
SQT
Rough texture
Irregular rim
6B
B
as. Body lengths of M. maclayi and Fe`i bananas were
signicantly greater than M. peekelii. Finally the ratio
of mean body length to crater width differentiated M.
maclayi from all other samples with the exception of
M. schizocarpa (Figure 9). Musa schizocarpa exhibited
Ethnobotany Research & Applications256
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Figure 7. 95% condence intervals
for mean crater widths of leaf
phytoliths differentiating Ensete
glaucum from all other bananas
except Musa maclayi and Musa
schizocarpa.
7
10
9
8
7
5
N=
species
7658
25
25
25
6
25 50 50
95% Cl crater dimension
7 labels
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
95% Cl crater dimension
7 labels
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
8
7658
25
25
25
25 50 50
22
20
18
16
10
N=
species
14
12
8titles
95% Cl long dimension
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
8titles
95% Cl long dimension
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
Figure 8. 95% condence intervals
for mean long dimensions of leaf
phytoliths differentiating Ensete
glaucum from four other bananas.
length. Although further compara-
tive studies are needed to include
a range of other species from ei-
ther section, current results sug-
gest the outlook is very promising
for tracking the complex history
of Musaceae in the archaeobo-
tanical record. Phytoliths can be
used to identify natural distribu-
tions of Musa and Ensete, differ-
entiate wild populations from fully
domesticated (seedless) popula-
tions and trace patterns of disper-
sal. However, based on this prem-
ise, mixed populations of wild and
cultivated bananas (a common
occurrence in Papua New Guin-
ea; Lentfer 2003b; Jean Kenne-
dy pers. comm.) and partially do-
mesticated populations prior to
the complete loss of seed for
some diploid cultivars commonly
produce seed cannot be differ-
entiated. In these circumstances it
is only the presence of Musaceae
species outside their natural rang-
es that might imply human trans-
mission and conrm evidence for
cultivation.
Where seed is absent from ar-
chaeobotanical assemblages, the
problem of identication becomes
more difcult and is reliant on dif-
ferentiation of the distinctive vol-
caniform morphotypes. From the
results of several studies (Lentfer
et al. 2003b, Mbida et al. 2001)
it is well-established that Ensete
species can be readily differenti-
ated from wild and domesticated
diploid and triploid Musa bananas
by morphotypic and morphomet-
ric means. However, differentiation
between Musa species is more
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
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257
9
7658
25
25
25
25 50 50
N=
species
4.0
3.5
3.0
2.5
2.0
1.5
9titles
95% Cl ratio (ld/td)
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
9titles
95% Cl ratio (ld/td)
M. banksii
M. acuminata
E. glaucum
M. fei
M. maclayi
M. paradisiaca
M. peekelii
M. schizocarpa
Figure 9. 95% condence intervals for mean long dimension (ld)/crater width (td)
ratios of leaf phytoliths differentiating Musa maclayi from all other bananas except
Musa schizocarpa.
complex and would be reliant on
a large sample size for any given
archaeobotanical assemblage.
Since triploid banana phytoliths
are generally larger than diploids
(Vrydaghs et al. 2009), there
could be scope for differentiating
domesticated triploid populations
from wild and cultivated diploid
populations by simply measuring
crater widths of archaeological
assemblages. Therefore, there
is potential for tracking banana
introductions and domestication.
Nevertheless, results show that
Eumusa and Australimusa sec-
tion bananas cannot be differen-
tiated at a general level and this
is problematic in regions where
bananas from both sections oc-
cur, either wild or cultivated. Indi-
cations are, however, that some
species and/or cultivars within
sections can be differentiated.
Most importantly, wild and do-
mesticated Eumusa bananas,
M. acuminata (AA) and M. bal-
bisiana (BB), can be differenti-
ated and there might be poten-
tial for tracking the introduction
of Musa acuminata bananas in
mainland Southeast Asia west
of the Philippines where M. bal-
bisiana dominates native banana populations. A similar
potential for discrimination is indicated for Australimusa;
M. maclayi and M. peekelii could not only be differenti-
ated from each other but also from the Australimusa do-
mesticate Fe`i. Therefore, there may be good potential for
tracking Australimusa banana dispersals and patterns of
domestication for the near Oceania region, east of Papua
New Guinea, where Australimusa bananas have dominat-
ed wild and cultivated populations.
The ‘New Guinea Banana Project’
Good potential for differentiating between banana phy-
toliths is indicated from morphometric and morphotyp-
ic analyses. Nevertheless, preliminary studies point to a
wide variation of morphotypes and additional study of a
larger sample incorporating additional species and culti-
vars is required to determine the extent of this variation
and further explore the potential for a more denitive set of
criteria for differentiation. The ‘New Guinea Banana Proj-
ect’ commenced in 2002 with collection of more than 100
wild and cultivated bananas from mainland Papua New
Guinea, New Britain and New Ireland (Table 3). Volcani-
form leaf phytoliths from 58 accessions were selected for
a more rigorous analysis (Table 4) than previously under-
taken, describing more morphological features (Table 5).
Digital images and measurements and morphometric de-
tails from 50 phytoliths per accession have been recorded
and saved on a readily accessible database. This in itself
is useful for identication of morphotypes during routine
analysis. Statistical analyses have yet to be completed.
Firstly, data will be lumped according to the same criteria
as Ball et al. (2006) and examined using the same statis-
tical procedure as Ball et al. (2006) and Vrydaghs et al.
(2009) for direct comparison. Subsequently, the analysis
will be re-run to incorporate the full set of morphotypic at-
tributes and morphometries.
Conclusions
Recent research (Denham et al. 2003, Lebot 1999, Per-
rier et al. 2009) shows that the New Guinea region has
played a key role in the development of the domesticated
banana, and complex origins and multiple dispersals for
banana cultivars within the southeast Asian/Pacic region
are indicated (Kennedy 2008). Phytoliths hold the key to
tracing the history of banana cultivation and domestica-
tion in the archaeobotanical record, and morphometric
and morphotypic analyses show good potential for differ-
entiation of phytoliths. Seed phytoliths can be readily dif-
Ethnobotany Research & Applications258
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Table 3. List of banana accessions collected from Papua New Guinea in 2002 for the ‘New Guinea Banana Project’, with associated ethnobotanical observations.
Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
Nari 1047 ABB Cape Gloucester Lowlands ?
Nari 146 AAB ENB Lowlands Luba
Nari 186 AAT Madang Lowlands Sar Amele
Nari
NBI 10
ABB ? Lowlands ?
Nari 168 AAB Central Province Lowlands ?
Nari
NBL 20
ABB Bougainville Lowlands ?
Nari
OBB 5
AAA Madang Lowlands Lakem
Nari 171 ABB Madang (orig.
Milne Bay)
Lowlands ?
Nari 142 AA ENB Lowlands Tagomor
Nari
NB9 11
ABB Morobe Lowlands Gana sumdu
Nari
NBI 18
AAB Chimbu Highlands Kunambo
Nari 206 AAB Western
Highlands
Highlands Rukumamb
tamby
Nari 164 AA Madang Lowlands Sihir
Nari
OBN 14
AAA ENB Lowlands Towberne
Nari 064 AA East Sepik Lowlands ?
WNB1 Ensete
glaucum
(Roxb.)
Cheesman
Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Tapupu Bakovi Seeds used for making beads.
WNB2 AA? Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Kikiyou Bakovi Cooked and also eaten ripe.
WNB3 ABB Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Tukuru Bakovi Cooked and also eaten ripe.
WNB4 ABB? Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Tamane buro Bakovi Cooked and also eaten ripe.
WNB5 AA? Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Maya Bakovi Cooking banana.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
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259
Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
WNB6 Musa textilis
Née
Tamare Village Lowlands 5
o
27
I
0.4
II
150
o
5
I
3.6
II
Tuain Pisin Pseudostem & sheath used for
making rope. Suckers sold to
villagers by agricultural supply
stores. Used as cash crop.
WNB7 ? Tamare Village Lowlands 5
o
27
l
0.4
ll
150
o
5
I
3.6
II
Hale vudi Bakovi Ornamental banana with red leaves.
WNB8 AA? Tamare Village Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Karuke vudi
(Karuke
=crab’s hand)
or Mrs. banana
Bakovi Eaten ripe.
WNB9 AA? Tamare Village Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
sausage
banana
English Cooked and also eaten ripe.
WNB10 Musa
acuminata
ssp. banksii
(Ridl.) Kiew.
Garu plantation Lowlands 5
o
31
l
12.9
ll
149
o
58
l
29.4
ll
Wil banana Pisin Wild banana growing along
margins of swamp.
WNB11 E. glaucum Tabairikau Lowlands 5
o
28
l
25.2
ll
150
o
27
l
27.2
ll
Vudu vudu Kuanua Collected from forest and grown
in garden with other bananas.
Necklaces made from seed. Sap red.
ENB1 ABB Tavui No.2 Lowlands 4
o
08
l
47.0
ll
152
o
10
l
02.4
ll
Yawa Kuanua Large seeds often present. Fruit
cooked and also eaten ripe. Fruit
also fed to pigs. Stems and leaves
used for mumu. Fruit used for
making an alcoholic beverage. The
ripe fruit is placed in drum and after
heating to release juices the liquid is
fermented. It is distilled into a clean
drum and put into small bottles.
The process takes two weeks.
ENB2 ABB Tavui No. 2 Lowlands 4
o
08
l
47.0
ll
152
o
10
l
02.4
ll
Kalapua Kuanua Cooked and also eaten ripe.
Fruit fed to pigs. Stems and
leaves used for mumu.
ENB3 ABB Tavui No. 2 Lowlands 4
o
08
l
58.9
ll
152
o
10
l
27.1
ll
Tukuru Kuanua Fruit cooked. Male bud is also eaten.
ENB4 ? Tavui No. 2 Lowlands 4
o
13
l
26.3
ll
152
o
10
l
27.1
ll
Kudukudu Kuanua Eaten ripe and cooked.
ENB5 ? Malaguna Lowlands 4
o
08
l
58.9
ll
152
o
08
l
59.0
ll
Kakatur Kuanua Eaten ripe.
ENB6 ? Gela Gela Lowlands 4
o
24
l
38.0
ll
152
o
14
l
48.6
ll
Pok pok Kuanua Cooking banana. Leaves easily torn and
unsuitable for cooking and wrapping.
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Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
ENB7 ? Gela Gela Lowlands 4
o
24
l
36.7
ll
152
o
14
l
51.6
ll
Makau Kuanua Cooking banana. Leaves easily torn and
unsuitable for cooking and wrapping.
ENB8 AAA Gela Gela Lowlands 4
o
24
l
36.1
ll
152
o
14
l
49.1
ll
Katur Kuanua Cooking banana. Leaves
used for mumu.
ENB9 AA Gela Gela Lowlands 4
o
24
l
36.1
ll
152
o
14
l
49.1
ll
Pitu Kuanua Cooking banana. Leaves easily torn and
unsuitable for cooking and wrapping.
ENB10 ABB Takabua Mission
Station
Lowlands 4
o
22
l
29.8
ll
152
o
13
l
3.1
ll
Makala tukuru Kuanua Eaten ripe. Leaves used for mumu.
ENB11 ABB Malakuna No. 4 Lowlands 4
o
23
l
6.0
ll
152
o
12
l
18.3
ll
Yawa buka Kuanua Eaten ripe. Leaves used for mumu.
ENB12 BB? Takabua Mission
Station
Lowlands 4
o
22
l
29.8
ll
152
o
13
l
3.1
ll
Okaoko Kuanua Eaten ripe. Leaves used for mumu.
ENB13 Fe`i Raolo Lowlands 4
o
5
l
40.5
ll
152
o
4
l
31.2
ll
Auro=Vuro Kuanua Eaten ripe and cooked.
ENB14 Fe`i Malmaluan
Training College
Lowlands 4
o
17
l
49.6
ll
152
o
1
l
41.0
ll
Vuro Kuanua Eaten ripe and cooked. Leaves used
for mumu.
ENB15 AA L.A.E.S. Kerevat Lowlands 4
o
20
l
7.0
ll
152
o
1
l
41.0
ll
Gulum Kuanua Cooking banana. Leaves easily torn and
unsuitable for cooking and wrapping.
ENB16 Fe`i L.A.E.S. Kerevat Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
Vuro/Pitu Kuanua Eaten ripe. Leaves used for mumu.
