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TRANSCRIPT
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. [email protected]
(bell), seeds and flowers can also be eaten; the leaves are used for cooking, wrapping, serving food and for shel-ter from the sun and rain; fibers 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 fleshy, 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 significant 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 difficult 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 briefly 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 identification and is another step forward in solving the issues surrounding banana dispersal, cul-tivation and domestication, especially in the Pacific/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
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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 difficult 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.
Significantly, 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 afield as Africa and the Pacific. 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-cific 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 Pacific 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 Pacific (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 confirmed record with a positive identification 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 confirmed as banana (Horrocks et al. 2008). Most phytolith records to date have been based on the identification of distinctive volcaniform morphotypes from Musaceae leaves. However, identifica-tion beyond the family level has not been successfully at-tempted in most studies because of the difficulties 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 identified some Musaceae morphotypes in sediments dated to c. 10,000 cal B.P. as Australimusa. However, these identifications 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 confirm 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 fills 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 fills 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 identified to any particular section or spe-cies.
In addition to the Kuk site, Lentfer also confirmed 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.
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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
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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).
HG
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Apart from phytolith studies in New Guinea there are few other published accounts of banana identification 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 identified to genus Musa (Mbida et al. 2001, 2004). Further to the east, at Munsa, Uganda, both Musa and Ensete phytoliths were identified 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-Pacific 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 difficult. The record is derived most-ly from two broad categories of evidence: ecological and geographical. This is mostly reliant on archaeobotanical finds 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 influ-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 Pacific 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 difficult, 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 confirmed if it could be proven that bananas were indeed seedless, and therefore cultivars.
Identification 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 first set of exploratory analyses examined seventeen acces-
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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)
CultivarsEumusa 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 figures.
B
A
sions consisting of Ensete and wild and cultivated Australi-musa and Eumusa Section bananas (Table 1). Twenty-five 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 significantly 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
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Figure 3. 95% confidence 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 significantly smaller craters compared to body length than leaf phytoliths and can be discriminated at α = 0.05 regardless of species derivation.
Figure 4. 95% confidence 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.
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M. p
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lii(s
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1725
2425
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2216
226
25
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ii(s
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Ense
tesp
.(lf
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sete
sp.
(sk)
M. p
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lii(s
d)4B
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B
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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 AttributesCrater 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 significantly 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 significantly greater at α = 0.05 than M. acuminata ssp. banksii and Ensete (Figure 5).
Figure 5. 95% confidence 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 first 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 significantly 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 significantly 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 difficult to differentiate (Figures 7 and 8). Crater width of the wild M. maclayi morphotypes were significantly 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 significant difference at α = 0.05 was between Australimusa section banan-
22 16 22 25 6
40
30
20
10
0N=
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
0N=
species
95% Cl long dimension
M. banksii(sd)
M. peekelii(sd/fr)
Ensete sp.(lf)
Ensete sp.(sk)
M. peekelii(sd)
5
2216
2225
6
40 30 20 10 0 N=
spec
ies
95%
Cll
ong
dim
ensi
on
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
Ense
tesp
.(lf
)En
sete
sp.
(sk)
M. p
eeke
lii(s
d)5
8 6 4 2 0 N=
spec
ies
95%
Clc
rate
r di
men
sion
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
E. g
lauc
um(lf
)E.
gla
ucum
(sk)
M. p
eeke
lii(s
d)4A
1725
2425
7
8 6 4 2 0 N=
spec
ies
95%
Clc
rate
r di
men
sion
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
E. g
lauc
um(lf
)E.
gla
ucum
(sk)
M. p
eeke
lii(s
d)4A
1725
2425
7
8 6 4 2 0 N=
spec
ies
95%
Clc
rate
r di
men
sion
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
E. g
lauc
um(lf
)E.
gla
ucum
(sk)
M. p
eeke
lii(s
d)4A
1725
2425
7
8 6 4 2 0 N=
spec
ies
95%
Clc
rate
r di
men
sion
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
E. g
lauc
um(lf
)E.
gla
ucum
(sk)
M. p
eeke
lii(s
d)4A
1725
2425
7
8 6 4 2 0 N=
spec
ies
95%
Clc
rate
r di
men
sion
M. b
anks
ii(s
d)M
. pee
kelii
(sd/
fr)
E. g
lauc
um(lf
)E.
gla
ucum
(sk)
M. p
eeke
lii(s
d)4A
1725
2425
7
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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
-30 2-4 -2 864
PC 1PC 2
6A
23 1 0 -1 -2 -30
2-4
-28
64
PC 1
PC 2
6A
Musa banksii F. Muell.
Musa schizocarpa SimmondsMusa peekelii Lauterb.Musa paradisiaca L.Musa maclayi F. Muell. ex Mikl.-MaclayMusa fei F. Muell.Musa acuminata CollaEnsete 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
TR
THSHO
RAFHGRX
QBCB
RP MHLO
SQUB MORB TBBB DTXOB
OB
NOIRRT
SPH
STXREGR
PC 1 PC 2
SQT
Rough textureIrregular 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
TR
THSHO
RAFHGRX
QBCB
RP MHLO
SQUB MORB TBBB DTXOB
OB
NOIRRT
SPH
STXREGR
PC 1 PC 2
SQT
Rough textureIrregular rim
6B
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.80.2 -0
.80.
0-0
.6-0
.2-0
.40.
20.
40.
60.
8
RTX
IRR
R
TR
THSH
O
RA
FHG
RX
QB
CB
RP
MH
LO
SQU
BM
OR
BTB
BB
DTX
OB
OB
NO
IRR
TSP
H
STX
REG
R
PC 1
PC 2
SQTR
ough
text
ure
Irreg
ular
rim
6B
A
the greatest variability and could not be differentiated from any other taxon.
Lentfer’s findings broadly concur with those of Ball et al. (2006) and Vrydaghs et al. (2009), which have been con-fined 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 significantly 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 identified 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
TR
THSHO
RAFHGRX
QBCB
RP MHLO
SQUB MORB TBBB DTXOB
OB
NOIRRT
SPH
STXREGR
PC 1 PC 2
SQT
Rough textureIrregular rim
6B
0.8
0.6
0.4
0.0
-0.2
-0.4
-0.6
-0.80.2 -0
.80.
