In this tutorial we are going to deal with recognizing one of the most basic patterns of a plant population: whether individuals are spatially closer or further apart than would be expected if they were simply randomly distributed (i.e., the location of an individual does not improve the prediction of where other individuals may be)).

Investigate the spatial pattern in plant populations and discuss what underlying processes could generate the observed patterns. First of all, however, we need to define some concepts.

A spatial pattern is a predictable structure that can be detected and quantified. In general, a pattern is considered to be a different structure than a random one, however, in the case of spatial patterns (and others as well) the random pattern can also be considered a pattern, after all it has some predictability (eg in relation to the average number of individuals)) and can be detected and quantified. There are several metrics used to describe the distribution of individuals that are able to differentiate, with greater or lesser efficiency, the three basic spatial patterns: random, homogeneous and aggregated.

Detecting a spatial pattern can be important both for understanding the mechanisms that generate the pattern, for deciding on the sampling method and scale, and for planning the management of a population. Some desirable properties of a spatial pattern measure are:

• clearly differentiate the pattern;
• not be affected by: sample size, population density, or variation in sample size and shape;
• be statistically tractable: able to calculate the uncertainty of the value and test for differences between samples.

For this practice we will use a point randomness estimate called K-Ripley. First we will use simulated distribution data with different patterns and then use the same technique to detect the spatial pattern in a natural population.

In this practice we will quantify the spatial pattern using a multiscale method. Multiscale methods allow, with a single metric, to assess how the spatial pattern varies with scale. We will describe the spatial pattern for the total set of individuals in a population in a delimited area and we will assess the pattern from the scale of the neighborhood of individuals to the wider scale of the population.

For practice we will use a little program called Programita, made by researcher Thorsten Wiegand to quantify spatial patterns using multiscale measures based on distance between points. To download the Programita manual click

here.

In Programita there are several measurements that can be used to calculate the spatial pattern, we will use two of them: Ripley's L and O-ring.

Both are point-based approaches, which use the calculation of point-to-point distances within a bounded area. These measures can be used for univariate analyses, that is, identifying the pattern for a single class of points, or for bivariate analyses, which identifies the pattern between two types of points. Bivariate analyzes can be used in the context of populations to verify whether individuals of a given stage are spatially associated with another, or in the context of community structuring to analyze whether there is attraction or repulsion in the occurrence of one species in relation to another.

Ripley's K is a measure of the average density around each point. For each point in the study area, the density inside a circle of radius r centered on the point (gray area of ​​the figure) is calculated. Then, an average of these values ​​obtained for all points is calculated.

The operation is repeated for different r values, which allows continuous evaluation of the K value for different scales.

$$ K_{(r)} = \frac{\sum_{i\neq{j}}^{i}I({d_{ij}<r})}{n}\frac{1}{\lambda} $$

Where:

• $d_{ij}$ is the distance from point $i$ to point $j$;
• $I({d_{ij}<r})$ indicator function, being 1 if the point $j$ is at a distance less than $r$ from the point $i$ and 0 if the point $j$ is outside this radius $r$ around $i$;
• $n$ is the total number of points;
• $\lambda$ is the density of the points.

The visual interpretation of K is not very intuitive as it is a cumulative function associated with the area of ​​the circle relative to r. Ripley's L, in turn, is the standardization of this value:

$$ L_{(r)}= (\sqrt{\frac{K_{(r)}}{\pi}}-r) $$

This transformation makes the value of L for a completely random distribution always be 0 and L > 0 indicates aggregation, while L < 0 indicates homogeneous pattern.

The O-ring stat is similar to Ripley's L, but based on a ring rather than a circle. It is measured by counting the number of points in a ring of radius r and fixed width. In the same way as L-Ripley, the intensities for different ring sizes are also calculated, keeping the width fixed.

So we define $O(r)$ as: $$ O_{(r)} = L_{(r)} - L_{(r-l)}$$

Where:

• $r -l$ : is the radius minus the width of the ring ¹⁾

In complete spatial randomness $O(r) = \lambda$ (pattern intensity), when the pattern is aggregated $O(r) > \lambda$ and when it is homogeneous $O(r) < \lambda$

• 1. Download the files related to spatial pattern 1 or 2. If open a page showing the data, right-click the link to save the file to your computer:

• if you don't have programita installed, download programita here in the same folder as the data file;
• unzip the file programita.zip;
• 2x click to open the executable file ProgramitaJulio2006.exe.

Welcome to Programita! Now let's open the data that we are going to work with.