ENB17 AB? L.A.E.S. Kerevat Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
Taupen Kuanua Cooking banana.
ENB18 AAA L.A.E.S. Kerevat Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
Malam red Kuanua Eaten ripe and cooked.
ENB19 AS? L.A.E.S. Kerevat Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
ENB20 M. textilis L.A.E.S. Kerevat Lowlands 4
o
20
l
9.2
ll
152
o
1
l
52.1
ll
Pseudostem and sheath used for
making rope. Suckers sold by L.A.E.S.
to agricultural supply stores.
NI1 ABB? cf.
Tukuru
(Kokopo)
Kulangit Village,
Kavieng
Lowlands 2
o
33
l
43.1
ll
150
o
47
l
54.6
ll
Sukuru Tigak Cooking banana but also eaten ripe.
Leaves strong and used for mumu,
and wrapping. Male bud also eaten.
NI2 AA? Kulangit Village,
Kavieng
Lowlands 2
o
33
l
43.1
ll
150
o
47
l
54.6
ll
Papat (cf.
Garamut in
Kuanua)
Tigak Cooking banana - fruits boiled
and roasted. Leaves weak - used
for covering mumu only.
NI3 AA Kulangit Village,
Kavieng
Lowlands 2
o
33
l
43.1
ll
150
o
47
l
54.6
ll
Kikiyou Tigak Cooking banana - fruits boiled,
fried and roasted. Leaves weak -
used for covering mumu only.
NI4 ABB? Kulangit Village,
Kavieng
Lowlands 2
o
33
l
45.5
ll
150
o
47
l
54.1
ll
Wan kina,
Wan pound
Pisin Cooked in mumu and eaten raw.
NI5 ? Kulangit Village,
Kavieng
Lowlands 2
o
33
l
45.5
ll
150
o
47
l
54.1
ll
Boa Tigak Cooked in mumu and eaten
raw and boiled. Sweet.
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An update from Papua New Guinea
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261
Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
NI6 Fe`i? Kulangit Village,
Kavieng
Lowlands 2
o
33
l
45.5
ll
150
o
47
l
54.1
ll
Suluklamuk Tigak Eaten ripe and cooked. Yellow esh
with poorly formed seeds eaten
by ants. Turns urine yellow. Bunch
drooping. Possibly intermediate form.
Leaves strong and used in mumu.
NI7 ? Kulangit Village,
Kavieng
Lowlands 2
o
33
l
45.5
ll
150
o
47
l
54.1
ll
Darip Tigak Eaten ripe and cooked. Yellow esh.
Leaves not used.
NI8 AAA Kulangit Village,
Kavieng
Lowlands 2
o
33
l
41.7
ll
150
o
47
l
52.1
ll
Palang Tigak Eaten ripe and cooked. Yellow esh.
Leaves not used.
NI9 AA Kulangit Village,
Kavieng
Lowlands 2
o
33
l
41.7
ll
150
o
47
l
52.1
ll
Ngitingsakai Tigak Cooking banana - boiled and roasted.
Leaves not used. Fruit occurs in
continuous spiral around rachis.
NI10 AA Kulangit Village,
Kavieng
Lowlands 2
o
33
l
42.5
ll
150
o
47
l
56.4
ll
Ulungan Tigak Fruit roasted or boiled. Leaves not
used.
NI11 AAB Kulangit Village,
Kavieng
Lowlands 2
o
33
l
48.4
ll
150
o
47
l
56.1
ll
Bavaya Tigak Fruit cooked or eaten raw. Leaves used
for wrapping.
NI12 AA Miom/Caranas,
Kavieng
Lowlands 2
o
35
l
25.6
ll
150
o
49
l
41.4
ll
Papat Wung Tigak Fruit roasted or boiled. Leaves used for
mumu and wrapping.
N13 AAB Miom/Caranas,
Kavieng
Lowlands 2
o
35
l
25.6
ll
150
o
49
l
41.4
ll
White bavaya English/Tigak Mostly roasted.
NI14 Fe`i Namasalang Lowlands 3
o
2
l
45.4
ll
151
o
24
l
57.1
ll
Utafan Malik Cooked and eaten raw - sweet. Sap
used as a dye for malangan masks.
NI15 ? Lelet <1000m 3
o
16
l
20.7
ll
151
o
55
l
39.0
ll
Siaman/
German
Mandak Eaten ripe.
NI16 Fe`i Kaluan, Lelet <1000m 3
o
19
l
28.8
ll
151
o
54
l
22.9
ll
Manui/
Namnam
Manui
(Manus
Island)/
Mandak
Eaten ripe or cooked. Leaves used for
mumu and wrapping. Bunch drooping
but at an angle and almost horizontal.
NI17 Fe`i Kaluan, Lelet <1000m 3
o
17
l
58.3
ll
151
o
54
l
48.7
ll
Loskauk Mandak Eaten ripe or cooked. Leaves used for
mumu and wrapping. Bunch drooping
but at an angle and almost horizontal.
NI18 Fe`i Kabagong, Lelet <1000m 3
o
16
l
55.4
ll
151
o
55
l
30.2
ll
Loskauk Mandak Cooking banana. Flesh sulfur-yellow.
NI19 Musa maclayi
F. Muell. ex
Mikl.-Maclay
ssp. maclayi
Arg. var.
namatani Arg.
Karu <1000m 3
o
29
l
52.7
ll
152
o
11
l
46.9
ll
Agigilai Barok Wild. Said not to be eaten but sweet
bright yellow esh surrounds seeds.
Leaves used for wrapping and mumu.
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Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
NI20 M. maclayi?
Fe`i?
Karu <1000m 3
o
30
l
20.3
ll
152
o
12
l
21.2
ll
Barok Wild. Tall, robust banana with shiny
pseudostem. Not fruiting at time of
collection.
NI21 M. maclayi
ssp. maclayi
var. namatani
Karu <1000m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Agigilai Barok Wild. Said not to be eaten but sweet
bright yellow esh surrounds seeds.
Leaves used for wrapping and mumu.
NI22 Musa peekelii
Lauterb. ssp.
peekelii Arg.
Rosirik <1000m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Wawa Sokiri Wild robust banana. Bunch erect at rst
then drooping. Fruit cooked or roasted.
