| |

Received: 12 June 2018    Revised: 14 January 2019    Accepted: 22 January 2019

DOI: 10.1111/gfs.12413

ORIGINAL ARTICLE

Seeding strategies of bahiagrass and pintoi peanut affect

pasture establishment under weed competition

Joao M. D. Sanchez1,2 | Joao M. B. Vendramini1  | Maria L. Silveira1 |

Lynn E. Sollenberger3 | Jose C. B. Dubeux Jr4  | Philipe Moriel1 | Brent Sellers1 |
James K. Yarborough1 | Fabio C. Leite de Oliveira5

1
Range Cattle Research and Education
Center, University of Florida, Ona, Florida Abstract
2
CAPES Foundation, Ministry of Education Pintoi peanut (Arachis pintoi Krapov. & W.C. Greg.) is a warm-­season perennial leg-
of Brazil, Brasilia, Brazil
ume with potential for use in grass–legume mixtures in Florida; however, limited in-
3
Department of Agronomy, University of
Florida, Gainesville, Florida
formation exists about its establishment in mixtures with bahiagrass (Paspalum
4
North Florida Research and Education notatum Flügge). The objective of this experiment was to evaluate the establishment
Center, Marianna, Florida of bahiagrass cv. “Argentine” and pintoi peanut cv. “Amarillo” as monocultures or mix-
5
Department of Animal Science, Maringa
ture. The experiment was conducted in Ona, FL, from June to October of 2014 and
State University, Maringa, Brazil
2015. Treatments were a split-­plot design of seeding strategies (bahiagrass monocul-
Correspondence
ture, pintoi peanut monoculture or bahiagrass-­pintoi peanut mixtures; main plots)
Joao M. B. Vendramini, Range Cattle
Research and Education Center, University and two N fertilization strategies (30 or 80 kg/ha N; 30N and 80N; subplots), with
of Florida, Ona, FL.
four replicates. Measurements of plant density and frequency were taken every
Email: jv@ufl.edu
4 weeks after seeding. Ground cover and herbage mass (HM) measurements were
Funding information
taken 112 days after seeding. Pintoi peanut ground cover was affected by seeding
Coordenação de Aperfeiçoamento de
Pessoal de Nível Superior; National Institute strategy × N level interaction. Ground cover was greater with 80N than 30N when
of Food and Agriculture; Florida Agricultural
pintoi was seeded in monoculture (3.6% vs 1.5% respectively) but not when it was
Experiment Station
seeded with bahiagrass (2.1%). There was no effect of seeding or N strategy on pintoi
peanut proportion in HM (1.4%). Bahiagrass ground cover was not affected by seed-
ing or N strategy (15.9%); however, its proportion in the HM was greater in 80N than
30N (12.1% vs 9.4% respectively). Mixed seeding did not negatively affect the estab-
lishment of bahiagrass and pintoi peanut and greater N fertilization levels improved
some establishment parameters, with no negative effect for pintoi peanut.

KEYWORDS
bahiagrass, establishment, grass–legume mixture, pintoi peanut

1 |  I NTRO D U C TI O N
Mixing legumes with warm-­season perennial grasses is an effective al-
ternative management practice to supply grasslands with N and increase
forage nutritive value (Thomas, 1995); however, lack of persistence has
been one of the major factors limiting the use of warm-­season legumes
in tropical and subtropical regions (Pitman, Chambliss, & Kretschmer,
1988). Rhizoma peanut (Arachis glabrata Benth.) is a persistent and pro-
ductive warm-­season perennial legume used in Florida; however, its
vegetative propagation and slow establishment limit its use in grazing
systems (Castillo, Sollenberger, Blount, et al., 2013; Castillo, Sollenberger,
Ferrell, et al., 2013; Castillo et al., 2014; Mullenix et al., 2014).