0-0
.6-0
.2-0
.40.
20.
40.
60.
8
RTX
IRR
R
TR
THSH
O
RA
FHG
RX
QB
CB
RP
MH
LO
SQU
BM
OR
BTB
BB
DTX
OB
OB
NO
IRR
TSP
H
STX
REG
R
PC 1
PC 2
SQTR
ough
text
ure
Irreg
ular
rim
6B
B
as. Body lengths of M. maclayi and Fe`i bananas were significantly 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% confidence 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
5N=
species7658 25 25 25
6
25 50 50
95%
Clc
rate
r di
men
sion
7 la
bels
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
95%
Clc
rate
r di
men
sion
7 la
bels
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
8
7658 25 25 25 25 50 50
22
20
18
16
10N=
species
14
12
8titl
es
95%
Cll
ong
dim
ensi
on
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
8titl
es
95%
Cll
ong
dim
ensi
on
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
Figure 8. 95% confidence 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 confirm evidence for cultivation.
Where seed is absent from ar-chaeobotanical assemblages, the problem of identification becomes more difficult 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 50N=species
4.0
3.5
3.0
2.5
2.0
1.5
9titl
es95
% C
lrat
io (l
d/td
)
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
9titl
es95
% C
lrat
io (l
d/td
)
M. b
anks
ii
M. a
cum
inat
a
E. g
lauc
um
M. f
ei
M. m
acla
yi
M. p
arad
isia
ca
M. p
eeke
lii
M. s
chiz
ocar
pa
Figure 9. 95% confidence 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 definitive 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 identification 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/Pacific 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-
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Tabl
e 3.
Lis
t of b
anan
a ac
cess
ions
col
lect
ed fr
om P
apua
New
Gui
nea
in 2
002
for t
he ‘N
ew G
uine
a B
anan
a P
roje
ct’,
with
ass
ocia
ted
ethn
obot
anic
al o
bser
vatio
ns.
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
Nar
i 104
7A
BB
Cap
e G
louc
este
rLo
wla
nds
?N
ari 1
46A
AB
EN
BLo
wla
nds
Luba
Nar
i 186
AAT
Mad
ang
Low
land
sSa
rA
mel
eN
ari
NB
I 10
AB
B?
Low
land
s?
Nar
i 168
AA
BC
entra
l Pro
vinc
eLo
wla
nds
?N
ari
NB
L 20
AB
BB
ouga
invi
lleLo
wla
nds
?
Nar
i O
BB
5A
AA
Mad
ang
Low
land
sLa
kem
Nar
i 171
AB
BM
adan
g (o
rig.
Miln
e B
ay)
Low
land
s?
Nar
i 142
AA
EN
BLo
wla
nds
Tago
mor
Nar
i N
B9
11A
BB
Mor
obe
Low
land
sG
ana
sum
du
Nar
i N
BI 1
8A
AB
Chi
mbu
Hig
hlan
dsK
unam
bo
Nar
i 206
AA
BW
este
rn
Hig
hlan
dsH
ighl
ands
Ruk
umam
b ta
mby
Nar
i 164
AA
Mad
ang
Low
land
sSi
hir
Nar
i O
BN
14
AA
AE
NB
Low
land
sTo
wbe
rne
Nar
i 064
AA
Eas
t Sep
ikLo
wla
nds
?W
NB
1E
nset
e gl
aucu
m
(Rox
b.)
Che
esm
an
Tam
are
Villa
ge
Low
land
s5o 2
7I 0.4
II15
0o 5I 3
.6II
Tapu
puB
akov
iS
eeds
use
d fo
r mak
ing
bead
s.
WN
B2
AA
?Ta
mar
e Vi
llage
Lo
wla
nds
5o 27I 0
.4II
150o 5
I 3.6
IIK
ikiy
ouB
akov
iC
ooke
d an
d al
so e
aten
ripe
.W
NB
3A
BB
Tam
are
Villa
ge
Low
land
s5o 2
7I 0.4
II15
0o 5I 3
.6II
Tuku
ruB
akov
iC
ooke
d an
d al
so e
aten
ripe
.W
NB
4A
BB
?Ta
mar
e Vi
llage
Lo
wla
nds
5o 27I 0
.4II
150o 5
I 3.6
IITa
man
e bu
roB
akov
iC
ooke
d an
d al
so e
aten
ripe
.W
NB
5A
A?
Tam
are
Villa
ge
Low
land
s5o 2
7I 0.4
II15
0o 5I 3
.6II
May
aB
akov
iC
ooki
ng b
anan
a.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths: An update from Papua New Guinea
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259
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
WN
B6
Mus
a te
xtili
s N
éeTa
mar
e Vi
llage
Lo
wla
nds
5o 27I 0
.4II
150o 5
I 3.6
IITu
ain
Pis
inP
seud
oste
m &
she
ath
used
for
mak
ing
rope
. Suc
kers
sol
d to
vi
llage
rs b
y ag
ricul
tura
l sup
ply
stor
es. U
sed
as c
ash
crop
.W
NB
7?
Tam
are
Villa
geLo
wla
nds
5o 27l 0
.4ll
150o 5
I 3.6
IIH
ale
vudi
Bak
ovi
Orn
amen
tal b
anan
a w
ith re
d le
aves
.W
NB
8A
A?
Tam
are
Villa
ge
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Kar
uke
vudi
(K
aruk
e =c
rab’
s ha
nd)
or M
rs. b
anan
a
Bak
ovi
Eat
en ri
pe.
WN
B9
AA
?Ta
mar
e Vi
llage
Lo
wla
nds
5o 27l 0
.4ll
150o 5
l 3.6
llsa
usag
e ba
nana
Eng
lish
Coo
ked
and
also
eat
en ri
pe.
WN
B10
Mus
a ac
umin
ata
ssp.
ban
ksii
(Rid
l.) K
iew
.
Gar
u pl
anta
tion
Low
land
s5o 3
1l 12.