Leaves used to cover mumu.
NI23 M. peekelii
ssp. peekelii
Rosirik <1000m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Wawa Sokiri Wild robust banana. Bunch erect at rst
then drooping. Fruit cooked or roasted.
Leaves used to cover mumu.
WH1 Musa ingens
Simmonds
Tubang, Kauil 2000m 5
o
27
l
0.4
ll
150o5
l
3.6
ll
Wuluk donu Jiwaka Wild tall banana growing to 16m to
18m height at 2000m in creek gully.
Fruit with yellow esh and large seeds,
green turning yellow when ripe. Cuscus,
parrots and ying fox eat fruit. Leaves
used for making temporary shelters
and for covering mumu.
WH2 M. ingens Tubang, Kauil 2000m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Wuluk donu Jiwaka Wild tall banana growing to 16m to 18m
height at 2000m altitude in creek gully.
Fruit with yellow esh and large seeds.
Leaves used for making temporary
shelters and for covering mumu.
WH3 M. ingens Tubang, Kauil 2000m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Wuluk donu Jiwaka As above.
WH4 Fe`i Balga, Kuk 1630m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Rua mema
(rua = banana,
mema = red)
Cooking banana, roasted and boiled -
preferably roasted.
WH5 Musa
schizocarpa
Simmonds
Baiyer Zoo 1177m 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
? ? Wild banana growing in rainforest with
gingers, ferns, Rubiaceae shrubs, tree
ferns and wild taro in rainforest in shade
under canopy.
ES1 AAT? Ranimbo, Hawain Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Yesing (=wild
banana)
Mino Tall robust banana with edible fruit.
Feral rather than wild. Fruit eaten when
raw or cooked. Leaves used for mumu
and wrapping. Fruit also fed to pigs.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
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263
Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
ES2 AA? CCRI station,
Hawain
Lowlands 5
o
31
l
12.9
ll
149
o
58
l
29.4
ll
Yesing (=wild
banana)
Mino Fruit green turning yellow when ripe.
Leaves used as decoration in singsing
celebrations after heating on re to
make pliable.
ES3 M.
schizocarpa
CCRI station,
Hawain
Lowlands 5
o
28
l
25.2
ll
150
o
27
l
27.2
ll
Yesing (=wild
banana)
Mino Wild seeded banana with yellowish
sap.
ES4 M.
schizocarpa
Ranimbo, Hawain Lowlands 4
o
08
l
47.0
ll
152
o
10
l
02.4
ll
Yesing (=wild
banana)
Mino Wild seeded banana with clear sap.
Leaves used for mumu, wrapping food
and for singsing decoration as well as
roofs for bush houses.
ES5 Fe`i Koikin Village,
Wewak
Lowlands 4
o
08
l
47.0
ll
152
o
10
l
02.4
ll
Taweyawa Mino Cultivated banana with red sap.
Fruit large 4-5cm diameter, 10-23cm
long with bottle-necked apex and
pronounced ridging. Mostly cooked.
ES6 M. acuminata
ssp. banksii
Koikin Village,
Wewak
Lowlands 4
o
08
l
58.9
ll
152
o
10
l
27.1
ll
Yesing (=wild
banana)
Mino Wild seeded banana with clear sap.
Fruit turns yellowish when ripe. Not
eaten. Said to often colonize landslides
with gingers and grasses. Species
common along Sepik Highway towards
Maprik. Often grows in association with
M. schizocarpa. This specimen growing
on well-drained loam along roadside
next to gardens and close to regrowth
forest. One plant only.
ES7 ABBT? Japaraga 1,
Maprik Road
<1000m 4
o
13
l
26.3
ll
152
o
10
l
27.1
ll
Giant kalapua Pisin Cooking banana. Leaves used for
wrapping and mumu.
ES8 AA Japaraga 1,
Maprik Road
<1000m 4
o
08
l
58.9
ll
152
o
08
l
59.0
ll
Salamua,
pisang
jari buaya
(crocodile
ngers)
?, Mino Fruit eaten ripe or cooked. Leaves used
for mumu and dried leaves dried and
used for making cigarettes. Clear sap.
ES9 AA Japaraga 1,
Maprik Road
<1000m 4
o
24
l
38.0
ll
152
o
14
l
48.6
ll
Heifeli’ To`o Mino,
Madang
language?
Fruit eaten ripe or cooked. Leaves
used for mumu.
ES10 M. acuminata
x schizocarpa
Japaraga 1,
Nagam River
<1000m 4
o
24
l
36.7
ll
152
o
14
l
51.6
ll
Yesing (=wild
banana)
Mino Wild banana growing on sandy river
bank. Sap clear. Fruit not eaten.
ES11 M. acuminata
ssp. banksii
Japaraga 1,
Nagam River
<1000m 4
o
24
l
36.1
ll
152
o
14
l
49.1
ll
Yesing (=wild
banana)
Mino Wild banana growing in shady regrowth
forest with moss understory.
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Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
M1 Fe`i Motonau/Midiba,
Madang
cl 4
o
24
l
36.1
ll
152
o
14
l
49.1
ll
Wahin Ari Cultivated banana with red sap growing
on well-drained red/brown loam. Fruit
large with pronounced ridging and
sulfur-yellow esh. Mostly cooked. Fe`i
bananas once common in the district
but said to be gradually dying out, often
surviving in abandoned garden settings.
Notable Fe`i bananas are referred to
as “yelo pispis” in Pisin because urine
turns yellow after the fruit is eaten. Fruit
cooked and used for coloring other food
stuffs and enhancing avours. Red sap
used by children for making stencils on
clothing.
M2 AA Stewart Research
Station, Madang
Lowlands 4
o
22
l
29.8
ll
152
o
13
l
3.1
ll
Manameg
(red)
Amele Small cultivated banana. Cooking
banana referred to as plantain although
strictly speaking it isn’t. Leaves used
for mumu.
M3 AA? Stewart Research
Station, Madang
Lowlands 4
o
23
l
6.0
ll
152
o
12
l
18.3
ll
Kekiau Kuanua Cooking banana, boiled or roasted.
Leaves used for mumu and wrapping
food.
