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Crop Protection 26 (2007) 12191225


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Effects of alternative weed management strategies on Commelina diffusa


Burm. infestations in Fairtrade banana (Musa spp.) in St. Vincent and
the Grenadines
Wendy Ann P. Isaaca,, Richard A.I. Brathwaitea, Jane E. Cohenb, Isaac Bekelea
a
Department of Food Production, Faculty of Science and Agriculture, The University of the West Indies, St. Augustine, Trinidad and Tobago
b
Department of Life Sciences, Faculty of Pure and Applied Sciences, The University of the West Indies, Mona, Jamaica

Received 26 October 2006; accepted 27 October 2006

Abstract

Alternatives to herbicides are needed for weed management in banana cultivated under the Fairtrade system in the Windward Islands.
Field trials were conducted in established banana orchards under irrigated and non-irrigated regimes in the rainy and dry seasons,
20032004, to evaluate the effects of selected treatments on the weed composition response in elds previously infested with Commelina
diffusa Burm. The treatments consisted of three leguminous live mulches (Desmodium heterocarpon (L.) DC., Mucuna pruriens Bak. and
Arachis pintoi Krap. & Greg.), three non-living mulches (coffee hulls, banana mulch and clear plastic mulch), two herbicides (fomesafen
and glufosinate-ammonium) and two organic treatments (corn gluten meal and concentrated vinegar and acetic acid). Treatments
signicantly shifted the species composition away from Commelina diffusa to annual weed species such as Spermacoce latifolia Aubl. and
Cleome aculeate L. Banana mulch treatment provided the best control by signicantly reducing weed biomass and weed species
composition and was followed by Fomesafen, coffee hulls, clear plastic mulch and Desmodium heterocarpon. This study indicates further
research to rene mulched based weed management systems is needed in banana orchards.
r 2006 Elsevier Ltd. All rights reserved.

Keywords: Commelina diffusa; Fairtrade banana; Mulch; Weed composition; Weed biomass

1. Introduction need for more sustainable production systems, which use


fewer or no chemicals, and, in particular, exclude the use of
The banana (Musa spp.) is the mainstay of the traditional herbicides.
agricultural and rural-based community life of St. Vincent Commelina diffusa Burm. is an invasive weed specie that
and the Grenadines in the Windward Islands and accounts has become an aggressive weed in established banana elds
for 10% of the total exports. Bananas are primarily grown throughout St. Vincent and the Grenadines. The weed was
on small farms averaging two hectares in size in hilly areas, once encouraged as a ground cover to reduce soil erosion
owned and worked by over 1000 local family farmers. (Edmunds, 1971) but has since been rated undesirable,
Many of these farmers produce their banana using the being a host of the reniformis nematode, Rotylenchulus
Fairtrade market arrangement of the UK and Europe. reniformis (Robinson et al., 1997), the banana lesion
Fairtrade is a means of assisting small-scale and other nematode, Pratylenchus goodeyi, the burrowing nematode,
disadvantaged producers in developing countries to im- Radopholus similis (Queneherve et al., 2006) and the soil
prove their quality of life by providing a more protable borne fungus, Fusarium oxysporum (Waite and Dunlap,
and stable trade relationship. Fairtrade also emphasizes the 1953) which causes Fusarium wilt. These nematodes all
contribute to signicant reduction in banana production,
Corresponding author. Tel.: +1 868 662 2002x3323; particularly Radopholus similis which may reduce yields by
fax: +1 868 645 0479. more than 50% and decrease the productive life of banana
E-mail address: wisaac@fsa.uwi.tt (W.A.P. Isaac). elds (Queneherve et al., 2006).