9ll14
9o 58l 2
9.4ll
Wil
bana
naP
isin
Wild
ban
ana
grow
ing
alon
g m
argi
ns o
f sw
amp.
WN
B11
E. g
lauc
umTa
bairi
kau
Low
land
s5o 2
8l 25.
2ll15
0o 27l 2
7.2ll
Vudu
vud
uK
uanu
aC
olle
cted
from
fore
st a
nd g
row
n in
gar
den
with
oth
er b
anan
as.
Nec
klac
es m
ade
from
see
d. S
ap re
d.E
NB
1A
BB
Tavu
i No.
2Lo
wla
nds
4o 08l 4
7.0ll
152o 1
0l 02.
4llYa
wa
Kua
nua
Larg
e se
eds
ofte
n pr
esen
t. Fr
uit
cook
ed a
nd a
lso
eate
n rip
e. F
ruit
also
fed
to p
igs.
Ste
ms
and
leav
es
used
for m
umu.
Fru
it us
ed fo
r m
akin
g an
alc
ohol
ic b
ever
age.
The
rip
e fru
it is
pla
ced
in d
rum
and
afte
r he
atin
g to
rele
ase
juic
es th
e liq
uid
is
ferm
ente
d. It
is d
istil
led
into
a c
lean
dr
um a
nd p
ut in
to s
mal
l bot
tles.
Th
e pr
oces
s ta
kes
two
wee
ks.
EN
B2
AB
BTa
vui N
o. 2
Low
land
s4o 0
8l 47.
0ll15
2o 10l 0
2.4ll
Kal
apua
Kua
nua
Coo
ked
and
also
eat
en ri
pe.
Frui
t fed
to p
igs.
Ste
ms
and
leav
es u
sed
for m
umu.
EN
B3
AB
BTa
vui N
o. 2
Low
land
s4o 0
8l 58.
9ll15
2o 10l 2
7.1ll
Tuku
ruK
uanu
aFr
uit c
ooke
d. M
ale
bud
is a
lso
eate
n.E
NB
4?
Tavu
i No.
2Lo
wla
nds
4o 13l 2
6.3ll
152o 1
0l 27.
1llK
uduk
udu
Kua
nua
Eat
en ri
pe a
nd c
ooke
d.E
NB
5?
Mal
agun
aLo
wla
nds
4o 08l 5
8.9ll
152o 0
8l 59.
0llK
akat
urK
uanu
aE
aten
ripe
.E
NB
6?
Gel
a G
ela
Low
land
s4o 2
4l 38.
0ll15
2o 14l 4
8.6ll
Pok
pok
Kua
nua
Coo
king
ban
ana.
Lea
ves e
asily
torn
and
un
suita
ble
for c
ooki
ng a
nd w
rapp
ing.
Ethnobotany Research & Applications260
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
EN
B7
?G
ela
Gel
aLo
wla
nds
4o 24l 3
6.7ll
152o 1
4l 51.
6llM
akau
Kua
nua
Coo
king
ban
ana.
Lea
ves e
asily
torn
and
un
suita
ble
for c
ooki
ng a
nd w
rapp
ing.
EN
B8
AA
AG
ela
Gel
aLo
wla
nds
4o 24l 3
6.1ll
152o 1
4l 49.
1llK
atur
Kua
nua
Coo
king
ban
ana.
Lea
ves
used
for m
umu.
EN
B9
AA
Gel
a G
ela
Low
land
s4o 2
4l 36.
1ll15
2o 14l 4
9.1ll
Pitu
Kua
nua
Coo
king
ban
ana.
Lea
ves e
asily
torn
and
un
suita
ble
for c
ooki
ng a
nd w
rapp
ing.
EN
B10
AB
BTa
kabu
a M
issi
on
Sta
tion
Low
land
s4o 2
2l 29.
8ll15
2o 13l 3
.1ll
Mak
ala
tuku
ruK
uanu
aE
aten
ripe
. Lea
ves
used
for m
umu.
EN
B11
AB
BM
alak
una
No.
4Lo
wla
nds
4o 23l 6
.0ll
152o 1
2l 18.
3llYa
wa
buka
Kua
nua
Eat
en ri
pe. L
eave
s us
ed fo
r mum
u.E
NB
12B
B?
Taka
bua
Mis
sion
S
tatio
nLo
wla
nds
4o 22l 2
9.8ll
152o 1
3l 3.1
llO
kaok
oK
uanu
aE
aten
ripe
. Lea
ves
used
for m
umu.
EN
B13
Fe`i
Rao
loLo
wla
nds
4o 5l 4
0.5ll
152o 4
l 31.
2llA
uro=
Vuro
Kua
nua
Eat
en ri
pe a
nd c
ooke
d.
EN
B14
Fe`i
Mal
mal
uan
Trai
ning
Col
lege
Low
land
s4o 1
7l 49.
6ll15
2o 1l 4
1.0ll
Vuro
Kua
nua
Eat
en r
ipe
and
cook
ed.
Leav
es u
sed
for m
umu.
EN
B15
AA
L.A
.E.S
. Ker
evat
Low
land
s4o 2
0l 7.0
ll15
2o 1l 4
1.0ll
Gul
umK
uanu
aC
ooki
ng b
anan
a. L
eave
s eas
ily to
rn a
nd
unsu
itabl
e fo
r coo
king
and
wra
ppin
g.E
NB
16Fe
`iL.
A.E
.S. K
erev
atLo
wla
nds
4o 20l 7
.5ll
152o 1
l 48.
2llVu
ro/P
ituK
uanu
aE
aten
ripe
. Lea
ves
used
for m
umu.
EN
B17
AB
?L.
A.E
.S. K
erev
atLo
wla
nds
4o 20l 7
.5ll
152o 1
l 48.
2llTa
upen
Kua
nua
Coo
king
ban
ana.
EN
B18
AA
AL.
A.E
.S. K
erev
atLo
wla
nds
4o 20l 7
.5ll
152o 1
l 48.
2llM
alam
red
Kua
nua
Eat
en ri
pe a
nd c
ooke
d.
EN
B19
AS
?L.
A.E
.S. K
erev
atLo
wla
nds
4o 20l 7
.5ll
152o 1
l 48.