M4 M.
schizocarpa
Baitata Road,
Madang
Lowlands 4
o
22
l
29.8
ll
152
o
13
l
3.1
ll
Maror Em Wild banana growing on poorly drained
limestone soil in regrowth forest along
roadside. Growing near Musa peekelii
ssp. angustigemma. (NB. Site used for
making the lm “Robinson Crusoe”).
Fruit self-peels to give a star-like
appearance to the apex. Skin green
when ripe. Flesh white. Leaves used to
make temporary bush shelters.
M5 M. peekelii
ssp.
angustigemma
Baitata Road,
Madang
Lowlands 4
o
5
l
40.5
ll
152
o
4
l
31.2
ll
Dor Em Tall robust wild banana growing near M.
schizocarpa. Common in well-drained
locations on coral/limestone soils
in regrowth forest. Bunch drooping,
falling straight. Flowers eaten - mixed
with salt. Young pseudostem also
eaten opportunistically. Flowers and
pseudostem used to garnish taro.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
265
Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
M6 M. acuminata
ssp. banksii
Baitata Village,
Madang
Lowlands 4
o
17
l
49.6
ll
152
o
1
l
41.0
ll
Tilja Em Wild banana growing in association
with M. peekelii ssp. angustigemma
on coral/limestone soils at edges of
regrowth forest.
M7 M. acuminata
ssp. banksii
Barum junction,
Madang
Lowlands 4
o
20
l
7.0
ll
152
o
1
l
41.0
ll
Mabul kulal
(mabul=wild:
kulal=inedible,
left to dry)
Kein Wild banana in moderately dense
stand growing on well-drained soil
in regrowth forest along roadside. In
close association with M. schizocarpa
and AS hybrids.
M8 AS Barum junction,
Madang
Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
Mabul jus
(jus= water
snail because
the snail has
a pointed
shell and the
banana has a
long pointed
apex).
Kein Wild banana growing on well-drained
soil in regrowth forest along roadside.
Growing amongst mixed stand of
M. acuminata, M. schizocarpa and
hybrids.
M9 M.
schizocarpa
Barum junction,
Madang
Lowlands 4
o
20
l
7.5
ll
152
o
1
l
48.2
ll
Hirunag
(hiru=ginger;
nag=son)
Kein Wild banana growing on well-drained
soil in regrowth forest along roadside.
Growing amongst mixed stand of M.
acuminata and M. schizocarpa and AS
hybrids.
M10 AA Jobtou, Madang Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bagui Kein Cultivated species very much like
the wild M. acuminata ssp. banksii in
appearance. Male bud green.
MB1 E. glaucum Naura Village Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Gudu gudu Haigwai Growing in neglected coconut
plantation. Seeds used to make
necklaces.
MB2 M.
schizocarpa
Naura Village Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Lau moi moi
(lau=inedible;
moi=banana)
Haigwai Wild banana growing in overgrown
gardens. Leaves sometimes used for
covering food. Species also found at
Gomila Village.
MB3 M. maclayi
ssp. maclayi?
Gomila Village,
Ahioma
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bihibihiya
(bihibihi= wild
banana; bihi
= cultivated
banana)
Tawala Wild banana with dark, milky purple-red
sap and erect bunch. Fruit with sulfur-
yellow esh. Growing in cool, sheltered
gully in regrowth forest. Another small
stand found growing nearby on rocky
talis slope beside creek in old garden
bordering regrowth forest.
Ethnobotany Research & Applications266
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Code Type Location Altitude
(m)
Latitude
S
Longitude
E
Vernacular
Name
Language Notes
MB4 M. maclayi
ssp. maclayi?
Gomila Village,
Ahioma
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bihibihiya
(bihibihi= wild
banana; bihi
= cultivated
banana)
Tawala One of stand of wild bananas with
dark, milky purple-red sap growing
on talis slope in old garden alongside
creek. Small stands of species found in
dense regrowth forest on steep banks
bordering creek.
MB5 M. maclayi
ssp. maclayi?
Gomila Village,
Ahioma
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bihibihiya
(bihibihi= wild
banana; bihi
= cultivated
banana)
Tawala Wild banana with pink sap and erect
bunch, tight but lax at proximal end.
Fruit with sulfur-yellow esh. Growing
in gardens on raised sandy gravel
deposits bordered by creek channels.
MB6 M. maclayi
ssp. maclayi?
Daduwe area,
Northeast
Coast Road
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bihibihiya
(bihibihi= wild
banana; bihi
= cultivated
banana)
Tawala Wild banana in small stand of bananas
growing on well-drained soil along
roadside in tall grassland.
MB7 M. maclayi
ssp. maclayi?
Daduwe area,
Northeast
Coast Road
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Bihibihiya
(bihibihi= wild
banana; bihi
= cultivated
banana)
Tawala Wild banana growing in small scattered
stand on well-drained soil along roadside
in tall grassland. Some specimens
feature retention of bracts on the rachis
while others shed bracts. Notably
bract retention is a characteristic of M.
maclayi ssp. ailului Arg. but in other
respects the specimens t more closely
with M. maclayi ssp. maclayi according
to Argent’s key.
MB0 M. maclayi
ssp. maclayi?
Kuiaro Village,
China Strait
Lowlands 5
o
27
l
0.4
ll
150
o
5
l
3.6
ll
Specimen found on hillside in regrowth
forest recently cleared for gardening.
Also noted to be growing in forest areas
inland from the coast.
MB0 M. maclayi
ssp. maclayi?
Tue Village Lowlands 5
o
31
l
12.9
ll
149
o
58
l
29.4
ll
Lau moi moi
(lau=false;
moi=banana)
Daiamoni Said to be growing in bush in creek
catchment. Not collected.
MB8 AA Tue Village Lowlands 5
o
28
l
25.2
ll
150
o
27
l
27.2
ll
Kokoluyu Daiamoni Cultivated banana. Fruit eaten cooked
or ripe.
MB9 AA? AS? Tue Village Lowlands 4
o
08
l
47.0
ll
152
o
10
l
02.4
ll
Masi masi Daiamoni Cultivated banana. Fruit eaten cooked
or ripe.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths:
An update from Papua New Guinea
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
267
Table 4. Accessions selected for the ‘New Guinea Banana Project’ analyses.
