Polibotánica

ISSN: 1405-2768
[email protected]
Departamento de Botánica
México

Montejo-Mayo, Wilber; del-Val, Ek; Gomez-Romero, Mariela; de la Barrera, Erick; Lindig-

Cisneros, Roberto
INTERACTIONS BETWEEN DOMINANT HYDROPHYTIC SPECIES OF THE
WETLANDS OF WESTERN MEXICO MEDIATED BY FIRE AND NITRATE
CONCENTRATION
Polibotánica, núm. 40, agosto, 2015, pp. 153-161
Departamento de Botánica
Distrito Federal, México

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 Núm. 40, pp. 153-161, ISSN electrónico: 2395-9525; México, 2015
DOI: 10.18387/polibotanica.40.10
INTERACTIONS BETWEEN DOMINANT HYDROPHYTIC
SPECIES OF THE WETLANDS OF WESTERN MEXICO
MEDIATED BY FIRE AND NITRATE CONCENTRATION

INTERACCIONES ENTRE ESPECIES HIDRÓFITAS

DOMINANTES DE HUMEDALES DEL OCCIDENTE DE
MÉXICO MEDIADAS POR EL FUEGO Y LA CONCENTRACIÓN
DE NITRATOS

Wilber Montejo-Mayo1, Ek del-Val2, Mariela Gomez-Romero2,

Erick de la Barrera2, y Roberto Lindig-Cisneros2,3
1
Facultad de Biología, Universidad Michoacana de San Nicolás de Hidalgo, Morelia,
Michoacán, México.
2
Centro de Investigaciones en Ecosistemas, Universidad Nacional Autónoma de México.
Apartado Postal 27, Admón. 3, Santa María, CP 58091, Morelia, Michoacán, México.

SUMMARY complex and differed between sampling

years. For the first year, only the effect of
Wetland plant community dynamics are Typha on itself was significant. For the se-
strongly driven by abiotic factors such as cond year, the effect of Typha was negative
nutrient levels and fire. In wetlands where on Phrangmites, the effect of Schoenoplectus
invasive species are present assessing the was negative on itself and, the effect of
role of abiotic factors in plant-plant interac- Typha and Phragmites was negative on
tions is fundamental for both understanding Typha. The effect of fire was significant for
community dynamics and management. In Phragmites and Schoenoplectus, not bur-
this study the interaction between pairs of ned plant were taller. Interactions between
species was quantified between one non-in- plants were altered by fire, the presence of
vasive (Schoenoplectus americanus), one Schoenoplectus benefited the performance
that can become overdominant after human of Typha, and the opposite was also true.
disturbances (Typha domingensis) and one For early stages of these species growth,
that has an invasive lineage present in North the strongest competition occurs between
America (Phragmites australis), growing the native species, allowing the invasive
under different nitrate concentrations and to mostly respond to nitrate levels, and fire
subjected to fire. All species responded by have a strong effect also on the interactions
increasing growth to nitrate addition, but between the natives and not with the inva-
depending on the species, the response was sive.
significant for different variables. Phragmites
for height and root biomass, Typha for aerial Key words: wetland, negative interactions,
and root biomass and Schoenoplectus for all management, nutrients, ecological resto-
variables. Interactions between species were ration.

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 Núm. 40: 153-161 Agosto 2015