2ll
EN
B20
M. t
extil
isL.
A.E
.S. K
erev
atLo
wla
nds
4o 20l 9
.2ll
152o 1
l 52.
1llP
seud
oste
m a
nd s
heat
h us
ed fo
r m
akin
g ro
pe. S
ucke
rs s
old
by L
.A.E
.S.
to a
gric
ultu
ral s
uppl
y st
ores
.N
I1A
BB
? cf
. Tu
kuru
(K
okop
o)
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
3.1ll
150o 4
7l 54.
6llSu
kuru
Tiga
kC
ooki
ng b
anan
a bu
t als
o ea
ten
ripe.
Le
aves
stro
ng a
nd u
sed
for m
umu,
an
d w
rapp
ing.
Mal
e bu
d al
so e
aten
.N
I2A
A?
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
3.1ll
150o 4
7l 54.
6llPa
pat (
cf.
Gar
amut
in
Kua
nua)
Tiga
kC
ooki
ng b
anan
a - f
ruits
boi
led
and
roas
ted.
Lea
ves
wea
k - u
sed
for c
over
ing
mum
u on
ly.N
I3A
AK
ulan
git V
illag
e,
Kav
ieng
Low
land
s2o 3
3l 43.
1ll15
0o 47l 5
4.6ll
Kik
iyou
Tiga
kC
ooki
ng b
anan
a - f
ruits
boi
led,
fri
ed a
nd ro
aste
d. L
eave
s w
eak
- us
ed fo
r cov
erin
g m
umu
only.
NI4
AB
B?
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
5.5ll
150o 4
7l 54.
1llW
an k
ina,
W
an p
ound
Pis
inC
ooke
d in
mum
u an
d ea
ten
raw
.
NI5
?K
ulan
git V
illag
e,
Kav
ieng
Low
land
s2o 3
3l 45.
5ll15
0o 47l 5
4.1ll
Boa
Tiga
kC
ooke
d in
mum
u an
d ea
ten
raw
and
boi
led.
Sw
eet.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths: An update from Papua New Guinea
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
261
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
NI6
Fe`i?
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
5.5ll
150o 4
7l 54.
1llSu
lukl
amuk
Tiga
kE
aten
ripe
and
coo
ked.
Yel
low
fles
h w
ith p
oorly
form
ed s
eeds
eat
en
by a
nts.
Tur
ns u
rine
yello
w. B
unch
dr
oopi
ng. P
ossi
bly
inte
rmed
iate
form
. Le
aves
stro
ng a
nd u
sed
in m
umu.
NI7
?K
ulan
git V
illag
e,
Kav
ieng
Low
land
s2o 3
3l 45.
5ll15
0o 47l 5
4.1ll
Dar
ipTi
gak
Eat
en r
ipe
and
cook
ed.
Yello
w fl
esh.
Le
aves
not
use
d.N
I8A
AA
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
1.7ll
150o 4
7l 52.
1llPa
lang
Tiga
kE
aten
rip
e an
d co
oked
. Ye
llow
fles
h.
Leav
es n
ot u
sed.
NI9
AA
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
1.7ll
150o 4
7l 52.
1llN
gitin
gsak
aiTi
gak
Coo
king
ban
ana
- bo
iled
and
roas
ted.
Le
aves
no
t us
ed.
Frui
t oc
curs
in
co
ntin
uous
spi
ral a
roun
d ra
chis
.N
I10
AA
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
2.5ll
150o 4
7l 56.
4llU
lung
anTi
gak
Frui
t ro
aste
d or
bo
iled.
Le
aves
no
t us
ed.
NI1
1A
AB
Kul
angi
t Vill
age,
K
avie
ngLo
wla
nds
2o 33l 4
8.4ll
150o 4
7l 56.
1llB
avay
aTi
gak
Frui
t coo
ked
or e
aten
raw
. Lea
ves
used
fo
r wra
ppin
g.N
I12
AA
Mio
m/C
aran
as,
Kav
ieng
Low
land
s2o 3
5l 25.
6ll15
0o 49l 4
1.4ll
Papa
t Wun
gTi
gak
Frui
t roa
sted
or b
oile
d. L
eave
s us
ed fo
r m
umu
and
wra
ppin
g.N
13A
AB
Mio
m/C
aran
as,
Kav
ieng
Low
land
s2o 3
5l 25.
6ll15
0o 49l 4
1.4ll
Whi
te b
avay
aE
nglis
h/Ti
gak
Mos
tly ro
aste
d.
NI1
4Fe
`iN
amas
alan
gLo
wla
nds
3o 2l 4
5.4ll
151o 2
4l 57.
1llU
tafa
nM
alik
Coo
ked
and
eate
n ra
w -
swee
t. S
ap
used
as
a dy
e fo
r mal
anga
n m
asks
.N
I15
?Le
let
<100
0m3o 1
6l 20.
7ll15
1o 55l 3
9.0ll
Siam
an/
Ger
man
Man
dak
Eat
en ri
pe.
NI1
6Fe
`iK
alua
n, L
elet
<100
0m3o 1
9l 28.
8ll15
1o 54l 2
2.9ll
Man
ui/
Nam
nam
Man
ui
(Man
us
Isla
nd)/
Man
dak
Eat
en ri
pe o
r coo
ked.
Lea
ves
used
for
mum
u an
d w
rapp
ing.
Bun
ch d
roop
ing
but a
t an
angl
e an
d al
mos
t hor
izon
tal.
NI1
7Fe
`iK
alua
n, L
elet
<100
0m3o 1
7l 58.
3ll15
1o 54l 4
8.7ll
Losk
auk
Man
dak
Eat
en ri
pe o
r coo
ked.
Lea
ves
used
for
mum
u an
d w
rapp
ing.
Bun
ch d
roop
ing
but a
t an
angl
e an
d al
mos
t hor
izon
tal.
NI1
8Fe
`iK
abag
ong,
Lel
et<1
000m
3o 16l 5
5.4ll
151o 5
5l 30.
2llLo
skau
kM
anda
kC
ooki
ng b
anan
a. F
lesh
sul
fur-
yello
w.