Grass Forage Sci. 2019;1–8. © 2019 John Wiley & Sons Ltd |  1
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2       SANCHEZ et al.
Pintoi peanut is another warm-­season perennial legume that has
been studied in South America during recent decades, with docu-
mented persistence under grazing on acidic and low-­fertility soils
(Rao & Kerridge, 1994). In comparison with rhizoma peanut, it has
the advantage of producing viable seeds, with some genotypes pro-
ducing more than 1 Mg/ha of seed (Carvalho & Quesenberry, 2012).
Establishing seed-­propagated plants is usually more convenient and
less expensive than establishing vegetatively propagated plants;
therefore, pintoi peanut might be an alternative to rhizoma peanut
in subtropical regions.
Similar to rhizoma peanut, the establishment of pintoi peanut is
generally slower than the establishment of warm-­season grasses, al-
though it varies according to location and cultivar (de Assis, Valentim,
Carneiro, de Azevedo, & Ferreira, 2008; Carvalho & Quesenberry,
2012; Perin, Guerra, & Teixeira, 2003). Perin et al. (2003) evaluated
the establishment of BRA-­14951 pintoi peanut in Seropedica (RJ,
Brazil) with initial plant densities ranging from 2 to 16 m−1 of row,
planted in rows spaced 25 or 50 cm apart. The authors observed
that ground cover was dependent on the initial plant population and
recommended a density of 8 plants/m with 50-­cm spacing between
rows; however, the soil was fully covered after 224 days in all treat-
ments. Conversely, de Assis et al. (2008) observed that the ground
cover at 304 days after planting 21 accessions that included pintoi
peanut, Arachis repens Handro, and crosses between these two spe-
cies was <50%, ranging from 11% to 43%. Such variation is likely
related to genotype differences, as well as adaptation to the envi-
ronment where they were planted. Most of the research about the
establishment of pintoi peanut has been done with vegetative plant-
ing; however, Baruch and Fisher (1996) observed that initial growth
of pintoi was greater when it was planted by seed rather than vege-
tative material.
Bahiagrass is the main warm-­season forage used in beef cattle
operations in Florida (Inyang et al., 2010). Establishment of pin-
toi peanut in mixture with bahiagrass has already been reported
(Bowman, Wilson, & Gogel, 1998; Jones, 1993); however, infor-
mation about establishment strategies for this mixture is scarce.
According to Skinner (2005), the composition of mixed forage
swards is critically dependent on plant-­to-­plant interactions during
the establishment phase, with some positive and negative effects of
mixed seeding being observed in temperate forages. As reviewed by
Gates, Quarin, and Pedreira (2004), bahiagrass also has slow estab-
lishment, mostly due to dormancy of the seeds and the low vigour
and size of the seedlings. The slow initial establishment of bahiagrass
may minimize the competition and could represent an opportunity
for legumes planted simultaneously to spread and assure a greater
initial proportion in the canopy. However, information about the im-
pact of simultaneously seeding bahiagrass and pintoi peanut on the
individual establishment performance of each plant was not found.
One of the main difficulties in establishing grass–legume mixed
swards is the inherent limitations on post-­emergence chemical weed
control (Sanderson, Brink, Ruth, & Stout, 2012). Selective herbicides
used to control broadleaf weeds may suppress desirable legumes,
and proper weed control in mixed swards is often difficult to achieve.
In the establishment of rhizoma peanut strips on pre-­existing bahia-
grass pastures, Castillo et al. (2014) recommend the use of glypho-
sate in the fall, to kill the grass, followed by a no-­till planting of the
legume on spring. The authors also recommend the post-­emergence
use of imazapic or imazapic + 2,4-­D on peanut strips. However, at
the time of the present publication, there are no registered herbi-
cides to be used in pintoi peanut crops, either as monoculture or in
mixed swards. Therefore, research addressing the practical estab-
lishment of this legume in the current conditions should avoid weed-
ing techniques that are not feasible on a farm scale.
According to Chambliss and Sollenberger (1991), N fertilization
is required during the establishment phase of bahiagrass. The au-
thors recommend 30 kg N/ha to be applied as soon as the seedlings
emerge, with an extra N fertilization of 70 kg/ha between 30 and
50 days after emergence. On the other hand, the use of N fertil-
ization in the establishment of legume crops is controversial, as
it may increase shoot and root growth, but it may also decrease
emergence, nodulation and N fixation (Hojjati, Templeton, & Taylor,
1978; Shuler & Hannaway, 1993; Woodman, Lowther, Littlejohn, &
Horrell, 1998). French, Prine, Ocumpaugh, and Rice (1994) observed
that the effect of N fertilization during the establishment of rhi-
zoma peanut was variable, and indicated that positive results may
be related with poor N2 fixation during the initial phase. Castillo,
Sollenberger, Ferrell et al. (2013) observed that applying 50 kg N/
ha may increase rhizoma peanut ground cover during establish-
ment, although the results depended upon weed control strategy.
Therefore, it was hypothesized that establishment parameters of
pintoi peanut and bahiagrass in mixed planting would be similar to
those of the respective plants in monoculture and increasing N fer-
tilization will improve growth and establishment of both plants. The
objective of this experiment was to evaluate establishment param-
eters of pintoi peanut and bahiagrass as monocultures or mixtures,
with two N fertilization regimens.
2 | M ATE R I A L A N D M E TH O DS
2.1 | Experimental site
The study was conducted at the University of Florida -­Range Cattle
Research and Education Center, Ona, FL (27°26′N 82°55′W), from
June to October (112 days). Plots were planted in June 2014 and
2015 in different sites at the research centre, one site per year. The
predominant soil was a Smyrna sand (sandy, siliceous, hyperther-
mic, Aeric Alaquods). Prior to initiation of the study, mean soil pH
(in water) was 5.9. Mehlich-­1 extractable P, K, Mg and Ca concen-
trations in the Ap horizon (0 to 15-­cm depth) were 12, 7, 73 and
608 mg/kg respectively.
2.2 | Treatments, experimental design
and management
Treatments were a split-­plot design of three seeding strategies (bahia-
grass monoculture, pintoi peanut monoculture or bahiagrass-­pintoi
SANCHEZ et al. |
      3