RESUMEN en la interacción entre nativas y no con la

invasora.
La dinámica de comunidades vegetales de
humedales está influenciada por factores Palabras clave: humedal, interacciones ne-
abióticos, como los nutrimentos y el fue- gativas, manejo, nutrimentos, restauración
go. En humedales en donde hay especies ecológica
invasoras, determinar el papel de factores
abióticos en las interacciones planta-planta INTRODUCTION AND METHODS
es fundamental para entender la dinámica
de la comunidad y para su manejo. En Wetland plant communities are complex
este estudio, la interacción entre pares systems in which abiotic factors can sig-
de especies fue cuantificada entre una no nificantly alter its floristic composition
invasora (Schoenoplectus americanus), (Zedler and Kercher, 2004). Among the most
una que puede volverse sobredominante important of these factors are hydroperiod,
como consecuancia de disturbios huma- salinity, and nutrient concentration, especially
nos (Typha domingensis) y una que posee that of nitrogen and phosphorus (Cronk and
un linaje que es invasor en norteamérica Fennessy, 2001). It has been suggested that
(Phragmites australis), las cuales crecieron when wetlands are subjected to changes in
bajo diferentes concentraciones de nitrato y the chemical composition of their water
fueron sometidas al efecto del fuego. Todas supply, particularly in the form of increases
las especies respondieron a la adición de in nitrogen concentration, changes can occur
nitratos incrementando su crecimiento. in the richness and/or composition of species
Phragmites respondió en altura y biomasa that can facilitate the domination of the sys-
de raíces, Typha en biomasa aérea y de raíces tem by invasive species (Cronk, 1995), whe-
y Schoenoplectus para todas las variables. ther native or exotic (Zedler and Kercher,
Las interacciones entre especies fueron 2004). Among the native species that present
complejas y variaron entre años. Durante invasive behavior in wetlands many belong
el primer año, sólo el efecto de Typha en to the genus Typha which, in response to
si misma fue significativo. Para el segundo increases in nutrient concentration and other
año, el efecto de Typha fue negativo en human disturbances, can form dense monos-
Phrangmites, el efecto de Schoenoplectus pecific patches and thus displace other native
fue negativo en sí mismo y el efecto de species (Galatowitsch et al., 1999; Smith et
Typha y Phragmites fue negativo en Typha. al., 2001; Woo and Zedler, 2002; Levin et
El fuego fue significativo para Phragmites al., 2006). Others, such as certain lineages
y Schoenoplectus, las plantas no quemadas of Phragmites australis (Swearingen and
fueron más altas. Se detectaron efectos del Saltonall, 2010) are particularly aggressive
fuego en las interacciones, la presencia de invasives (Rickey and Anderson, 2004; Lae-
Schoenoplectus benefició el desempeño de gue et al., 2006; Saltonstall and Stevenson,
Typha, y viceversa. En etapas tempranas, las 2007). Phragmites australis is a species
interacciones más fuertes ocurrieron entre considered to be native to Eurasia by some
las especies no invasoras, permitiendo a la authors without recognizing subspecies (Lot
invasora responder a la concentración de et al., 2013), others (Saltonal et al., 2004)
nitratos, y el fuego tuvo su mayor efecto recognize one subspecies that they consi-

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Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico

der to be native to Canada and the United the wetland respond mainly to the addition
States of America, Phragmites australis of nitrogen; S. americanus also responds to
subsp. americanus, and a lineage present increased nitrogen, but also to phosphorus in
from the Southern United States to Central terms of the accumulation of aerial biomass
America with uncertain origin, although this (Escutia-Lara et al., 2010). The responses
last lineage has been proposed as a new of P. australis have not been evaluated in
species, Phragmites karka (Ward, 2010). this system but, in other wetlands, it has
Despite the taxonomical discrepancies, an been reported that this species responds
invasive form is recognized, characterized to increases in the concentrations of both
by morphological and population dynamics nitrogen and phosphorus and also to the
characteristics, in particular that forms dense interaction of these two nutrients (Boar,
clumps that exclude all other plant species 1996; Ennabili et al., 1998; Ehrenfeld,
(Sweringen and Saltonall, 2010). 2003). From field observations in the model
wetland (Escutia-Lara et al., 2009a) and
To understand the dynamics of the displace- from controlled experiments (Escutia-Lara
ment of native species by invasive species et al., 2009b; Escutia-Lara et al., 2010), it
in wetlands, it is necessary to quantify the can be suggested that the competitive capa-
effects of factors, such as nutrient contents city of P. australis is greater or equal to that
or fire, that can modify the interactions be- of T. domingensis and, in turn, greater than
tween species. The present study is based that of S. americanus; however, the mag-
on observations of the vegetation structure nitude and exact nature of the interactions
and dynamics in a model system, the spring between these species remains unknown.
wetlands of the Mintzita (101°17’47” W, The objectives of the present study were
19°38’43” N: 1930 masl) in the municipality to quantify the response of these species to
of Morelia, in Michoacán, México (Escutia- different concentrations of nitrate, as this is
Lara et al., 2009a). These wetlands are cha- the macronutrient to which all three species
racterized by a natural zonation consisting respond most markedly (Escutia-Lara et
of an area dominated by Schoenoplectus al., 2010) and to fire, in order to relate this
americanus close to the edge of the wetland, recurring disturbance factor to patterns of
and a zone dominated by Typha domingensis species growth.
that conforms to changes in the water level.
Between both of these areas, there is a zone Rhizomes of the three species were collected
in which the two species coexist and where at the study site; these were selected to be of
the distributions respond partially to the wa- similar size at the start of the experiment and
ter level throughout the wetland; however, were assigned at random to each of the re-
other factors, such as nutrient concentration plicates of the experiments described below.
and fire also seem to play an important role. Experiments were conducted to quantify
Phragmites australis forms dense clumps the effect of interactions between pairs of
consistent with those reported for the inva- species: Typha domingensis, Phragmites
sive lineage (Sweringer and Saltonal, 2010) australis and Schoenoplectus americanus.
in the wetlands irrespective of the dominant Each experimental unit consisted of two
native species. In a previous study, it was plants, either of the same species or of each
found that individuals of T. domingensis in possible pairing between them. Rhizomes