NI1
9M
usa
mac
layi
F.
Mue
ll. e
x M
ikl.-
Mac
lay
ssp.
mac
layi
A
rg. v
ar.
nam
atan
i Arg
.
Kar
u<1
000m
3o 29l 5
2.7ll
152o 1
1l 46.
9llA
gigi
lai
Bar
okW
ild. S
aid
not t
o be
eat
en b
ut s
wee
t br
ight
yel
low
fles
h su
rrou
nds
seed
s.
Leav
es u
sed
for w
rapp
ing
and
mum
u.
Ethnobotany Research & Applications262
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
NI2
0M
. mac
layi
? Fe
`i?K
aru
<100
0m3o 3
0l 20.
3ll15
2o 12l 2
1.2ll
Bar
okW
ild.
Tall,
rob
ust
bana
na w
ith s
hiny
ps
eudo
stem
. N
ot
fruiti
ng
at
time
of
colle
ctio
n.N
I21
M. m
acla
yi
ssp.
mac
layi
va
r. na
mat
ani
Kar
u<1
000m
5o 27l 0
.4ll
150o 5
l 3.6
llA
gigi
lai
Bar
okW
ild. S
aid
not t
o be
eat
en b
ut s
wee
t br
ight
yel
low
fles
h su
rrou
nds
seed
s.
Leav
es u
sed
for w
rapp
ing
and
mum
u.N
I22
Mus
a pe
ekel
ii La
uter
b. s
sp.
peek
elii
Arg
.
Ros
irik
<100
0m5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Waw
aS
okiri
Wild
robu
st b
anan
a. B
unch
ere
ct a
t firs
t th
en d
roop
ing.
Fru
it co
oked
or r
oast
ed.
Leav
es u
sed
to c
over
mum
u.N
I23
M. p
eeke
lii
ssp.
pee
kelii
Ros
irik
<100
0m5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Waw
aS
okiri
Wild
robu
st b
anan
a. B
unch
ere
ct a
t firs
t th
en d
roop
ing.
Fru
it co
oked
or r
oast
ed.
Leav
es u
sed
to c
over
mum
u.W
H1
Mus
a in
gens
S
imm
onds
Tuba
ng, K
auil
2000
m5o 2
7l 0.4
ll15
0o5l 3
.6ll
Wul
uk d
onu
Jiw
aka
Wild
tal
l ba
nana
gro
win
g to
16m
to
18m
hei
ght
at 2
000m
in
cree
k gu
lly.
Frui
t with
yel
low
fles
h an
d la
rge
seed
s,
gree
n tu
rnin
g ye
llow
whe
n rip
e. C
uscu
s,
parr
ots
and
flyin
g fo
x ea
t fru
it. L
eave
s us
ed f
or m
akin
g te
mpo
rary
she
lters
an
d fo
r cov
erin
g m
umu.
W
H2
M. i
ngen
sTu
bang
, Kau
il20
00m
5o 27l 0
.4ll
150o 5
l 3.6
llW
uluk
don
uJi
wak
aW
ild ta
ll ba
nana
gro
win
g to
16m
to 1
8m
heig
ht a
t 200
0m a
ltitu
de in
cre
ek g
ully.
Fr
uit w
ith y
ello
w fl
esh
and
larg
e se
eds.
Le
aves
us
ed
for
mak
ing
tem
pora
ry
shel
ters
and
for c
over
ing
mum
u.
WH
3M
. ing
ens
Tuba
ng, K
auil
2000
m5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Wul
uk d
onu
Jiw
aka
As
abov
e.W
H4
Fe`i
Bal
ga, K
uk16
30m
5o 27l 0
.4ll
150o 5
l 3.6
llR
ua m
ema
(rua
= b
anan
a,
mem
a =
red)
Coo
king
ban
ana,
roa
sted
and
boi
led
- pr
efer
ably
roas
ted.
WH
5M
usa
schi
zoca
rpa
Sim
mon
ds
Bai
yer Z
oo11
77m
5o 27l 0
.4ll
150o 5
l 3.6
ll?
?W
ild b
anan
a gr
owin
g in
rai
nfor
est w
ith
ging
ers,
fern
s, R
ubia
ceae
shr
ubs,
tree
fe
rns
and
wild
taro
in ra
info
rest
in s
hade
un
der c
anop
y.E
S1
AAT
?R
anim
bo, H
awai
nLo
wla
nds
5o 27l 0
.4ll
150o 5
l 3.6
llYe
sing
(=w
ild
bana
na)
Min
oTa
ll ro
bust
ban
ana
with
edi
ble
fruit.
Fe
ral r
athe
r tha
n w
ild. F
ruit
eate
n w
hen
raw
or c
ooke
d. L
eave
s us
ed fo
r mum
u an
d w
rapp
ing.
Fru
it al
so fe
d to
pig
s.
Lentfer - Tracing Domestication and Cultivation of Bananas from Phytoliths: An update from Papua New Guinea
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
263
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
ES
2A
A?
CC
RI s
tatio
n,
Haw
ain
Low
land
s5o 3
1l 12.
9ll14
9o 58l 2
9.4ll
Yesi
ng (=
wild
ba
nana
)M
ino
Frui
t gr
een
turn
ing
yello
w w
hen
ripe.
Le
aves
use
d as
dec
orat
ion
in s
ings
ing
cele
brat
ions
afte
r he
atin
g on
fire
to
mak
e pl
iabl
e.E
S3
M.
schi
zoca
rpa
CC
RI s
tatio
n,
Haw
ain
Low
land
s5o 2
8l 25.
2ll15
0o 27l 2
7.2ll
Yesi
ng (=
wild
ba
nana
)M
ino
Wild
se
eded
ba
nana
w
ith
yello
wis
h sa
p.E
S4
M.
schi
zoca
rpa
Ran
imbo
, Haw
ain
Low
land
s4o 0
8l 47.
0ll15
2o 10l 0
2.4ll
Yesi
ng (=
wild
ba
nana
)M
ino
Wild
see
ded
bana
na w
ith c
lear
sap
. Le
aves
use
d fo
r mum
u, w
rapp
ing
food
an
d fo
r sin
gsin
g de
cora
tion
as w
ell a
s ro
ofs
for b
ush
hous
es.