TA B L E   1   Establishment parameters of bahiagrass and pintoi the total surface of each plot after seeding. All plots were fertilized
peanut as affected by seeding strategy with 30 kg/ha each of N, P2O5 and K 2O, 2 weeks after planting. In

Seeding strategy addition, 2 kg/ha of the micronutrient mixture F-­503 (B—24 g/kg,

Cu—24 g/kg, Fe—144 g/kg, Mn—60 g/kg, Zn—56 g/kg) was applied.
Variables Monoculture Mixturea SE p valueb
This was the only fertilization received by 30N plots, while a second
Bahiagrass N fertilization of 50 kg/ha N was applied to 80N subplots 8 weeks
Frequency (%) 38.7 33.7 5.67 0.14 after planting.
Ground cover 17.5 14.3 5.42 0.39 As there was no selective herbicide registered for use in pintoi
(%) peanut crops at the time of planting the present experiment, all plots
Herbage mass 194 192 56.2 0.65 were mowed every 28 days at 7.5 cm for weed control. The height
(kg/ha) was chosen to avoid excessive defoliation of seedlings of both de-
Herbage mass 10.6 10.9 2.40 0.65 sired species.
proportion
(%)
Pintoi peanut 2.3 | Response variables
Density 6.7 5.4 0.73 0.20
Measurements of plant density and frequency were taken every
(plants/m2)
4 weeks after seeding. A 1-­ × 1-­m quadrat, subdivided into one hun-
Frequency (%) 11.7 10.2 5.57 0.11
dred 10 × 10-­cm squares, was placed in each plot, with flags perma-
Herbage mass 22 26 10.7 0.47
nently marking its position throughout the experiment. The number
(kg/ha)
of plants of pintoi peanut and bahiagrass (according to the treat-
Herbage mass 1.4 1.4 0.53 0.94
ment) in the full area of the quadrat was recorded for plant density.
proportion
(%) Stolons present in the evaluation area that originated from plants
outside the evaluation area were also counted in the plant density
Notes. Data are 2-­year least square means across 4 months.
a
Mixture: mixed seeding of bahiagrass and pintoi peanut. b p Value refers measurement. Frequency was expressed as the proportion of the
to the effect of seeding strategy within each response variable on the 10-­ × 10-­cm squares where pintoi peanut or bahiagrass plants were
transformed scale. present. Presence was counted when stems or stolons of plants
were present.
peanut mixture; main plots) and two N fertilization strategies (30 or Ground cover and herbage mass (HM) evaluations were done at
80 kg N/ha; referred to as 30N and 80N respectively; sub-­plot), with the termination of the experimental period each year, in October
main plots distributed in a randomized complete block design with 2014 and 2015. Ground cover was assessed by visual observation
four replicates. Plots were 4 × 4 m, subdivided into two 2 × 4-­m sub- of two 0.25-­m2 rings placed in the 1-­ × 1-­m marked area. Forage was
plots, with 1-­m alleys between main plots. The bahiagrass cultivar was harvested at 1 cm above soil level with a hand clipper in the same
Argentine, and the pintoi peanut cultivar was Amarillo. Seeding rates two 0.25-­m2 rings. The harvested material was dried in a forced-­air
were 25 and 10 kg/ha for bahiagrass and pintoi peanut respectively. dryer at 55°C until constant weight and then hand-­separated into
Seeding rate of bahiagrass was chosen to be relatively high to ensure bahiagrass, pintoi peanut or weed fractions, which were used for de-
establishment without chemical weed control, while seeding rate of termination of HM and species proportion in HM.