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 Núm. 40: 153-161 Agosto 2015

were planted in five liter pots (25 cm high were conducted with the statistical program
x 16 cm wide) filled with commercial peat. R (R Development Core Team, 2011).
Thirty pots were planted with each one of
the six possible species combinations (three RESULTS
with interspecific and three with intraspeci-
fic pairs) giving a total of 180 experimental Following burning of the aerial parts of the
units. In addition, for each species-interac- plants in the second growing season, all
tion treatment, the effect of five treatments species responded to the addition of nitrate
of fertilization with potassium nitrate and, in some cases, to the interaction treat-
(KNO3) was tested: 0, 0.08, 0.16, 0.24 and ments and burning. For P. australis, there
0.32 g/l of cultivation medium. In this way, was a significant response in plant height
six replicates were produced for each treat- to the added fertilizer (fig. 1a; F(1,78) =
ment of the combination of competition x 71.2, P < 0.000001). The effect of the inte-
fertilization. Addition of KNO3 was carried ractions (fig. 1b) was significant (F(2,78) =
out every 15 days for a period of six months 6.6, P = 0.002): the intraspecific interaction
(May to October). During the dry season produced the greatest average plant height
(February of the following year), 15 pots (138 cm), followed by the interaction with
were chosen at random from each interac- S. americanus (120 cm) and with T. do-
tion treatment (three for each fertilization mingensis (115 cm). Burning of the aerial
X interaction combination of treatments) parts did not affect height in P. australis.
in order to burn the aerial biomass, while In the case of aerial biomass (fig. 1b), sig-
the other 15 were left as a control. In June, nificant differences were observed due to
all remaining plants were harvested and the addition of nitrate (F(1,78) = 118.3, P <
aerial and root biomass recorded. In order 0.000001) and to the effect of the interaction
to assess the importance of the interaction (F(2,78) = 6.9, P = 0.002). These followed
between the different species, three response the same pattern observed in plant height
variables were analyzed: plant height, and (41 g for intraspecific interaction, 32 g for
aerial and root biomass. In the case of same the interaction with S. americanus and 26
species interactions, the response variable g for the interaction with T. domingensis).
considered was the average value from Significant differences (F(1,78) = 8.32,
both plants in each pot. For different species P = 0.005) were found with the burning
interactions, the variable was measured in treatment, since greater quantities of aerial
the individual of the species on which the biomass accumulated in those plants that
effect of the interaction was to be determi- had not been burned (36 g), relative to those
ned. Analysis was conducted with general that had (30 g). Root biomass (fig. 1c) only
linear models, considering fertilization as a increased with increased nitrate addition
continuous variable, and the interaction and (F(1,78) = 18.4, P < 0.0001).
aerial biomass burning as categorical varia-
bles. Tests of hypothesis were carried out to Schoenoplectus americanus presented a
generate ANOVA tables (Crawley, 2007) more complex set of responses than P.
and to determine the effect of each of the australis. In terms of height (fig. 1d), this
variables and their interactions. All analyses species responded to both nitrate addition