0261-2194/$ - see front matter r 2006 Elsevier Ltd. All rights reserved.
doi:10.1016/j.cropro.2006.10.019
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1220 W.A.P. Isaac et al. / Crop Protection 26 (2007) 12191225

Commelina diffusa has gained noxious weed status for There is a paucity of information on the management of
other reasons too. Banana growers in the Windward weeds, including C. diffusa, by other means than herbi-
Islands, since the early 1970s have relied primarily upon cides. Banana growers in the Windward Islands urgently
herbicides as 2,4-D, paraquat and glyphosate (Feakin, need to nd alternatives to herbicides in order to sustain
1971; Hammerton, 1981). Heavy reliance and continuous their export capacity and remain Fairtrade compliant. The
use of these synthetic herbicides have resulted in a number objectives of this study were (i) to evaluate alternative
of problems, including the development of herbicide strategies for the management of C. diffusa and other weed
resistant biotypes of C. diffusa. Since the introduction of species in Fairtrade banana, and (ii) to evaluate the weed
Fairtrade, over 5 years ago, with its prohibition on the use composition response to these selected treatments.
of herbicides, farmers have controlled weeds manually,
using the cutlass or using the weed whacker. Use of the 2. Materials and methods
cutlass and weed whacker has intensied the problem of C.
diffusa by exacerbating the spread of its propagules 2.1. Experimental site and preparation
(Richard Brathwaite, Pers. Comm. December, 2003). These
Islands are characterized by hilly topography and a climate Field trials were conducted in 2003 and 2004 (rainy and
with ideal moisture conditions for the proliferation of C. dry seasons) in established banana elds at four sites in St.
diffusa. Because many of the banana plantations have been Vincent. They were conducted under irrigated and non-
farmed for several years with virtually no crop rotations or irrigated regimes on two of the major soil types of St.
tillage practices further stabilization of C. diffusa popula- Vincent. Site I (non-irrigated/rainy season) was a (Greggs
tions has occurred. loam and clay loam (pH 5.8) with a classication of high
C. diffusa is a perennial, shade tolerant monocotyledo- level yellow earth which is predisposed to severe erosion,
nous creeper propagated mainly by seeds or asexually by and had a slope of over 301 (Watson et al., 1958). Site II
stem sections that readily root at the nodes and produce (irrigated/rainy season) (pH 5.2), Site III (irrigated/dry
autonomous, erect shoots (Fournet and Hammerton, 1991; season) (pH 4.42) and Site IV (non-irrigated/dry season)
Myint, 1994; Wilson, 1981). Its vigorous growth habit (pH 5.2) were Soufriere cindery gravely sandy loam (recent
allows C. diffusa to form dense, pure stands that can volcanic ash), predisposed to slight erosion and had a slope
compete easily with low growing crops smothering them of 051 (Watson et al., 1958). Trials I and II were
(Holm et al., 1977). Wilson (1981) reports, in addition, that conducted during the rainy season in 2003 and trials III
the stems of C. diffusa have a high moisture content and and IV during the dry season in 2004. All four sites had
once it is well rooted a plant can survive for long periods similar cropping histories of being under banana for more
without moisture. This is evident in young banana than 5 years and with the mat spacing being 2.2 m  2.2 m.
plantations in St. Vincent where much of the shoot systems Prior to the application of the application of treatments,
dries out when exposed to sun during the dry season. each site was clean weeded and all the weed stubble
However, at the onset of rains and when the canopy of the removed. C. diffusa was the dominant weed specie found at
banana closes, the weed re-establishes rapidly from the the experimental sites before trials were established.
running parts of the shoots. The average monthly precipitation during the experi-
Because of its vigorous growth habit, which allows the C. ment is given in Fig. 1.
diffusa to form dense pure stands, it can smother low
growing crops such as vegetables, pulses and cereals by 2.2. Treatments
smothering them (Holm et al. 1977). C. diffusa is thought
to be very competitive in established banana elds The experimental design was a randomized complete
although little research has been done on such competition block design with two replicates at each site. Treatments
or on its allelopathic potential. Burns (2004) reported that were arranged under banana plants in 4.5 m  4.5 m plots.
another invasive species, C. benghalensis L. had higher The experiment consisted of (1) two herbicide treatments:
growth rates at high nutrient availability and water (i) glufosinate-ammonium and (ii) fomesafen which were
availability when compared to a related non-invasive (but applied at early post-emergence (at the weed 35 leaf
alien) congener, C. bracteosa Hassk. Interestingly, Shcher- stage). These herbicides were applied with a backpack
bakova (1974) reported severe stunting in C. diffusa caused sprayer which delivered 269-L/ha at kPa pressure using a
by high nitrogen and Urich et al. (2003) reported altered fan-nozzle (TJ-8002). (2) Three non-living mulch treat-
growth and physiological characteristics for different C. ments: (i) banana mulch (traditional practice) applied at a
erecta L. clones with increased phosphorus supply. Weeds depth of 56 cm using fully green and senescing leaves, (ii)
in general can severely affect the growth of young banana coffee hulls applied at a depth of 35 cm and (iii) clear
plantations (Seeyave and Phillips, 1970). This has been plastic mulch using high-density, transparent polyethylene
conrmed by work done since the 1960s in the West Indies tarp at 0.5 mils (50 gauge) thick for a 6-week period. (3)
with the main effect of weed competition on banana plants Three live mulch treatments: (i) Arachis pintoi (wild
being to delay maturity (Kasasian and Seeyave, 1968; peanuts) planted by seed and stem cuttings drilled into
Simmonds, 1959). the soil in rows 16 cm apart with 5 seeds per hole;
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600 total dry mass (DM) recorded. Weed species were identied
from a publication by Fournet and Hammerton (1991) and
conrmation of all weed species was made by the National
500 Herbarium of Trinidad and Tobago.