pintoi peanut was based on literature (Cook, Jones, & Williams, 1994).
The N fertilization strategies were based on the recommendations for
2.4 | Statistical analysis
bahiagrass (Chambliss & Sollenberger, 1991). The authors recommend
the application of 30 kg N/ha when seedlings emerge, which was the Response variables were analysed by fitting mixed-­effects models
basis for the 30N treatment. A second N application of 50 kg/ha was using the MIXED procedure of SAS (SAS Institute Inc., 2014; version
chosen as a moderate dose that is compatible with fertilization rates 9.4). Effect of seeding strategy on response variables was used to
usually adopted by local producers, totalling 80 kg N/ha. compare each plant species seeded as monoculture versus mixture
The experimental area was sprayed with glyphosate [N-­(phos- (e.g., plant density of bahiagrass seeded in monoculture vs. bahia-
phomethyl) glycine] at 1.44 kg/ha, around 30 days before planting. grass in mixture with pintoi peanut), except for the proportion of
After the sod was killed, the area was cultivated with a rototiller weeds in ground cover and dry matter. The models for plant density
until there was no remaining vegetation on the soil surface, with and frequency included seeding strategy, N fertilization strategy
the process being repeated 1 days before the planting. This pre-­ and month as fixed-­effect variables, while the models for botani-
sowing practice was chosen as it is commonly adopted by produc- cal composition included only effects of seeding and N fertilization
ers when establishing bahiagrass pastures in South Florida. Seeds strategies. The random term for both models included year, block
of bahiagrass were manually broadcast over the soil surface while and their interaction. When month was included in the model, it
seeds of pintoi peanut were placed in rows spaced 0.5-­m apart, at was analysed as a repeated measurement and the covariance struc-
approximately 5-­cm depth. Soil was raked and firmed by stepping on ture selected based on the least Akaike information criterion value.
 |
4       SANCHEZ et al.

TA B L E   2   Establishment parameters of bahiagrass and pintoi (a) Pintoi density

peanut as affected by N fertilization levels 9
a a
8
N fertilization a

Plant density (plants/m2)

7
Response variables 30Na 80N SE p valueb 6 b
5
Bahiagrass
4
Density (plants/ 49.3 41.7 8.04 0.18 3
m2) 2
Frequency (%) 35.6 36.8 5.65 0.65 1
0
Ground cover (%) 15.7 16.1 5.42 0.83
July Aug. Sep. Oct.
Herbage mass 143 242 56.2 <0.01* Month
(kg/ha)
Herbage mass 9.4 12.1 2.40 0.06 (b) Pintoi frequency
proportion (%) 25
Pintoi peanut a
20 a
Density (plants/ 5.4 6.6 0.70 0.11

Frequency (%)
m 2) b b
15
Frequency (%) 9.9 12.0 5.94 0.04*
10
Herbage mass 22 26 10.7 0.21
(kg/ha)
5
Herbage mass 1.3 1.4 0.53 0.88
proportion (%) 0
July Aug. Sep. Oct.
Notes. Data are 2-­year least square means across 4 months. Month
a
30N: 30 kg/ha N 14 days after seeding; 80N: 30 kg/ha N 2 week after
seeding plus 50 kg/ha N 8 week after seeding. b p Value refers to the ef- F I G U R E   1   Pintoi peanut plant density (a) and frequency (b) as
fect of N fertilization levels within each response variable on the trans- affected by month (p < 0.03 and 0.02 respectively). Data are 2-­year
formed scale. least square means. Means followed by similar lowercase letters are
*Statistically significant p value (p ≤ 0.10). not different on the transformed scale (p > 0.10)