Programita accepts text files with .dat and .asc extensions. They are files in text format, separated by tabs (or spaces). The data file has the following structure:

The first line contains general information about the data file:

• minimum value of x;
• maximum value of x;
• minimum value of y;
• maximum value of y; and
• total number of individuals

From the second line on, are the data of the points that will be analyzed:

• first column with the x coordinates of the individuals;
• second column with the y coordinates of the individuals;
• third column with points of pattern 1 identified by 1 and pattern 2 by 0 ³⁾;
• fourth column with the points of pattern 1 identified by 0 and pattern 2 by 1 ⁴⁾.

In the case of univariate data, the third column will always be 1 and the fourth column will always be 0. For bivariate data the third and fourth columns will have values ​​of 0 and 1 according to the point pattern.

• 1. Check if the Input data file window is showing the .dat files. If not, check that the programita executable file is in the same folder as the .dat files.

• 2. in the menu on the left select the file padrao“0X”all.dat. In the case X will be 1 or 2 depending on your choice;

• 3. In How ​​your data are organized select List
• 4. Let's start using Ripley's L so in Which method to use select Circle
• 5. In Select modus of data select List with coordinates in the grid. When selecting this option, a window will appear with the option Select a new cell size:

• 6. If you have less than 500 points, change the proposed cell size to 1. Otherwise, leave it at the program's default.
• 7. Done all that, you should be like this:

• 8. You can now take a deep breath and hit the Calculate index button;

The program's visual output is a map where individuals appear in red dots, following the coordinates of the data file. The graph in the upper right corresponds to the L-Ripley value for different radii. In this graphic output it is possible to analyze how the spatial pattern varies according to the scale.

However, this is not enough to state at what scales the population is aggregated. For this we need to compare the observed result with the pattern that would be generated by the completely random distribution of points. This null model is called complete spatial randomness. To generate this model by simulation it is necessary to relocate the same number of points randomly in the same area. If we do this over and over again, it is possible to generate a confidence envelope (similar to the confidence interval) in which the random distribution pattern is found. If the observed values ​​are contained within the envelope we can conclude that our pattern is not different from the random one.

To do this you must:

• 9. select the option Calculate confidence limits and;
• 10. in the window Select a null model select the null model Pattern 1 and 2 random;
• 11. Make sure your screen looks like the picture and click the Calculate index button again.

• 12. Do the same procedure, but in Which method to use select Ring
• 13. Compare the results between the L-Ripley and the O-Ring.

O Programita permite a análise de padrão de pontos de uma classe em relação a outra. Para isso é necessário diferenciar os pontos no arquivo de dados, utilizando 0 ou 1 nas colunas 3 e 4, como mostra a figura abaixo, em um arquivo que distinguia indivíduos adultos de juvenis:

Vamos agora analisar o padrão dos pontos associados (PROLE) em relação aos parentais (PAR), seguindo o mesmo procedimento anterior.

• 1. selecione o arquivo com a separação de classes de pontos parentais e associados: padrao“0X”bi.dat;
• 2. em What do you want to do selecione a opção Point-pattern analysis
• 3. em How your data are organized selecione List
• 4. neste caso, estamos interessados na análise do padrão em escala cumulativa para entender até que distância há agregação, por isso, em Which method to use selecione L-Ripley
• 5. em Select modus of data selecione List with coordinates no grid
• 6. para testarmos se existe agregação dos pontos PROLE em relação ao PAR , utilizaremos o envelope de confiança. selecione a opção Calculate confidence limits e selecione o modelo nulo Pattern 1 fix, 2 random.
• 7. rode a análise apertando: Calculate index
• 8. interprete os resultados.

O Palmiteiro (Euterpe edulis Mart.) é uma espécie muito característica das florestas atlânticas e costuma ocorrer com densidades altas em áreas mais preservadas. Vamos agora analisar os dados referentes a uma população de palmitos que ocorre em uma parcela de floresta de Restinga na Ilha do Cardoso, Cananéia -SP. Os dados foram coletados nos anos de 2009/2010 em uma área de 10,24ha (320m x 320m).

Preparamos três arquivos no formato lido pelo Programita:

1. dados de indivíduos juvenis (diâmetro do tronco entre 1 e 5 cm): juvenil.dat
2. dados de indivíduos adultos (diâmetro do tronco > 5 cm): adulto.dat
3. juvenis e adultos (padrão 1 adulto, padrão 2 juvenil): juvenil_adulto.dat

Utilizando as ferramentas disponíveis no Programita para descrever os padrões espaciais:

• da população total de palmito;
• apenas dos juvenis e;
• apenas dos adultos.

Investigue se a distribuição dos juvenis está associada a dos adultos.