ES
5Fe
`iK
oiki
n Vi
llage
, W
ewak
Low
land
s4o 0
8l 47.
0ll15
2o 10l 0
2.4ll
Taw
eyaw
aM
ino
Cul
tivat
ed
bana
na
with
re
d sa
p.
Frui
t la
rge
4-5c
m d
iam
eter
, 10
-23c
m
long
w
ith
bottl
e-ne
cked
ap
ex
and
pron
ounc
ed ri
dgin
g. M
ostly
coo
ked.
ES
6M
. acu
min
ata
ssp.
ban
ksii
Koi
kin
Villa
ge,
Wew
akLo
wla
nds
4o 08l 5
8.9ll
152o 1
0l 27.
1llYe
sing
(=w
ild
bana
na)
Min
oW
ild s
eede
d ba
nana
with
cle
ar s
ap.
Frui
t tu
rns
yello
wis
h w
hen
ripe.
Not
ea
ten.
Sai
d to
ofte
n co
loni
ze la
ndsl
ides
w
ith
ging
ers
and
gras
ses.
S
peci
es
com
mon
alo
ng S
epik
Hig
hway
tow
ards
M
aprik
. Ofte
n gr
ows
in a
ssoc
iatio
n w
ith
M. s
chiz
ocar
pa. T
his
spec
imen
gro
win
g on
wel
l-dra
ined
loa
m a
long
roa
dsid
e ne
xt to
gar
dens
and
clo
se to
reg
row
th
fore
st. O
ne p
lant
onl
y.E
S7
AB
BT?
Japa
raga
1,
Map
rik R
oad
<100
0m4o 1
3l 26.
3ll15
2o 10l 2
7.1ll
Gia
nt k
alap
uaP
isin
Coo
king
ba
nana
. Le
aves
us
ed
for
wra
ppin
g an
d m
umu.
ES
8A
AJa
para
ga 1
, M
aprik
Roa
d<1
000m
4o 08l 5
8.9ll
152o 0
8l 59.
0llSa
lam
ua,
pisa
ng
jari
buay
a (c
roco
dile
fin
gers
)
?, M
ino
Frui
t eat
en ri
pe o
r coo
ked.
Lea
ves
used
fo
r m
umu
and
drie
d le
aves
drie
d an
d us
ed fo
r mak
ing
ciga
rette
s. C
lear
sap
.
ES
9A
AJa
para
ga 1
, M
aprik
Roa
d<1
000m
4o 24l 3
8.0ll
152o 1
4l 48.
6llH
eife
li’ T
o`o
Min
o,
Mad
ang
lang
uage
?
Frui
t ea
ten
ripe
or
cook
ed.
Leav
es
used
for m
umu.
ES
10M
. acu
min
ata
x sc
hizo
carp
aJa
para
ga 1
, N
agam
Riv
er<1
000m
4o 24l 3
6.7ll
152o 1
4l 51.
6llYe
sing
(=w
ild
bana
na)
Min
oW
ild b
anan
a gr
owin
g on
san
dy r
iver
ba
nk. S
ap c
lear
. Fru
it no
t eat
en.
ES
11M
. acu
min
ata
ssp.
ban
ksii
Japa
raga
1,
Nag
am R
iver
<100
0m4o 2
4l 36.
1ll15
2o 14l 4
9.1ll
Yesi
ng (=
wild
ba
nana
)M
ino
Wild
ban
ana
grow
ing
in s
hady
regr
owth
fo
rest
with
mos
s un
ders
tory
.
Ethnobotany Research & Applications264
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
M1
Fe`i
Mot
onau
/Mid
iba,
M
adan
gcl
4o 24l 3
6.1ll
152o 1
4l 49.
1llW
ahin
Ari
Cul
tivat
ed b
anan
a w
ith re
d sa
p gr
owin
g on
wel
l-dra
ined
red
/bro
wn
loam
. Fr
uit
larg
e w
ith
pron
ounc
ed
ridgi
ng
and
sulfu
r-ye
llow
fles
h. M
ostly
coo
ked.
Fe`
i ba
nana
s on
ce c
omm
on i
n th
e di
stric
t bu
t sai
d to
be
grad
ually
dyi
ng o
ut, o
ften
surv
ivin
g in
aba
ndon
ed g
arde
n se
tting
s.
Not
able
Fe`
i ba
nana
s ar
e re
ferr
ed t
o as
“yel
o pi
spis
” in
Pis
in b
ecau
se u
rine
turn
s ye
llow
afte
r the
frui
t is
eate
n. F
ruit
cook
ed a
nd u
sed
for c
olor
ing
othe
r foo
d st
uffs
and
enh
anci
ng fl
avou
rs. R
ed s
ap
used
by
child
ren
for m
akin
g st
enci
ls o
n cl
othi
ng.
M2
AA
Ste
war
t Res
earc
h S
tatio
n, M
adan
gLo
wla
nds
4o 22l 2
9.8ll
152o 1
3l 3.1
llM
anam
eg
(red
)A
mel
eS
mal
l cu
ltiva
ted
bana
na.
Coo
king
ba
nana
refe
rred
to a
s pl
anta
in a
lthou
gh
stric
tly s
peak
ing
it is
n’t.
Leav
es u
sed
for m
umu.
M3
AA
?S
tew
art R
esea
rch
Sta
tion,
Mad
ang
Low
land
s4o 2
3l 6.0
ll15
2o 12l 1
8.3ll
Kek
iau
Kua
nua
Coo
king
ba
nana
, bo
iled
or
roas
ted.
Le
aves
use
d fo
r m
umu
and
wra
ppin
g fo
od.
M4
M.
schi
zoca
rpa
Bai
tata
Roa
d,
Mad
ang
Low
land
s4o 2
2l 29.
8ll15
2o 13l 3
.1ll
Mar
orE
mW
ild b
anan
a gr
owin
g on
poo
rly d
rain
ed
limes
tone
soi
l in
regr
owth
fore
st a
long
ro
adsi
de. G
row
ing
near
Mus
a pe
ekel
ii ss
p. a
ngus
tigem
ma.