2.4. Analysis of data


400
Data were subjected to analyses of variance (ANOVA)
Rainfall (mm)

using SPSS (Version 11.0 for windows) with a signicance


300 level of 0.05. Means were separated using least signi-
cance differences (LSD) also at the 0.05 signicance level.
Values for the WCE score were converted to percentages
200
for ANOVA. Analyses were performed on weed counts at
21, 35 and 49 DAT, weed scores at 63 DAT and weed
species biomass at 112 DAT to test the effects of the
treatments.
100

3. Results and discussion


0
A S O N D J F M A M J 3.1. Influence of the weed management treatments on overall
MONTH weed density and weed control efficiency
Rabacca Data Dumbarton Data Long Term Means
All treatments reduced weed density compared to the
Fig. 1. Rainfall/precipitation pattern for the AugustJune experimental UWC up to 49 DAT (Table 1). At 21 DAT, the herbicide
period, 20032004 and the long term means (LTM) from Rabacca (Sites and non-living mulch treatments were as effective at
IIIV) and Dumbarton Field Stations (Site I). suppressing weed growth as the HWC and at 35 and 49
DAT were actually more effective. Of the other treatments,
only the D. heterocarpon live mulch gave such good weed
(ii) Mucuna pruriens (velvet beans) drilled into the soil in control, being similar to the HWC at 35 and 49 DAT
rows 30 cm apart with 3 seeds per hole; (iii) Desmodium
heterocarpon var ovalifolium (CIAT 13651) broadcast at a
rate of 5 kg/ha. (4) Two organic treatments: (i) corn gluten Table 1
Effects of treatments on weed density at 21, 35 and 49 DAT; weed control
meal a pre-emergent weed blocker and a slow release scores at 63 DAT and dry matter production 112 DAT (composite of four
fertilizer (9-1-0), which controls emerging weeds and trials in 2003 and 2004)
provides nutrients to the crop, applied at a rate of 10 kg/
ha and (ii) concentrated vinegar and acetic acid, a post- Treatment No. weeds/x m2 Weed score % DM (g/m2)
emergent contact herbicide, sprayed directly to weeds at 21 DAT 35 DAT 49 DAT 63 DAT 112 DAT
the 35 leaf stage. (5) Two control treatments: (i) hand-
weeded control (HWC) which was hand weeded once every AP 26.0c 39.0d 45.5d 52.1e 83.8f
4 weeks and (ii) an unweeded control (UWC) which was BB 8.5b 24.0c 49.0d 48.1ef 83.2f
BM 0.5a 2.5a 2.5a 95.6a 3.2a
left unweeded from weeks 1 to 16. CH 0.0a 1.5a 2.5a 94.5a 24.7c
CWB 27.0c 50.5e 80.0e 42.6f 102.2g
2.3. Measurements CWC 2.0g 14.0b 20.5c 66.8d 41.4d
DH 13.0b 17.0b 18.0bc 86.7b 59.0e
To determine the suppressive effects of the various FF 0.0a 0.0a 1.0a 77.8c 11.0ab
FGA 0.0a 4.0a 11.5b 73.1cd 44.8d
treatments, weed counts were taken at 21, 35 and 49 days MP 38.0d 62.5f 82.0e 43.3f 81.6f
after the application of treatments (DAT) using xed PM 0.0a 0.0a 0.0a 72.4cd 19.0bc
quadrats (0.50 m2), with one placed randomly in each plot. UWC 47.5e 85.0g 127.0f 27.3g 90.5fg
Weed control efciency (WCE) were taken at 63 DAT SEM 3.6 4.4 5.2 5.3 15.9
using a scale from 1 to 10 where 1 represented poor WCE LSD (0.05) 5.4 5.9 6.5 6.3 11.5
Treat *site ** * NS * *
and 10 represented excellent WCE. Weed species biomass
(g dry mass/m2) was sampled using a 1 m2 quadrat Abbreviations: AP, Arachis pintoi; BB, burnout; BM, banana mulch; CH,
randomly placed in each plot at 112 DAT. These weeds coffee hulls; CWC, clean weed control; CWB, corn weed blocker; DH,