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Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico

Fig. 1. Response of Phragmites australis, Schoenoplectus americanus and Typha domingensis

to the treatments of nitrate addition, competition and burning at the end of the experiment.
Each panel (a to h) shows the relationship between the response variable and the con-
centration of added nitrate, and the means and standard errors for the response variable
as a function of the competition interaction or the burning treatments. Slope curves are
shown only when the effect of the nitrate addition was significant. In cases where some of
the interactions between factors were significant, adjustment curves are shown for each
combination of levels. Different letters denote significant differences in means according
to the Tukey test.

157
 Núm. 40: 153-161 Agosto 2015

(F(1,78) = 99.1, P < 0.000001) and to the plex pattern of responses to treatments: there
interaction with other individuals (F(2,78) were significant effects of nitrate addition
= 3.24, P = 0.045), since the plants reached (F(1,78) = 91.0, P < 0.000001), plant-plant
the tallest average height (95 cm) in the interactions (F(2,78) = 8.0, P = 0.0006) and
presence of P. australis, followed by those of the combination of factors (F(2,78) = 4.1,
grown with T. domingensis (92 cm) and P = 0.02), since the intraspecific interaction
finally by those grown in the intraspecific treatment with fire produced a lower root
treatment (86 cm). They also responded biomass (4.6 g). Finally, the interaction be-
to the burning treatment (F(1,78) = 9.1, P tween the three factors; addition of nitrate,
= 0.003), with plants that had been burned plant-plant interactions and burning was
reaching an average height of 96 cm, while also significant (F(2,78) = 5.5, P = 0.005),
non-burned plants only reached 86 cm. as reflected in the slopes of the curves of
There were also significant effects of the root biomass as a function of nitrate addi-
interaction between nitrate addition and bur- tion for each combination of the levels of
ning treatment (F(1,78) = 4.15, P = 0.045), experimental factors.
since non-burned plants presented lower
rates of height increase than burned plants, Typha domingensis responded to fewer
with increasing concentrations of nitrates, as factors than the other two species, since
reflected in the slopes of the curves. The in- for no variable was there a response to the
teraction between competition and burning burning treatment. For plant height (fig. 1g),
treatment was also significant (F(2,78) = this species responded to the addition of
5.73, P < 0.004) because the burned plants nitrate (F(1,78) = 17.0, P < 0.0001) and to
from the treatment of competition with T. plant-plant interactions (F(2,78) = 18.3, P <
domingensis reached a greater height (104 0.000001) since the plants reached greatest
cm) than any other combination of levels height on interaction with S. americanus
of the two factors while non-burned plants (137 cm) and presented the shortest height
of the same competition treatment were the with P. australis (118 cm). Accumulation
shortest (80 cm). of aerial biomass (fig. 1h) also responded to
nitrate addition (F(1,78) = 12.0, P < 0.0008),
Regarding aerial biomass (fig. 1e), S. and to plant-plant interactions (F(2,78) =
americanus responded to the addition of 14.5, P < 0.0001), since the plants in intras-
nitrate (F(1,78) = 43.1, P < 0.000001) and pecific treatments accumulated the least
to competition (F(2,78) = 56.2, P < 0.001), biomass on average (3 g), followed by plants
accumulating more biomass in the inters- grown with P. australis (5.8 g) and finally
pecific treatment with P. australis (4.0g) those grown with S. americanus (9.0 g);
relative to the other two treatments (2.4 g the accumulation of root biomass only res-
with T. domingensis and 2.3 g in intraspe- ponded to plant-plant interactions (F(2,78)
cific competition). In the burning treatment, = 7.5, P = 0.001), and followed the same
more aerial biomass was accumulated in the pattern observed in the aerial biomass; 11.8
non-burned plants (3.35 g) than in burned g in intraspecific competition, 14.1 g with
plants (2.5 g). Regarding root biomass (fig. P. australis and 23.9 g with S. americanus).
2f), S. americanus presented the most com-