(NB
. Site
use
d fo
r m
akin
g th
e fil
m “
Rob
inso
n C
ruso
e”).
Frui
t se
lf-pe
els
to
give
a
star
-like
ap
pear
ance
to
the
apex
. S
kin
gree
n w
hen
ripe.
Fle
sh w
hite
. Lea
ves
used
to
mak
e te
mpo
rary
bus
h sh
elte
rs.
M5
M. p
eeke
lii
ssp.
an
gust
igem
ma
Bai
tata
Roa
d,
Mad
ang
Low
land
s4o 5
l 40.
5ll15
2o 4l 3
1.2ll
Dor
Em
Tall r
obus
t wild
ban
ana
grow
ing
near
M.
schi
zoca
rpa.
Com
mon
in
wel
l-dra
ined
lo
catio
ns
on
cora
l/lim
esto
ne
soils
in
re
grow
th
fore
st.
Bun
ch
droo
ping
, fa
lling
stra
ight
. Fl
ower
s ea
ten
- m
ixed
w
ith
salt.
Yo
ung
pseu
dost
em
also
ea
ten
oppo
rtuni
stic
ally.
Fl
ower
s an
d ps
eudo
stem
use
d to
gar
nish
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
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
M6
M. a
cum
inat
a ss
p. b
anks
iiB
aita
ta V
illag
e,
Mad
ang
Low
land
s4o 1
7l 49.
6ll15
2o 1l 4
1.0ll
Tilja
Em
Wild
ban
ana
grow
ing
in a
ssoc
iatio
n w
ith M
. pe
ekel
ii ss
p. a
ngus
tigem
ma
on c
oral
/lim
esto
ne s
oils
at
edge
s of
re
grow
th fo
rest
.M
7M
. acu
min
ata
ssp.
ban
ksii
Bar
um ju
nctio
n,
Mad
ang
Low
land
s4o 2
0l 7.0
ll15
2o 1l 4
1.0ll
Mab
ul k
ulal
(m
abul
=wild
: ku
lal=
ined
ible
, le
ft to
dry
)
Kei
nW
ild
bana
na
in
mod
erat
ely
dens
e st
and
grow
ing
on
wel
l-dra
ined
so
il in
reg
row
th f
ores
t al
ong
road
side
. In
cl
ose
asso
ciat
ion
with
M.
schi
zoca
rpa
and
AS
hyb
rids.
M8
AS
Bar
um ju
nctio
n,
Mad
ang
Low
land
s4o 2
0l 7.5
ll15
2o 1l 4
8.2ll
Mab
ul ju
s (ju
s= w
ater
sn
ail b
ecau
se
the
snai
l has
a
poin
ted
shel
l and
the
bana
na h
as a
lo
ng p
oint
ed
apex
).
Kei
nW
ild b
anan
a gr
owin
g on
wel
l-dra
ined
so
il in
reg
row
th f
ores
t al
ong
road
side
. G
row
ing
amon
gst
mix
ed
stan
d of
M
. ac
umin
ata,
M
. sc
hizo
carp
a an
d hy
brid
s.
M9
M.
schi
zoca
rpa
Bar
um ju
nctio
n,
Mad
ang
Low
land
s4o 2
0l 7.5
ll15
2o 1l 4
8.2ll
Hiru
nag
(hiru
=gin
ger;
nag=
son)
Kei
nW
ild b
anan
a gr
owin
g on
wel
l-dra
ined
so
il in
reg
row
th f
ores
t al
ong
road
side
. G
row
ing
amon
gst
mix
ed s
tand
of
M.
acum
inat
a an
d M
. sch
izoc
arpa
and
AS
hy
brid
s.M
10A
AJo
btou
, Mad
ang
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Bag
uiK
ein
Cul
tivat
ed
spec
ies
very
m
uch
like
the
wild
M.
acum
inat
a ss
p. b
anks
ii in
ap
pear
ance
. Mal
e bu
d gr
een.
MB
1E
. gla
ucum
Nau
ra V
illag
eLo
wla
nds
5o 27l 0
.4ll
150o 5
l 3.6
llG
udu
gudu
Hai
gwai
Gro
win
g in
ne
glec
ted
coco
nut
plan
tatio
n.
See
ds
used
to
m
ake
neck
lace
s.M
B2
M.
schi
zoca
rpa
Nau
ra V
illag
eLo
wla
nds
5o 27l 0
.4ll
150o 5
l 3.6
llLa
u m
oi m
oi
(lau=
ined
ible
; m
oi=b
anan
a)
Hai
gwai
Wild
ba
nana
gr
owin
g in
ov
ergr
own
gard
ens.
Lea
ves
som
etim
es u
sed
for
cove
ring
food
. S
peci
es a
lso
foun
d at
G
omila
Vill
age.
MB
3M
. mac
layi
ss
p. m
acla
yi?
Gom
ila V
illag
e,
Ahi
oma
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Bih
ibih
iya
(bih
ibih
i= w
ild
bana
na; b
ihi
= cu
ltiva
ted
bana
na)
Taw
ala
Wild
ban
ana
with
dar
k, m
ilky
purp
le-r
ed
sap
and
erec
t bun
ch. F
ruit
with
sul
fur-
yello
w fl
esh.
Gro
win
g in
coo
l, sh
elte
red
gully
in r
egro
wth
fore
st. A
noth
er s
mal
l st
and
foun
d gr
owin
g ne
arby
on
rock
y ta
lis s
lope
bes
ide
cree
k in
old
gar
den
bord
erin
g re
grow
th fo
rest
.
Ethnobotany Research & Applications266
www.ethnobotanyjournal.org/vol7/i1547-3465-07-247.pdf
Cod
eTy
peLo
catio
nA
ltitu
de
(m)
Latit
ude
SLo
ngitu
de
EVe
rnac
ular
N
ame
Lang
uage
Not
es
MB
4M
. mac
layi
ss
p. m
acla
yi?