were cut at ground level with hand clippers and dried at Desmodium heterocarpon; FF, Flexstar Fomesafen; FGA, Finale
Glufosinate ammonium; MP, Mucuna pruriens; PM, clear plastic mulch;
60 1C in a forced draught oven for 48 h to a constant mass. UWC, unweeded control; DAA, days after application; DM, dry matter.
Weeds were sorted by taxa (species or genus, depending on *Po0.05; **Po0.01; NS, not signicant. (ag) Means followed by the
their similarity). Uncommon species were pooled and their same letter in the column do not differ signicantly (Po0.05).
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although not quite as effective as the non-living mulches or hampering proper establishment of the cover crop. Weed
Fomesafen. density in this treatment was high for most of the
Fomesafen and glufosinate-ammonium gave a fast kill of monitoring period; it was not until 12 weeks that any
emerging weeds, including C. diffusa. Fomesafen provided signicant weed suppression occurred. M. pruriens vigor-
long lasting and almost complete suppression of weed ously established itself, weed density was also high for this
growth for 49 DAT and its WCE at 63 DAT was treatment as most of its growth was in climbing the banana
moderately good at 78%. However, this herbicide was plant as reported by Buckles et al. (1998). The vines had to
also phytotoxic to young banana plantlets as well as leaves be removed often from bananas and their effectiveness in
of mature plants (430%). A single application gave suppressing weeds was low. Additionally, the vines began
effective control of C. diffusa up to 112 DAT. For the senescing after producing pods allowing further weed
other herbicide, glufosinate-ammonium, weed density had establishment. Buckles et al. (1998) noted that for effective
increased somewhat by 49 DAT, but its WCE at 63 DAT weed control with M. pruriens elds should be planted over
was similar to that of fomesafen. However, it was effective a 3-year period as exudates from senescing leaves have a
in controlling C. diffusa and a repeat application of herbicidal effect.
glufosinate-ammonium may be necessary for adequate The two organic treatments which included the vinegar
control of this species. and acetic acid and corn gluten meal were not as effec-
All three non-living mulches gave excellent suppression tive as other treatments in suppressing weed populations.
of weed growth for 49 DAT, even better than the HWC by Weed density under the corn gluten meal treatment
35 DAT. The WCE of the two dead mulches, banana increased from 53% to 62% from 35 to 49 DAT,
mulch and coffee hulls, at 49 DAT was around 95%. It is respectively, which was similar to the increase in the
probable that the relatively large amount of material UWC from 55% to 66% in the unweeded control at the
covering the soil surface was sufcient to suppress weed same DAT. Concentrated vinegar and acetic acid caused
seed germination and seedling growth. However, it is very rapid browning and drying of plant foliage within a
possible that exudates released from the decaying plant 24 h period with almost complete drying in 1 week. By
material also play a role. It is possible that WCE of the about 14 DAT plants began to recover from this initial
decomposing coffee hulls is not only due to the amount of injury as evidenced by an increasing amount of green
material applied on the soil surface but also to exudates tissues in plants. Burning was more pronounced on