Section Number
analyzed
Accession code
Wild species
Eumusa Musa acuminata ssp. banksii 5 ES11, ES6, M7, M6, ENB10
Musa schizocarpa 5 ES3, WH5, MB2, M9, ES4
Musa acuminata ssp. banksii x schizocarpa 2 M8, ES10
Australimusa Musa peekelii 2 M5, NI22
Musa maclayi 5 NI21, MB7, MB4, MB3, MB5
Musa textilis 2 ENB20, WNB6
Ingentimusa Musa ingens 3 WH1, WH2, WH3
N/A Ensete glaucum 4 WNB1, WNB11, MB1, MB1/2
Cultivars*
Eumusa AA 5 NI12, ENB13, Nari064,
Nari164, MB8
AAA 5 NI8, NariOBB5, NariNB420,
NariOBN14, ENB18
BB? 1 ENB12
AAB 5 NI13, Nari206, Nari146, NI11, NB1
ABB 5 Nari1047, NariNBG11, NariNBI10,
Nari171, NariNBL20
AB 1 ENB17
Australimusa TT(Fe`i) 5 WH4, ES5, ENB13, M1, ENB14
Eumusa x
Australimusa
AAT 2 Nari186, ES1?
ABBT? 1 ES7
*Genome labels for diploid, triploid and polyploid cultivars: A = acuminata, B = balbisiana, T = Australimusa.
Table 5. Expanded list of categories and attributes used in analysis of volcaniform leaf phytoliths for the ‘New Guinea
Banana Project’.
Category Attribute
Base, 3D shape tabular, blocky, spherical( globose), platy (planar)
Base, 2D shape round, oval, square, rectangle, quadrilateral, boat (oblong), irregular
Body length -
Body width -
Body height short (h<1/3 length), medium (h=1/3 to ½ length), tall (h≥1/2 length)
Crater length -
Crater width -
Crater shape (dorsal view) round (orbicular), oval, quadrilateral, irregular
Rim present, absent
Rim shape regular, irregular, skirt (crenate)
Sides straight, convex, concave, straight/concave, concave/convex, straight/convex
Body texture psilate, granulate, verrucate, nodulose, tuberculate, psilate/verrucate, psilate/granulate,
granulate/verrucate
Base ornamentation absent, short (<1.25µm), medium (1.25-2.5µm), long (>2.5µm), long/tuberculate/
dendritic, short/medium, medium/long
Ethnobotany Research & Applications268
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
ferentiated from phytoliths generated by other plant parts,
and the absence of seed phytoliths in an archaeobotani-
cal assemblage can signal presence of seedless/domes-
ticated bananas. However, parthenocarpy and sterility,
processes inherent in banana domestication, evolved
over several generations, and populations of seeded cul-
tivated diploid bananas are still common in Papua New
Guinea. Therefore, it can be difcult to determine the sta-
tus of bananas (wild or cultivated) in the archaeobotanical
record unless there is unequivocal evidence for bananas
either being outside their natural range, associated with
archaeological and pedogenic features indicative of cul-
tivation, and/or associated with other known cultigens. In
the absence of diagnostic seed phytoliths identication is
reliant on volcaniform leaf phytoliths. Studies show that
some species and cultivars can be discriminated accord-
ing to crater size and body length and by the presence
of certain rare morphotypes. Nevertheless, there is large
morphotype variation within and between species; conse-
quently, identication is currently reliant on large sample
sizes rarely encountered in fossil assemblages. There-
fore, more work is needed to clarify the extent of variability
across the geographic range and within different habitats.
The ‘New Guinea Banana Project’ commenced in 2002
with the eld collection of samples and resultant analysis
of 58 additional accessions. The outcome of these analy-
ses should help to resolve many of the outstanding issues
regarding the differentiation of volcaniform leaf phytoliths
in the archaeobotanical record, particularly in the Pacic/
Papua New Guinea region.
Acknowledgements
I am most grateful to the Pacic Science Foundation
and the University of Queensland for providing the fund-
ing for this project. My sincere thanks to Clair Harris for
her assistance. I also thank Jeff Daniells, Bill Boyd and
Robin Torrence who assisted with the eld collection and
the PNG government ofcials and staff (NARI, DAL, DPI,
CCEA and CCRI) in particular Valentine Kambori, Rosa
Kambuou and Jeff Wiles for their support and kind assis-
tance with getting the project up and running, and Janet
Paofa, Martin Gunther, David Minemba, James Maima,
Moses Woruba, Will Akus, Joseph Wasem, Marcel Toko-
rewaga, Kauwe Murley, Luke Nama, Louis Kurika, March
Tovue, Joy Penias, Scola Singit, Nick Eki and Manau Peni
for their kind assistance. I also thank Nick Lyons for his
kind hospitality and all the people who showed us around
their gardens and allowed us to take samples.
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... Satu temuan menarik dari sampel residu gerabah adalah fitolit bertipe volcani form (Gambar 8) yang mengarah pada tanaman pisang Musa sp. atau Ensete sp (Ball, Vrydaghs, Van Den Hauwe, Manwaring, & De Langhe, 2006;Horrocks & Rechtman, 2009;Lentfer, 2009;Mindzie et al., 2001). ...
... Gambar 8. (a) dan (b) adalah fitolit berbentuk volcani form pada residu gerabah serta (c) adalah referensi fitolit pisang yang diambil dari Perrier et al., 2011;Mindzie et al., 2001, Horrocks & Rechtman, 2009, Lentfer, 2009. (Sumber : Alifah et al., 2020 Temuan fitolit tanaman Zingiberaceae pada residu artefak batu menunjukkan intensitas yang sangat tinggi. ...
... Pemanfaatan pisang sebagai sumber pangan lazim dilakukan karena buah, bunga, dan batang semuanya merupakan bagian yang dapat dikonsumsi. Selain itu, pisang dapat dimanfaatkan untuk kepentingan non-pangan, seperti keperluan pengolahan makanan (daun, batang semu), bahan tali (batang semu), serta manik-manik dan mata uang (biji) (Burkill, 1936;Lentfer, 2003Lentfer, , 2009. Situs arkeologi yang memiliki temuan fitolit pisang dengan pertanggalan yang cukup tua telah menunjukkan bahwa pisang merupakan salah satu tanaman penting seperti di Situs Kuk Swamp, Papua Nugini, yang sejauh ini merupakan bukti domestikasi pisang tertua pada kala Holosen sekitar 10000 BP (Denham et al., 2003). ...