Autoregressive covariance structure was used in pintoi peanut plant
density and frequency, while toeplitz structure was used in bahia-
grass plant density and frequency. Normality of residues and ho-
mogeneity of variances were tested using conditional studentized
residual plots and data transformed if ANOVA assumptions were
violated. Logarithmic transformation was used for bahiagrass plant
density, ground cover, HM and HM proportion, as well as pintoi pea-
nut frequency, ground cover and HM. Square transformation was
used in weed ground cover and HM percentage. Means reported are
non-­transformed least square means. Treatments were considered
different when p ≤ 0.10 by LSD test.
3 |   R E S U LT S
3.1 | Plant density
Pintoi peanut plant density was not affected by either seeding
(Table 1) or N fertilization (Table 2); however, it was affected by
month (Figure 1). Density increased from July to August (4.3 vs.
6.1 plants/m2 respectively) and then remained similar in September
and October (6.7 and 7.1 plants/m2 respectively). Bahiagrass plant
density was not affected by N fertilization (Table 2); however, it was
affected by a seeding strategy × month interaction (Figure 2). The
interaction occurred because in July, bahiagrass monoculture plots
had greater plant density than mixed plots (35.4 vs. 14.4 plants/
m2 respectively), but such difference was not detected in August,
September and October (means = 47.3, 62.9, and 47.1 plants/m2 re-
spectively). Plant density increased from July to August and remained
constant through the other months for both seeding treatments.
3.2 | Frequency
Pintoi peanut frequency was not affected by seeding strategy
(Table 1); however, it was affected by N fertilization (Table 2) and
month (Figure 1). There was greater pintoi frequency for 80N than
30N (12.0% vs. 9.9% respectively). The frequency was similar be-
tween July and August (7.4% and 8.4% respectively), increased
in September (12.8%) and remained similar in October (14.8%).
Bahiagrass frequency was not affected by either seeding (Table 1)
or N fertilization (Table 2) strategies; however, it was affected by
month (Figure 2). The frequency increased from July to September
(from 23.7% to 45.6%), remaining similar in October (44.8%).
3.3 | Ground cover
Pintoi peanut ground cover was affected by a seeding strategy × N
fertilization interaction (Table 3), which happened because greater
pintoi peanut ground cover was observed in 80N than 30N in pin-
toi peanut monoculture only. Conversely, pintoi peanut ground
SANCHEZ et al.       5|
cover was not affected by N fertilization when it was seeded in TA B L E   3   Pintoi peanut ground cover as affected by seeding
mixture with bahiagrass. Bahiagrass ground cover was not af- strategy × N fertilization (p < 0.02)
fected by either seeding (Table 1) or N fertilization (Table 2) strat- Seeding strategy
egy (mean = 15.9%).
Monoculture Mixture SE

N fertilization %
3.4 | Herbage mass and botanical composition
a
30N 1.5 ac 2.2 a 0.94
Neither pintoi peanut HM or proportion in HM was affected by seed-
80N 3.6 a 2.0 b
ing (Table 1) or N fertilization (Table 2) strategy. Herbage mass of
p valueb <0.01* 0.93
bahiagrass was not affected by seeding strategy (Table 1); however,
a
it was affected by N fertilization (143 vs. 242 kg/ha for 30N and 30N: 30 kg/ha N 14 days after seeding; 80N: 30 kg/ha N 2 week after
seeding plus 50 kg/ha N 8 week after seeding; Mixture: mixed seeding of
80N respectively; Table 2). Similarly, the proportion of bahiagrass in
pintoi peanut and bahiagrass. b p Value refers to the effect of N fertiliza-
tion levels within each seeding strategy on the transformed scale.
(a) Bahiagrass density c
Means followed by similar lowercase letters within rows are not differ-
90 ent (p > 0.10).
A
80 *Statistically significant p value (p ≤ 0.10).
a A
Plant density (plants/m2)