released from the decaying material. Relating this to the broadleaved weeds, than C. diffusa and grasses and the
caffeine found in coffee, Rizvi et al. (1980) described latter quickly recovered and grew vigorously. Concentrated
caffeine as a natural herbicide selectively inhibiting vinegar and acetic acid have been reported to burndown
germinating seed of Amaranthus spinosus L. After the top growth only and not affect root growth such that
clear plastic mulch was removed at 42 DAT the stressed perennials may regenerate after a single application and
and etiolated weeds, which had germinated under the require additional treatment (Preston Sullivan, Appropri-
plastic recovered, causing an increase in weed density. ate Technology Transfer for Rural Areas (ATTRA),
However, the suppressive effects of this mulch still were September, 2003 unpublished data). This product is
seed at 63 DAT, with a WCE of 72%. Marenco and considered a contact herbicide, since its active ingredient
Lustosa (2000) reported an increase in seed germination of is not translocated within the plant and its primary site of
C. benghalensis L. when clear plastic was removed in a trial injury is surface contact with green plant tissue. Repeat
using clear plastic mulch for soil solarization in Brazil. To applications and thorough plant coverage would be
maintain the 56 cm thickness of the banana mulch, leaves necessary for effective management of perennial weeds
had to be frequently replaced. The practicality of this is a using concentrated vinegar and acetic acid. Control using
concern on a commercial scale, as there are unlikely to be corn gluten meal was also short term, causing death to
sufcient leaves available, as discussed by Cintra and germinating broadleaved weeds only and being a slow-
Borges (1988) in studies conducted in Brazil. The release fertilizer it presumably supplied nitrogen to grassy
availability of sufcient quantities of coffee hulls would weeds. However, in at least two locations, this poor weed
be a similar concern. control was confounded by removal of corn gluten meal by
Of the three cover crop treatments, D. heterocarpon the feeding activities of rats and domestic chickens. Both
suppressed the highest number of weeds at all 3 dates after concentrated vinegar and acetic acid and corn gluten meal
application. This was due to the fact that D. heterocarpon cannot be recommended for effective control of C. diffusa
gave better coverage and was therefore more competitive and other weeds in banana elds.
than A. pintoi and M. pruriens. The rapid establishment of Weed control scores at 63 DAT also showed similar
D. heterocarpon quickly suppressed emerging weeds which trends to the weed density at 49 DAT (Table 1 and Fig. 2).
was contrary to ndings by Bradshaw and Lanini (1995) The two dead mulches (banana mulch, 96%, and coffee
who noted that this cover crop required more than 2 hulls, 95%) showed excellent weed control. WCE was also
months for establishment and control of weeds in high (from 87% to 72%) in D. heterocarpon, followed by
Nicaraguan coffee. A. pintoi fell prey to predatory Rattus fomesafen, glufosinate-ammonium and clear plastic mulch
norvegicus (rats) and Gallus domesticus (domestic chickens) (from 87% to 72%). The hand-weeded treatment at 67%
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100 80