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Around 400 bc , pottery- and iron-producing populations immigrated into the Inner Congo Basin (ICB) and subsequently spread upstream some major tributaries of the Congo River. Until recently, their subsistence was almost completely unknown. We present an archaeobotanical study of three sites in the ICB covering parts of the Early Iron Age (ca. 400 bc - ad 650) and of the Late Iron Age (LIA) as well as subrecent times (ca. ad 1300–2000). We studied 82 flotated samples of botanical macroremains, and 68 soil phytolith samples, recovered from the terra firme sites Iyonda and Mbandaka, and the floodplain fishing camp site of Bolondo. The EIA assemblage from Iyonda yielded domesticated Cenchrus americanus (pearl millet), Vigna unguiculata (cowpea), Canarium schweinfurthii , Elaeis guineensis (oil palm), several wild plants, and parenchyma fragments tentatively attributed to Dioscorea sp. (yams). The exploitation of these plants originated in the savannas and forest-savanna ecotones of West Africa. The presence of C. americanus in LIA contexts at Bolondo and Mbandaka, dated to ca. ad 1350–1550, indicates that its cultivation is not dependent on a seasonal climate with a distinct dry season, contrary to previous views. The role of C. americanus as a staple is difficult to assess; it might have been used for special purposes, e.g. beer brewing. In spite of extensive screening, we did not detect any banana phytoliths in the EIA samples. Musa phytoliths were only present in LIA contexts after ca. ad 1400, leaving room for the possibility that the introduction and spread of Musa spp. AAB ‘Plantain’ in the ICB was a late phenomenon.
... Henriet et al. (2006) [7] reported Si concentration and uptake in banana increased with silica supply. The plants are known to produce phytoliths (discrete silicon deposits) in the leaves and pseudostem, which persist in the soil long after plant death, leading to their use in archaeological studies to track the cultivation and consumption of banana [13]. When silicon is present in the soil solution at high concentrations, banana plants absorb it passively but switch to active uptake via theoretical transporters when concentrations are low [6]. ...
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Field experiment was conducted in Farmers holding located at Rajagopalapuram village under Kuttalam taluk, Mayiladuthurai district, Tamilnadu, India belonging to Padugai series (Typic ustifluvent) low in available silicon to elucidate the response of banana in terms of yield, nutrition and quality due to silicon nutrition applied through potassium silicate. The treatment consists of T1 - NPK (RDF), T2 - NPK + Potassium silicate (FS) - 0.25%, T3- NPK + Potassium silicate (FS) - 0.50%, T4 - NPK + Potassium silicate (FS) - 1.00%, T5­ - NPK + Potassium silicate (FS) - 0.25%, T6 - NPK + Potassium silicate (FS) - 0.50%, T7­ - NPK + Potassium silicate (FS) - 1.00%, T8 - NPK + Potassium silicate (SA) - 50 kg ha-1, T9 - NPK + Potassium silicate (SA) - 100 kg ha-1 and T10 - NPK + Potassium silicate (SA) – 150 kg ha-1. From T2 to T4 foliar spray was done at 3rd and 5th month and from T5 to T7, foliar spray was done at 3rd, 5th and 7th month. The test crop is Banana Var. Grand Naine. The results showed that the highest fruit yield was noticed with 50 kg Si ha-1 (96.0 t ha-1) and it reduced with Si levels. Among foliar treatments, fruit yield increased with silicon concentrations (0.25 to 1%). The highest fruit yield was observed with 1% Si applied at 3rd and 5th month (92.7 t ha-1). Foliar spray of 0.25 and 0.5% Si applied thrice increased the fruit yield compared to twice application at same concentration. But fruit yield declined when Si was sprayed at 1% thrice. Soil application of silicon recorded higher nutrient uptake over foliar addition. The nutrient uptake was maximum with 150 kg Si ha-1. With respect to foliar treatments, the highest uptake was noticed with foliar spray of 1% Si applied at 3rd and 5th month. Soil application of 50 kg Si ha-1 recorded the highest fruit quality viz., TSS, total sugars, reducing and non-reducing sugar and leaf protein. The highest fruit quality was noticed with foliar spray of 1% Si applied at 3rd and 5th month.
... banksii does not grow. Similar archaeological remains were identified in the Bismarck archipelago and Vanuatu that are both outside the natural range of the species, pleading for a human introduction (Lentfer 2009;Tromp et al. 2020). The existence of M. acuminata ssp. ...
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This study is an unprecedent exploration of the diversity of 226 diploid bananas genotyped with restriction-site-associated DNA sequencing data (RADseq) to clarify the processes that led to the creation of edible diploid AA bananas. This wide set included 72 seedy bananas, mostly M. acuminata from different genepools, and 154 edible, i.e. parthenocarpic and sterile, AA accessions obtained from genebanks and recent collecting missions. We highlighted the geographic organisation of the diversity of edible AAs and confirmed the admixed nature of many and further conducted introgressions tests within AAs from South East Asia and New Guinea. Lastly, taking advantage of the presence of an important number of M. acuminata ssp. banksii (22) and of AA from Papua New Guinea (76) in the set, we investigated the patterns of differentiation between wild and cultivated bananas seemingly belonging to the same genepool. We discovered a few cultivated AAs that may be of pure origins both in South-East Asia and in New Guinea. We also detected two undefined parental genepools in South East Asia for which regions of origin could be Thailand and a region between north Borneo and the Philippines, respectively. Finally, we suggest the existence of a third genepool in New Guinea island that might be a source population for both edible AAs and the local M. acuminata ssp. banksii.
... Wild bananas are absent from the flora of Mabuyag 37 ; today bananas only occur as garden cultivars near present-day settlements and are not found on the site. Additionally, phytoliths typical of banana seeds were not observed in the microfossil assemblage 44 , further supporting a scenario involving the introduction of cultivars rather than wild plants. The older microfossil evidence lacks definitive association with the archaeological remains of former cultivation practices, such as a terrace, and the absence of a palaeoecological signature of environmental transformation hinders more definitive interpretations of land-use practices. ...