70 Aa a
60
50 Ab the HM was not affected by seeding strategy (Table 1), but it was
40 A greater in 80N when compared to 30N (12.1% vs. 9.4% respectively;
30 A a Table 2).
20 a A
Bahia Bahia/Pintoi
10
B a
0 b 3.5 | Weed ground cover and proportion in HM
July Aug. Sep. Oct.
Month Weed ground cover was only affected by seeding strategy (Table 4).
Pintoi peanut monocultures had greater weed ground cover than
(b) Bahiagrass frequency
bahiagrass-­
pintoi peanut and bahiagrass monoculture plots. The
60
a a proportion of weeds in HM was affected by seeding and N fertiliza-
50 tion strategies (Table 4). Similar to weed ground cover, pintoi peanut
b monocultures had greater weed proportion in HM than bahiagrass-­
Frequency (%)

40
c pintoi peanut and bahiagrass monoculture plots (Table 4). The pro-
30
portion of weeds in HM was slightly greater in 30N compared to
20
80N (94.1% vs. 91.0% respectively; p < 0.02).
10

0
July Aug. Sep. Oct.
4 | D I S CU S S I O N
Month

F I G U R E   2   Bahiagrass plant density (a) as affected by seeding
strategy × month interaction (p < 0.04) and frequency (b) as
affected by month (p < 0.01). Data are 2-­year least square means.
Means followed by similar uppercase letters within each month and
lowercase letters within each seeding strategy are not different on
the transformed scale (p > 0.10)
Most of the pintoi peanut seed germination occurred within the first
2 weeks of the experiment, with a germination rate of nearly 60%
(data not shown), which was similar to previous reports of Ferguson
(1994). The number of seedlings was the main driver of plant den-
sity during the first month after planting. In the following months,

TA B L E   4   Weed ground cover and

Seeding strategy
proportion in herbage mass as affected by
seeding strategies of bahiagrass and pintoi Pintoi peanut Mixture Bahiagrass SE
peanut
Response variables % p valuea