80
60
Mean Weed Score %

Weed DM g/m2
60

40
40

20
20

0
AP BM CWB DH FGA PM
BB CH CWC FF MP UWC 0
Treatments
AP BM CWB DH GA PM
BB CH CWC FF MP UWC
Fig. 2. Effect of treatments on mean weed score% (weed control Treatments
efciency) at 63 DAT.
Watergrass Broadleaves Grasses

Fig. 3. Effect of various treatments on the mean weed dry matter (DM) g/
m2 for watergrass, all broadleaved and grass weeds.
was similar to the latter two treatments. A. pintoi,
concentrated vinegar and acetic acid, M. pruriens and corn
gluten meal were less efcient (5243%) in controlling 3.3. Response of weed flora composition to the weed
weeds, but still better than the UWC at 27%. management treatments

3.2. Weed dry matter production of C. diffusa as influenced The removal of C. diffusa before treatment application
by the treatments at 112 DAT led to an increase in the weed ora composition. A total of
28 families were represented among the weeds occurring at
At 112 DAT there were also large and signicant the four sites with Poaceae, Asteraceae and Cyperaceae
differences between treatments (Table 1) in weed DM. being most common (Table 2). Fifty species in 46 genera
Lowest levels were found in banana mulch (3.2 g/m2) were identied and 74% of these were species with annual
followed by fomesafen (11 g/m2), clear plastic mulch life histories. Only 26% of the species were perennials. At
(19.0 g/m2) and coffee hulls (24.7 g/m2). These values were both sites C. diffusa (Commelinaceae) had been the
slightly lower than that for the HWC. Glufosinate- predominant weed before the sites were clean weeded.
ammonium produced similar levels, while the other Of the 50 weed species found in these trials, 28 were found
treatments gave slightly higher DM levels than the HWC. at Site I, 25 at Site II, 25 at Site III, 22 at Site IV with just
Highest levels were found in corn gluten meal (102.2 g/m2) seven common to all four sites. The common weeds were, six
which was similar to that of the UWC (90.5 g/m2). Weed broadleaved weeds, Pycreus polystachys (Rottb) Beauv.,
DM was also high (and similar to the UWC) in A. pintoi Phyllanthus tenellus Roxb., Oxalis barrelieri L., Peperomia
(83.8 g/m2) followed by concentrated vinegar and acetic pellucida (L.) Kunth., Spermacoce latifolia Aubl., Laportea
acid (83.2 g/m2) and M. pruriens (81.6 g/m2). aestuans (L.) Chew and C. diffusa (Table 2).
The abundance of C. diffusa in treatments was sig- The weed species composition shifted signicantly from
nicantly different with none found in coffee hull and C. diffusa to a number of annual weed species after the
banana mulch treatments at 112 DAT (Fig. 3). However, application of any of the treatments. At Site I S. latifolia
C. diffusa was signicantly suppressed by broadleaved was the most abundant weed species found in treatment
and grass weeds in all treatments. Broadleaved weed levels plots followed by Cyathula prostrata (L.) Blume. Other
were more abundant than grass weed levels in all weeds found at lower levels included Drymaria cordata (L.)
treatments (Fig. 3). Lowest DM of broadleaved weeds Willd. ex. Schult, Panichum trichoides Sw., L. aestuans and
were found in banana mulch followed by fomesafen, coffee Nautilocalyx melittifolius (L.) Wiehler. Cleome rutidosper-
hulls and clear plastic mulch and highest levels in the ma DC. and Cleome acuelata L. were most abundant at
UWC, M. pruriens, corn gluten meal, A. pintoi, concen- treatment plots in Site II. Cyperus spp. were found in
trated vinegar and acetic acid and D. heterocarpon. Grasses higher levels in the UWC, M. pruriens, A. pintoi and
were highest under corn gluten meal and concentrated concentrated vinegar and acetic acid. Eleusine indica (L.)
vinegar and acetic acid, both signicantly different to Gaertn. was prevalent at Site III with high levels in corn
the UWC. gluten meal and concentrated vinegar and acetic acid
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Table 2
Weeds recorded at the trial Sites IIV in Fairtrade banana elds in St. Vincent over the 2003/2004 growing season