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Montejo-Mayo, W. et al.: Interactions between dominant hydrophytic species of the Wetlands of Western Mexico

DISCUSSION diminished growth, since taller and more

robust plants were produced when grown
The results show, as expected, that all the in the presence of individuals of the same
study species responded positively to the species. For T. domingensis, the contrary
addition of nitrate; however, in the pre- was true as the presence of S. americanus
sence of other individuals, performance benefited its performance after burning and
variables differed depending on species. for S. americanus, the plants most favored
Interspecific interactions are not important were those grown with T. domingensis. The
for Phragmites australis and Schoenoplectus positive effects of ash from S. americanus in
americanus since they present the same the growth of T. domingensis seedlings have
rates of growth as when they are growing been reported (Lopez-Arcos et al., 2012). It
alongside individuals of their own species. therefore appears that competition between
For Typha domingensis, the situation is these two species is less when growing to-
different since intraspecific interactions gether and there is a likelihood of fire. The
were more detrimental than interspecific modification of these competitive interrela-
ones. Given that P. australis is invading the tions by anthropogenic disturbance, reported
wetland, it was expected that P. australis in previous studies for other wetland species
would be a superior competitor to both T. (Woo and Zedler, 2002) as being of benefit
domingensis and S. americanus and would to those species with invasive attributes, was
suppress their performance, but this did not not reflected in this experiment. It is worth
in fact occur. The addition of nutrients alone noting that P. australis is always taller and of
cannot explain the increase in abundance greater biomass than the other two species,
of P. australis in the Mintzita wetland, as making it likely in the long term that compe-
nutrients do for other invasive species in tition for light becomes a determining factor
other wetlands (Woo and Zedler, 2002). In in its invasion of the site, and therefore this is
the case of other species that present inva- a factor that must be fully taken into account
sive behavior, it has been documented that in any future studies to predict the behavior
they require anthropogenic disturbances in of this invasive species.
order to increase their numbers (Jesson et
al., 2000; Fireswyck and Zedler, 2007). CONCLUSIONS

In this experiment, the added disturbance The series of experiments indicate that nitrate
is related to the burning of vegetation, a availability altered the intensity of plant-
practice that currently takes place in Mint- plant interactions but that fire was more
zita (Escutia-Lara et al., 2009a). With this instrumental in altering these, in particular
disturbance, the plant response to nutrients that presence of Schoenoplectus benefited
remained surprisingly unchanged; the three Typha. Since several species of the later
species grew in line with the increases in genus have shown invasive behavior, this
nitrate concentration. However, the inter- study suggests an additional mechanism that
relations associated with plant-plant inter- deserves more attention under natural condi-
actions did change. For burned P. australis tions, since the controlled conditions of the
plants, interspecific interactions caused experiments undertaken limits its realism.

159
 Núm. 40: 153-161 Agosto 2015

Finally, fire was a significant factor altering Escutia-Lara, Y.; E. de la Barrera, Y. Martí-
growth for Phragmites and Schoenoplectus, nez de la Cruz, y R. Lindig-Cisneros,
benefiting more the former than the later, 2010. “Respuesta a la adición de ni-
therefore, strongest competition occurred trógeno y fósforo en el crecimiento de
between natives, leaving Phragmites, the Typha domingensis y Schoenoplectus
invasive, free to respond to increased nitrate americanus”. Boletín de la Sociedad
availability by growing more. Botánica de México, 87: 93-97.

A CKNOWLEDGEMENTS Escutia-Lara, Y.; M. Gómez-Romero, y R.

Lindig-Cisneros, 2009b. “Nitrogen
This work was funded by the Universidad and phosphorus effect on Typha do-
Nacional Autónoma de México through the mingensis Presl. rhizome growth in a
project PAPIIT IN203608 and CONACYT matrix of Schoenoplectus americanus
(SEP-CONACYT-2008-101335). (Pers.) Volkart ex Schinz and Keller”.
Aq. Bot., 90: 74-77
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Recibido: 17 octubre 2013. Aceptado: 23 enero 2015.

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