Weed ground cover 86.1 ab 75.9 b 71.0 b 6.83 0.03*

Weed in HM 98.7 a 89.7 b 89.4 b 2.13 <0.01*

Notes. HM: herbage mass; Mixture: mixed seeding of bahiagrass and pintoi peanut.
a
p-­Value refers to the effect of seeding strategy in each response variable on the transformed scale.
b
Means followed by similar lowercase letters within rows are not different (p > 0.10).
*Statistically significant p value (p ≤ 0.10).
 |
6       SANCHEZ et al.
the increase in plant density may have resulted from germination of
new seedlings, but it was also affected by the spread of plants that
as seedlings were outside the sampling area. Plant-­to-­plant interac-
tions are very important factors affecting growth of mixed canopies,
including the establishment phase (Iannucci, 2006; Skinner, 2005).
Skinner (2005) observed that decreased emergence and growth of
temperate legumes may be observed when they are planted in mix-
tures with temperate grasses; however, such effects seemed to inter-
act with other factors, such as environmental conditions. Literature
regarding the same effect in pintoi peanut has not been found. In the
present experiment, there were no negative effects of mixed seed-
ing on pintoi plant density, indicating that bahiagrass did not impact
germination and mortality of pintoi peanut seedlings. Additionally,
despite the high proportion of weed ground coverage and herbage,
soil surface was not totally covered until the last week of July, when
most of pintoi peanut seeds had already germinated. Therefore, data
indicate that pintoi peanut germination and mortality were not af-
fected to large extent by weed competition as well. Woodman et al.
(1998) observed that N fertilization can decrease the germination
of some temperate legume species if applied with the seeds. The
authors speculate that the cause might be related to toxicity of N
compounds or osmotic effects nearby the germinating seeds. In the
present experiment, the N fertilizer was applied on the soil surface,
while pintoi seeds were placed 5-­cm deep. Additionally, the first N
fertilization (30 kg/ha) was applied to all the plots 5–10 days after
the emergence of the first pintoi peanut seedlings, with the second
application (50 kg/ha) being applied when most of them had already
emerged; therefore, it is unlikely that the second N fertilization
caused damage to the plants, justifying the lack of N effect.
The increase in plant density of bahiagrass over time was expected
because seeds of bahiagrass germinate gradually (Gates & Mullahey,
1997; Gates et al., 2004). According to West and Marousky (1989),
the delay in germination may be related to physiological processes
affecting embryo maturity and physical barriers present in the seed
preventing its imbibition and growth. However, no evidence was
found in the literature explaining the lower plant density observed in
the mixed-­seeding plots in July. Bahiagrass seedlings started germi-
nating approximately 1 week after pintoi peanut seedlings had first
emerged. Therefore, plant-­
to-­
plant interactions decreasing plant
density of bahiagrass in July are possible. Even though bahiagrass
plant density was affected by seeding strategy only in July, weed
encroachment was intense in these months. Therefore, it is possible
that seeding strategy effects would happen in other months if weed
competition was suppressed. In previous research, timing of N fertil-
ization did not affect bahiagrass seedling emergence (Busey, 1992),
similar to what was observed in the present experiment.
The frequency observed in July is mostly related to the germi-
nation of seeds, as most seedlings individually accounted for only
one frequency percentage unit. Pintoi peanut is a prostrate growing
plant which produces long stolons that expand laterally (Carvalho
& Quesenberry, 2009), increasing its frequency without impacting
plant density. The lack of seeding strategy effect on pintoi peanut
frequency supports the hypothesis of the current study; therefore,
the initial spread of pintoi peanut was not affected by the inclusion of
bahiagrass in the canopy. However, weed pressure was intense in all
the plots since the first experimental week, and weeds were a major
ground cover component (Table 4). It is possible that in weed-­free
conditions, the frequency of pintoi monoculture would differ from
pintoi seeded in mixture with bahiagrass. The increase in pintoi pea-
nut frequency with greater N fertilization levels has already been re-
ported in the establishment of other legume species, and it might be
related to greater N availability in the soil when root nodulation and
N fixation in seedlings are not fully developed (Hojjati et al., 1978).
Bahiagrass seedlings are small (Gates et al., 2004), and the fre-
quency observed in the first month was a direct result of number
and distribution of germinating seeds. Unlike pintoi peanut, the in-
crease in bahiagrass frequency along the months seems to be con-
sistent with the increase in plant density, which indicates that the
increase of frequency related to growth of individual seedlings was
likely small. In fact, comparing the averages of plant density and fre-
quency, each bahiagrass plant accounted for 0.8 percentage units
of frequency, while each pintoi plant accounted for 1.8 percentage
units. However, the lower plant density for bahiagrass-­pintoi peanut
in July was not reflected in frequency. Similar to discussed for pintoi
peanut, weed pressure may have impacted bahiagrass frequency as
well. The lack of N fertilization effect on frequency is consistent with
plant density results.
Reasons for the seeding strategy × N fertilization strategy inter-
action affecting pintoi peanut ground cover are not clear. The lack
of response to N in the presence of bahiagrass might be related to
the greater capacity of grasses to retain fertilizer N when in greater
proportion in grass–legume mixtures (Nyfeler, Huguenin-­Elie, Suter,
Frossard, & Lüscher, 2011). However, despite greater presence of
