Alien pine changing plant community structure in a coastal dune ecosystem

Felcia Miranda Fischer, Juliano Morales de Oliveira2 & Valrio de Patta Pillar1
1 Universidade Federal do Rio Grande do Sul, Quantitative Ecology 2 Universidade do Vale do Rio dos Sinos, Vegetal Ecology

feliciafischer@yahoo.com.br
Aims Our aim is to evaluate how Pinus taeda invasion is related to topography, vegetation structure and composition on costal psammophilous grasslands in southern Brazil. In this sense, we hope to better understand the invasion process and its effects on the plant communities. Methods
distancia na composio residual Residual composition distance

We tested three regression models using variable response the Euclidean distance in species composition and as predictor variables (1) the individual's age of P. taeda, (2) age and residual pine needle litter coverage (the variation of soils needle cover not explained by the individual age), and (3) age, needle coverage and residual crown diameter (range in diameter not related to age and needle coverage). Results Most of the sampled points, with or without invasion, occurred in the relief type marshland, with few points on the slopes and tops. We could not find any points for invasion of P. taeda in the top position. But there was no significant difference between positions in relief for the presence of P. taeda. We also found no significant difference in the coverage percentage of shrub and herbaceous between points with and without P. taeda. Was found in the survey a total of 47 non graminoids species. We found a significant difference in species composition among plots under P. taeda and control plots (P = 0.005). The ordination of species composition data using principal coordinates analysis (Fig. 2) separated plots under individuals of P. taeda and control plots, and this distinction became even more evident as increased age of the individual. We realized that the most abundant species in control plots were generally negatively correlated with axis 1, while those more abundant under the canopy of P. taeda are positively correlated with the same axis. This shows that this ordination axis expressed composition variation explained by the presence of P. taeda. The partial regression models (Fig. 5) indicated the difference in species composition between control plots and plots under P. taeda increased with the age of individuals (Fig. 5a) and with needle ground cover (Fig. 5b), but was not influenced by pine crown diameter (considering the portion of variance unrelated to the former factors). Pine age and needle cover explained about one third of composition distance variation (R2= Ocorrncia de espcies 0.32; P=0.108). 3
50 40

5 Composition distance distncia na composio

y = 0,0657x + 1,7862 R2 = 0,238

0 0 5 10 15 20 25 idade (anos)

The study was conducted at Itapeva State Park, a natural protected area, in the coastal region (29 20'S and 44 45'W). The study area was a psammophilous grassland in wet lowlands, formed by sandy deposits of Holocene origin, parallel to the coastline. The area was located between the foredunes and mobile dunes inland. The area presents dune mounds, slightly undulating, covered with varied grass and shrub density. Sampling points were located at each vertex on a grid of 100 x 100 m in the entire area of the wet lowlands of the park, totaling 121 points. In a radius of 25 m around each point we recorded the presence of P. taeda individuals. Next, we considered only those sampling points of the grid with the presence of P. taeda. In each of these points we defined two plots of 10 x 10 m, one centered on each of the individuals of P. taeda and another random control plot. In each plot we described the environment on the micro relief, which was classified as top, marshland and hillside. We tested whether plots in different positions in the landscape differed as to the presence of P. taeda, using analysis of variance with randomization.

Age (years)
3 2 1 0 -1 -2 -3 -40 y = 0,014x + 0,1997 R2 = 0,0643

-20

20

40

60

80

acculas needles Residual pineresidual (%) litter (%)

Figure 5: linear regression model to explain the variation in floristic composition on plots with and without Pinus taeda. a) composition distance due to individuals age. b) Distance from residual composition (by removing the part that is explained by age) for soil covering by residual needles litter (by removing the part that is explained by age). Model considering the two predictor variables: distance on the composition x = 1.8 + 0.07 + 0.01 x age residual needles. R = 0.32 P = 0.108.
Discussion The lack of significant difference between the positions of points in relief with P. taeda may be due to the small number of sampled points on the slopes and tops, or the establishment of P. taeda is indifferent to the positions in the field, on this scale evaluated. The result of comparing the points with and without P. taeda as the density of herbaceous vegetation and shrubs may indicate that there is no influence of native vegetation on the establishment and development of P. taeda, or that resources and conditions that define the density of vegetation (both herbaceous shrub about) are not the same determinants of invasion of P. taeda. This result contradicts results found in other studies, where coverage of shrubs reduced the spread of Pinus sp. in fields. Photosynthesis rates of P. taeda are related to soil moisture and light, and growth is inhibited by shading in forests interior. Vegetation density does not appear to be sufficient to inhibit invasion of P. taeda. This study proved that there is a noticeable change in floristic composition in the area of direct influence of individuals of P. taeda. The results suggest that these organisms generate a microenvironment protected from sun and wind, and possibly provide perches for birds dispersing. Thereby it provides the establishment of tree species (as Myrsine parvifolia and Sapium glandulosum) and / or shade tolerant (as Rumohra adiantiformis) and creepers plants that require a more developed woody environment (as Smilax campestris). The presence of Pinus taeda also appears to inhibit the growth of grassland dependent species of high incidence of sunlight, as Tibouchina urbanii and Asteraceae species, like Pterocaulon angustifolium and Baccharis radicans. According to the regression model, age and litter needle cover are factors that explain the difference in species composition between plots located under the canopy of P. taeda and control plots. The relation with age indicates that the older the individual of P. taeda longer the time that this is influencing the community to which it belongs. But differences in composition are due by the presence of P. taeda or if the difference is related to preexisting conditions that could affect both? In this study, the results show that, at least in density, vegetation does not vary according to the presence of P. taeda (in a portion larger than the canopy). While under the canopy floristic composition varies. Corroborating this result, we proved that the time of establishment and the amount of pine needles litter are factors related to change in floristic composition, a strong evidence it is the individual of P. taeda that causes changes in the floristic composition. Conclusion The native vegetation has not shown resistance to the establishment of individuals of P. taeda. This alien species alter the natural vegetation, facilitating the development of some species and compressing the other in an ever smaller area. This process may even locally extinct some species, if they are confined to areas invaded by P. taeda.