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Multiproxy archaeobotanical analyses (starch granule, phytolith and microcharcoal) of an abandoned agricultural terrace at Wagadagam on Mabuyag Island, Torres Strait, Australia, document extensive, low-intensity forms of plant management from at least 2,145–1,930 cal yr bp and intensive forms of cultivation at 1,376–1,293 cal yr bp. The agricultural activities at 1,376–1,293 cal yr bp are evidenced from terrace construction, banana (Musa cultivars) cultivation and dramatic transformations to the local palaeoenvironment. The robust evidence for the antiquity of horticulture in western Torres Strait provides an historical basis for understanding the diffusion of cultivation practices and cultivars, most likely from New Guinea. This study also provides a methodological template for the investigation of plant management, potentially including forms of cultivation that were practiced in northern Australia before European colonization. Multiproxy archaeobotanical analyses of an abandoned agricultural terrace at Wagadagam document extensive, low-intensity forms of plant management from at least 2,145–1,930 cal yr bp and intensive forms of cultivation at 1,376–1,293 cal yr bp.
... In part, this reflects limited preservation of macrobotanical remains of many tropical cultivars in wet tropical and subtropical environments, excepting nut shells and fruit stones. In part, though, this reflects uncertainties regarding the microbotanical (primarily phytolith and starch granule) diagnostics of domestication for most crop plants in the region, with bananas being a possible exception (Lentfer, 2009;Vrydaghs et al., 2009). ...
Book
Theory in the Pacific, the Pacific in Theory explores the role of theory in Pacific archaeology and its interplay with archaeological theory worldwide. The contributors assess how the practice of archaeology in Pacific contexts has led to particular types of theoretical enquiry and interest, and, more broadly, how the Pacific is conceptualised in the archaeological imagination. Long seen as a laboratory environment for the testing and refinement of social theory, the Pacific islands occupy a central place in global theoretical discourse. This volume highlights this role through an exploration of how Pacific models and exemplars have shaped, and continue to shape, approaches to the archaeological past. The authors evaluate key theoretical perspectives and explore current and future directions in Pacific archaeology. In doing so, attention is paid to the influence of Pacific people and environments in motivating and shaping theory-building. Theory in the Pacific, the Pacific in Theory makes a significant contribution to our understanding of how theory develops attuned to the affordances and needs of specific contexts, and how those contexts promote reformulation and development of theory elsewhere. It will be fascinating to scholars and archaeologists interested in the Pacific region, as well as students of wider archaeological theory.
... Thus, it is not possible to undertake a systematic comparison of the data for ancient phytoliths. Furthermore, Premathilake and Hunt fail to address methodological issues identified in previous research that has found the differentiation of wild bananas, cultivated diploids and cultivated triploids extremely difficult using morphometric or morphological criteria (Ball et al., 2006;Lentfer, 2009;Vrydaghs et al., 2009;Lentfer and Denham, 2017). Given these methodological uncertainties, there is no robust basis for discriminating wild and domesticated bananas in the phytolith assemblage at Fahien. ...
Article
A recent publication on the phytolith assemblage at Fahien rockshelter, Sri Lanka (Premathilake and Hunt) is argued to represent: the exploitation of wild Musa acuminata and M. balbisiana during the Late Pleistocene; the introduction of edible diploid cultivars from the Southeast Asia–New Guinea region during the early to mid‐Holocene; the generation and cultivation of triploid banana hybrids on Sri Lanka before 6194–5994 cal bp; and the subsequent spread of derived triploid cultivars to mainland India and westward to Africa. A careful review of the archaeobotanical research presented by Premathilake and Hunt, in the context of broader multidisciplinary evidence (agronomy, archaeobotany, genetics and linguistics) for the domestication and spread of banana cultivars, indicates that three main aspects of their argument are problematic: the lack of clarity in the characterization of banana domestication in the past; the methods used to discriminate phytoliths into banana taxa; and the promotion of Sri Lanka as a source region rather than a recipient of banana cultivars. Following reconsideration, the Fahien evidence is consistent with previous interpretations for the origins of diploid and significant triploid cultivars outside of Sri Lanka and dispersal to that island.
Article
Starch granules can be well preserved in a variety of archaeological contexts, for example, in residues and sediments. Therefore, starch analysis has potential to provide another means of tracking the exploitation, dispersal and domestication of Musa bananas and Ensete. Starch granule morphotypes from fruits of Ensete glaucum and wild and cultivated Australimusa and Eumusa bananas were analyzed in this preliminary study. Numerous starch granule morphotypes were present in every sample analyzed. One hundred and nine morphotypes, representing 38 morphotype groups (variants) were described. Of these, several are specific to the samples analyzed and others occurred in more than one sample. They can be used to discriminate between different genera, sections, species and cultivars. Raphides were also numerous in wild Australimusa bananas. Although additional studies are required to determine levels of specificity, this preliminary study shows that starch analysis (and raphide presence and abundance) can be used in a similar way to phytolith analysis in the identification of Musaceae and has extremely good potential as a tool for tracing the prehistory of bananas in the archaeological record.
Article
Against the backdrop of a comprehensive archaeobotanical review, we present linguistic evidence that allows us to develop a model of the dispersal of bananas westwards from New Guinea. This westward dispersal is a window on pre-Austronesian contact between New Guinea and regions to the west. The linguistically-derived model is compatible with archaeobotanical and botanical data. Our interpretation accords with evidence for maritime interactions from the early Holocene in western New Guinea and eastern Indonesia.
Article
Random genomic probes were used to detect restriction fragment length polymorphisms (RFLPs) in 26 accessions of Musa representing eight species from Papua New Guinea (PNG), M. textilis, M. jackeyi and one accession of Ensete. Ninety-eight phylogenetically informative characters were scored and analyzed cladistically and phenetically. Results generally agreed with previous morphology-based phylogenetic analyses. However, the closest wild relative of the edible M. fehi (fe'i banana) appears to be M. lolodensis. Musa angustigemma is sister species with M. boman and M. jackeyi and is distinct from M. peekelii, with which it is often united. Musa boman is unambiguously placed in section Australimusa. The diploid parthenocarpic landraces of section Musa unique to PNG are closely related to, but apparently distinct from, M. acuminata ssp. banksii. The evolution of the fe'i bananas and the M. acuminata-derived diploid landraces of PNG are discussed.