bahiagrass than pintoi peanut in mixed plots (10.8% vs. 1.4% of HM,
respectively, Table 1), the majority of the canopy, both in terms of
HM or ground cover, was represented by weeds (Table 4). Therefore,
other aspects related to different plants might have taken place in
this interaction as well.
Overall ground cover of pintoi peanut in the present experiment
was less than previous data presented in the literature. de Assis et al.
(2008) observed that Amarillo pintoi peanut covered more than 35%
of the soil at 112 days after planting. Perin et al. (2003) evaluated
the effect of several seedling arrangements on the establishment of
BRA-­14951 pintoi peanut and observed that the seedling density of
4 plants/m2 (similar to observed in this experiment) covered around
40% of soil surface at 112 days after planting. However, Carvalho
and Quesenberry (2012) reported that several accessions of pin-
toi peanut had slow establishment in North Florida, with none of
the tested genotypes covering more than 50% of the soil around
1 year after planting. Reasons for the differences in ground cover
between the current study and previous literature are likely related
to the differences in weed pressure, as the authors mentioned above
removed weeds manually while such practice was not done in the
present study.
The ground cover measurements were taken once at 112 days
after seeding, which corresponds to the October evaluation.
SANCHEZ et al.
Therefore, lack of treatment effects on ground cover at this date
is consistent with the plant density and frequency measurements.
The lack of seeding strategy effect on HM and botanical compo-
sition is in agreement with the previous response variables. Seeding
pintoi peanut in monoculture did not increase its proportion in HM
when compared to bahiagrass-­pintoi peanut mixture, indicating that
pintoi peanut was not negatively affected by the mixture with bahia-
grass during the initial establishment. The increase in pintoi peanut
frequency caused by greater N fertilization levels was not consistent
with the lack of the same effect on HM and botanical composition.
Unlike the observations for other response variables, greater N
fertilization inputs increased both HM and the proportion of bahia-
grass in the HM. Positive response of bahiagrass to N fertilization level
has been reported in literature (Blue, Dantzman, & Impithuksa, 1980).
The presence of bahiagrass was the main factor decreasing the
weed presence in the ground cover or HM even though the weed
proportion in the present experiment was greater than observed in
previous literature (Busey, 1992; Jaramillo et al., 2018). According to
Sanderson et al. (2012), the use of grass–legume mixtures decreased
weed proportion in HM when compared with grass or legume mono-
cultures in Pennsylvania. The authors also observed that species
composition is an important factor affecting the weed suppression
in mixed canopies, with weed proportion in HM being negatively
correlated with the proportion of orchardgrass in the seed mixture.
Jaramillo et al. (2018) observed minimal weed pressure when pin-
toi peanut was overseeded in established bahiagrass pastures. Such
practice was not evaluated in the current study and the reports of
Jaramillo et al. (2018) indicated it has potential to decrease weed
pressure during the establishment of pintoi peanut-­bahiagrass mixed
swards; however, literature comparing these techniques was not
found. Weed control in pintoi peanut plantings is an important sub-
ject that deserves attention if the plant is to be used in commercial
operations. Currently, there are no selective herbicides registered in
the USA to weed control in pintoi peanut.
According to Busey (1992), bahiagrass establishment may be
achieved under weedy conditions if proper fertilization is applied.
The author observed increase in HM of bahiagrass swards from the
first to the second year after seeding, even with weed competition
in early establishment phases. In fact, Chambliss and Sollenberger
(1991) observed that obtaining a well-­established bahiagrass sward
may take two growing seasons. Therefore, it is possible that exper-
imental units with bahiagrass in the present experiment would be
fully established in the coming growing season, even with the ob-
served weed encroachment. Contrasting with bahiagrass, literature
indicating the feasibility of establishing pintoi peanut under high
weed pressure was not found.
5 | CO N C LU S I O N S
Seeding bahiagrass and pintoi peanut either as monocultures or
grass–legume mixtures did not affect most of the establishment
variables in plots where weeds were controlled only by mowing
|
      7
every 28 days. Additional application of N fertilization 8 weeks
after planting did not negatively affect pintoi peanut, but rather
improved its spread, increased bahiagrass HM and decreased
weed proportion in the HM. Pintoi peanut may be seeded simul-
taneously with bahiagrass with no measurable impairment of ini-
tial establishment of either species. Based on these data, seeding
pintoi peanut in mixture with bahiagrass rather than in monocul-
ture is a strategy to decrease weed encroachment during initial
establishment.
AC K N OW L E D G M E N T S
We thank Frank A. Kuwahara, Cecilio V. Soares Filho and Ulysses
Cecato for their support during the field evaluations. We thank the
CAPES foundation for sponsoring the first author's Ph.D. programme
and USDA/NIFA and Florida Agricultural Experiment Station for
partially funding the research project.
C O N FL I C T O F I N T E R E S T
The authors have no conflict of interest to declare.
ORCID
Joao M. B. Vendramini  https://orcid.org/0000-0003-3383-1620
Jose C. B. Dubeux  https://orcid.org/0000-0001-8269-9959
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How to cite this article: Sanchez JMD, Vendramini JMB,
Silveira ML, et al. Seeding strategies of bahiagrass and pintoi
peanut affect pasture establishment under weed
competition. Grass Forage Sci. 2019;00:1–8. https://doi.
org/10.1111/gfs.12413