Gom
ila V
illag
e,
Ahi
oma
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Bih
ibih
iya
(bih
ibih
i= w
ild
bana
na; b
ihi
= cu
ltiva
ted
bana
na)
Taw
ala
One
of
stan
d of
wild
ban
anas
with
da
rk,
milk
y pu
rple
-red
sa
p gr
owin
g on
tal
is s
lope
in o
ld g
arde
n al
ongs
ide
cree
k. S
mal
l sta
nds
of s
peci
es fo
und
in
dens
e re
grow
th fo
rest
on
stee
p ba
nks
bord
erin
g cr
eek.
M
B5
M. m
acla
yi
ssp.
mac
layi
?G
omila
Vill
age,
A
hiom
aLo
wla
nds
5o 27l 0
.4ll
150o 5
l 3.6
llB
ihib
ihiy
a (b
ihib
ihi=
wild
ba
nana
; bih
i =
culti
vate
d ba
nana
)
Taw
ala
Wild
ban
ana
with
pin
k sa
p an
d er
ect
bunc
h, ti
ght b
ut la
x at
pro
xim
al e
nd.
Frui
t with
sul
fur-
yello
w fl
esh.
Gro
win
g in
gar
dens
on
rais
ed s
andy
gra
vel
depo
sits
bor
dere
d by
cre
ek c
hann
els.
MB
6M
. mac
layi
ss
p. m
acla
yi?
Dad
uwe
area
, N
orth
east
C
oast
Roa
d
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Bih
ibih
iya
(bih
ibih
i= w
ild
bana
na; b
ihi
= cu
ltiva
ted
bana
na)
Taw
ala
Wild
ban
ana
in s
mal
l sta
nd o
f ban
anas
gr
owin
g on
w
ell-d
rain
ed
soil
alon
g ro
adsi
de in
tall
gras
slan
d.
MB
7M
. mac
layi
ss
p. m
acla
yi?
Dad
uwe
area
, N
orth
east
C
oast
Roa
d
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Bih
ibih
iya
(bih
ibih
i= w
ild
bana
na; b
ihi
= cu
ltiva
ted
bana
na)
Taw
ala
Wild
ban
ana
grow
ing
in s
mal
l sca
ttere
d st
and o
n wel
l-dra
ined
soil a
long
road
side
in
ta
ll gr
assl
and.
S
ome
spec
imen
s fe
atur
e re
tent
ion
of b
ract
s on
the
rach
is
whi
le
othe
rs
shed
br
acts
. N
otab
ly
brac
t ret
entio
n is
a c
hara
cter
istic
of M
. m
acla
yi s
sp.
ailu
lui
Arg
. bu
t in
oth
er
resp
ects
the
spec
imen
s fit
mor
e cl
osel
y w
ith M
. mac
layi
ssp
. mac
layi
acc
ordi
ng
to A
rgen
t’s k
ey.
MB
0M
. mac
layi
ss
p. m
acla
yi?
Kui
aro
Villa
ge,
Chi
na S
trait
Low
land
s5o 2
7l 0.4
ll15
0o 5l 3
.6ll
Spe
cim
en fo
und
on h
illsi
de in
regr
owth
fo
rest
rec
ently
cle
ared
for
gar
deni
ng.
Als
o no
ted
to b
e gr
owin
g in
fore
st a
reas
in
land
from
the
coas
t.M
B0
M. m
acla
yi
ssp.
mac
layi
?Tu
e Vi
llage
Low
land
s5o 3
1l 12.
9ll14
9o 58l 2
9.4ll
Lau
moi
moi
(la
u=fa
lse;
m
oi=b
anan
a)
Dai
amon
iS
aid
to b
e gr
owin
g in
bus
h in
cre
ek
catc
hmen
t. N
ot c
olle
cted
.
MB
8A
ATu
e Vi
llage
Low
land
s5o 2
8l 25.
2ll15
0o 27l 2
7.2ll
Kok
oluy
uD
aiam
oni
Cul
tivat
ed b
anan
a. F
ruit
eate
n co
oked
or
ripe
.M
B9
AA
? A
S?
Tue
Villa
geLo
wla
nds
4o 08l 4
7.0ll
152o 1
0l 02.
4llM
asi m
asi
Dai
amon
iC
ultiv
ated
ban
ana.
Fru
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Table 4. Accessions selected for the ‘New Guinea Banana Project’ analyses.
Section Number analyzed
Accession code
Wild speciesEumusa Musa acuminata ssp. banksii 5 ES11, ES6, M7, M6, ENB10
Musa schizocarpa 5 ES3, WH5, MB2, M9, ES4Musa acuminata ssp. banksii x schizocarpa 2 M8, ES10
Australimusa Musa peekelii 2 M5, NI22Musa maclayi 5 NI21, MB7, MB4, MB3, MB5Musa textilis 2 ENB20, WNB6
Ingentimusa Musa ingens 3 WH1, WH2, WH3N/A Ensete glaucum 4 WNB1, WNB11, MB1, MB1/2
Cultivars*Eumusa AA 5 NI12, ENB13, Nari064,
Nari164, MB8AAA 5 NI8, NariOBB5, NariNB420,
NariOBN14, ENB18BB? 1 ENB12AAB 5 NI13, Nari206, Nari146, NI11, NB1ABB 5 Nari1047, NariNBG11, NariNBI10,
Nari171, NariNBL20AB 1 ENB17
Australimusa TT(Fe`i) 5 WH4, ES5, ENB13, M1, ENB14Eumusa 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 AttributeBase, 3D shape tabular, blocky, spherical( globose), platy (planar)Base, 2D shape round, oval, square, rectangle, quadrilateral, boat (oblong), irregularBody 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, irregularRim present, absentRim shape regular, irregular, skirt (crenate)Sides straight, convex, concave, straight/concave, concave/convex, straight/convexBody texture psilate, granulate, verrucate, nodulose, tuberculate, psilate/verrucate, psilate/granulate,
granulate/verrucateBase ornamentation absent, short (<1.25µm), medium (1.25-2.5µm), long (>2.5µm), long/tuberculate/
dendritic, short/medium, medium/long
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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 difficult 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 identification 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, identification 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 field 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 Pacific/Papua New Guinea region.
Acknowledgements
I am most grateful to the Pacific 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 field collection and the PNG government officials 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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