Figure 1: Coastal psamophilous grassland with invasion of Pinus taeda. Figure 2: Localization of Itapeva State Park in South America and the sample units in the area.

30 20 10 0
od o Ac na e a m Pt vi el er s la oc de c os au a cu lo m n b an Ti bo gu e ns uc st i hi na fo liu Ep m ur id vi en ll e dr um an Ti a bo fu u lg en Ba chi na s cc ha ur ba ri s ni ra i D C di es en ca m t ns od el la iu as m ad i atic C or sc a di en a R d um cu ra e ns oh ss ra a d av ic Sm ia a ila nti fo x c a rmi M s m yr pe si ne st PA pa ri s rv R i C EL foli a A V AZ IA
Control controle P. taeda P.taeda

In the plots, we also assessed the percentage of soil covered by shrubs (up to 1 m high) and herbs (lass then 1 m height). We compared the average cover of herbs and shrubs in plots with and without P. taeda at each point. For this we applied paired analysis of variance with permutation test by restricting the permutations within each pair. Furthermore, we selected a total of 15 individuals of2,5 P. taeda for age determination, for canopy diameter measure and 2,0 for evaluation of pine needle litter deposition and also its effects 1,5 on plant community. For determining the age of each individual of P. taeda we toke trunk cross sections. In laboratory, the cross 1,0 sections were treated and we counted each growth ring as one year, and in this way determined the year of the individual 0,5 establishment. 0,0 Under each tree, we estimated the ground cover by pine needles litter, in 4 plots of 1x1m, under its canopy. For -0,5 evaluation of community composition, we marked a 1 x 1 m plot under the canopy and another paired plot of the same size -1,0 randomly located. In each plot the composition of non graminoids -1,5 species was accessed. We used multivariate analysis of variance -1,5 with permutation test, restricting the permutations within each pair (with P. taeda and control) to verify whether communities under the canopies were different from those outside canopies. We also explored patterns of species composition by means of ordination by principal coordinates analysis. Further, with the species composition data we calculated the Euclidean distances for each pair of plots. We used linear regression analysis with randomization test to evaluate how this difference in composition varied according to the age of the pine individual, ground cover by needles, and canopy diameter.
PCoA 2 (13%)

Figure 3: Species with at least 20% of occurrence in any of the treatments (P.taeda and control) plots and percentage of plots without any species.
Controle Pinus
Pinus 9 9

2,5
2,5

Controle Control

Pinus P. taeda 9

Controle

2,0
2,0

12
12

1,5
1,5

12 10
10

PCoA 2 (13%)

1,0
PCoA 2 (13%) 1,0

12 12
2
2

12

Desmodium barbatum (0,66) Tibouchina urvilleana (0,65) Epidendrum fulgens (0,62) Baccharis spicata (0,60) Gamochaeta americana (0,60) Macroptilium prostratum (0,60) Dodonaea viscosa (0,60)

5 3 3 1 2 1

5 5 15 13 15 13 10 11 33 5 10 11 5 66 14 13 8 6 3 3 8 8 10 14 138 6 44 1515 0,0 0,0 44 77 2 1 12 2 14 14 7 1 7 15 -0,5 131 -0,5 10

0,5 -1,0

0,5

8-1,0

6 14

11 4

138 6 7

15

11

11
9

-1,5 -1,5 -1,0 -1,5 -1,0

14 7 -1,5

9
0,0 0,5 1,0 1,5 2,0

Centella asiatica (-0,44) Rumohra adiantiformis (-0,40) Myrsine parvifolia (-0,39) Smilax campestris (-0,36)
2,0

-0,5

-0,5

0,0 0,5 PCoA 1 (15%)

1,0

1,5

11 PCoA 1 (15%) 9

Pterocaulon angustifolium (-0,68) Baccharis radicans (-0,5) -1,0 Tibouchina urbanii (-0,49) -0,5 0,0 0,5 1,0 1,5
PCoA 1 (15%)

Smilax campestris (0,62) Myrsine parvifolia (0,60) 2,0 Rumohra adiantiformis (0,55) Sapium glandulosum (0,50)

Figure 2: Principal coordinates analysis (PCoA) ordination for non-graminoid species composition in psammophily vegetation, individuals under the canopy of Pinus taeda and outside the canopy (control). Axes 1 and 2 explain 15% and 13% of total variation. The size of the dots representing the tree age. Its also indicated the species with greater correlation with the axis.

QUANTITATIVE ECOLOGY