Species Family Common name Life history Site

Amaranthus dubius Mart. Amaranthaceae Bhaji a III and IV


Cyathula prostrate (L.) Blume Amaranthaceae Cyathula a I
Catharanthus roseus G. Don Apocynaceae Periwinkle a III
Ageratum conyzoides L. Asteraceae Billy Goat Weed a I
Bidens pilosa L. Asteraceae Railway Daisy a I, II and IV
Emilia sonchifolia (L.) DC. Asteraceae Consumption Weed a I, II and III
Synedrella nodiflora (L.) Gaertn. Asteraceae a IV
Vernonia cinerea (L.) Less. Asteraceae Inammation Bush a I and II
Cleome aculeata L. Capparidacea Consumption bush a II
Cleome rutidosperma DC. Capparidacea Spider Plant a II
Drymaria cordata (L.) Willd. ex. Schult Caryophyllaceae West Indian Chickweed a I and II
Ipomoea quamoclit L. Convolvulaceae Sweet William a II, III and IV
Merremia aegyptia (L.) Urb. Convolvulaceae a I
Commelina diffusa Burm. Commelinaceae Watergrass p I, II, III and IV
Mormordica charantia L. Cucurbitaceae Wild carilla a I, II and IV
Cyperus cyperoides (L.) Kuntze. Cyperaceae p III
Cyperus esculentus L. Cyperaceae Nut Grass p I, II and III
Cyperus kyllingia Endl. Cyperaceae Spike hedge p I, II and IV
Cyperus luzulae (L.) Retz. Cyperaceae Razor grass p III
Pycreus polystachys (Rottb.) Beav. Cyperaceae p I, II, III and IV
Croton lobatus L. Euphorbiaceae Croton a II, III and IV
Phyllanthus tenellus Roxb. Euphorbiaceae Seed under leaf a I, II, III and IV
Centrosema pubescens Benth. Fabaceae p II and IV
Desmodium incanum DC Fabaceae Spanish Clover p I
Indigofera hirsuta L. Fabaceae Hairy indigo a III and IV
Nautilocalyx melittifolius (L.) Wiehler Gerneriaceae St. Vincent p I
Marsypianthes chamaedrys (Vahl.) O. Kuntze Labiatae a I and III
Spigelia anthelmia L. Loganiaceae Worm Bush a II
Sida rhombifolia L. Malvaceae Broom Weed a I
Mimosa pudica L. Mimosaceae Timari a III and IV
Boerhavia diffusa L. Nyctaginaceae Hog Feed a II
Ludwigia octovalvis (Jacq.). Raven Oenotheraceae Wild Clove p I
Oxalis barrelieri L. Oxalidaceae a I, II, III and IV
Microtea debilis Sw. Phytoaccaceae Demoiselle a III and IV
Peperomia pellucida (L.) Kunth. Piperaceae Shinning bush a I, II, III and IV
Piper dilatatum L. C. Rich. Piperaceae Elder Bush p I
Cenchrus echinatus L. Poaceae Burgrass a III and IV
Eleusine indica (L.) Gaertn. Poaceae Fowl Foot Grass a II and III
Chloris barbata Link Poaceae Purple top a III
Dactylotenium aegyptum (L.) Beauv. Poaceae Beach wiregrass a III
Panichum trichoides Sw. Poaceae Tickle Grass a I and II
Paspalum conjugatum Berg. Poaceae Sourgrass a III
Pityrogramma calomelanos (L.) Link Pteridaceae Fern p I
Portulaca oleraceae L. Portulacaceae Purslane a I and II
Hedyotis corymbosa (L.) Lam. Rubiaceae Mille-graines a II, III and IV
Spermacoce latifolia Aubl. Rubiaceae Button Weed a I, II, III and IV
Lindernia diffusa (L.) Wettst. Scrophulariaceae Spreading false pimpernel a IV
Thelpteris dantata (Forssk.) Thelypteridaceae Downy Maiden p I
Laportea aestuans (L.) Chew Urticaceae Sting-a-nettle a I, II, III and IV
Phenax sonneratii (Poir.) Wedd. Urticaceae a I and IV
Total number of species 50
Total number of genera 46 Total number of a 37
Total number of families 28 Total number of p 13
Total species at site I 28
Total species at site II 25
Total species at site III 25
Total species at site IV 22
Total species common to all sites 7

Life histories shown as annual (a) and perennial (p).


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treatments. At Site IV Croton lobatus L. was the most Buckles, D., Triomphe, B., Sain, G., 1998. Cover crops in hillside
abundant weed found in almost all treatments. agriculture: Farmer Innovation with Mucuna. International Develop-
The re-establishment of C. diffusa was suppressed in all ment Research Centre (IDRC), Mexico City, Mexico: International
Maize and Wheat Improvement Centre (CIMMYT) Ottawa, Canada.
treatments, including the HWC. The results indicate that Burns, J.H., 2004. A comparison of invasive and non-invasive dayowers
prior to the experiment, interspecic competition by the (Commelinaceae) across experimental nutrient and water gradients.
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