34

Estimating Plant Population, Dispersion, and Association Part A - Data Collection

Introduction. One of the most fundamental problems faced by community and population ecologists is that of measuring population sizes and distributions. These data are important for comparing differences between communities and species. They are necessary for impact assessments

(measuring effects of disturbance) and restoration ecology (restoring ecological systems). They are also used to set harvest limits on commercial and game species (e.g. fish, deer, etc.). In most cases it is either difficult or simply not possible to census all of the individuals in the target area. The only way around this problem is to estimate population size using some form of sampling technique. There are numerous types of sampling techniques. ome are designed

for specific types of organisms (e.g. plants vs. mobile animals). !s well there are numerous ways of arriving at estimates from each sampling technique. !ll of these procedures have advantages and disadvantages. In general, the accuracy of an estimate depends on ") the number of samples ta#en, $) the method of collecting the samples, %) the proportion of the total population sampled. ampling is viewed by statistical ecologists as a science in its own right. In most cases, the ob&ect is to collect as many randomly selected samples as possible (so as to increase the proportion of the total population sampled). The accuracy of an estimate increases with the num er of samples ta!en. This is because the number of individuals found in any given sample will vary from the number found in other samples. 'y collecting numerous samples, the effect of these variations can be averaged out. The purpose for collecting the samples randomly is to a"oid iasing the data. (ata become biased when individuals of some species are sampled more frequently, or less frequently, than e)pected at random. uch biases can cause the population size

to be either over estimated or under estimated, and can lead to erroneous estimates of population size. *opulation size generally refers to the number of individuals present in the population, and is self+e)planatory. (ensity refers to the number of individuals in a given area. ,or ecologists density is usually a more useful measure. This is because density is standardized per unit area, and therefore, can be correlated with environmental factors or used to compare different

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populations. The spatial distribution of a population is a much more complicated matter. 'asically, there are three possible types of spatial distributions (dispersions) (see diagrams below). In a random dispersion, the locations of all individuals are independent of each other. In a uniform dispersion, the occurrence of one individual reduces the li#elihood of finding another individual nearby. In this case the individuals tend to be spread out as far from each other as possible. In a clumped dispersion, the occurrence of one individual increases the li#elihood of finding another individual nearby. In this case, individuals tend to form groups (or clumps).
o o

o oo oo o o o

o o

o oo

o o oo ooooo oo o o o ooo oo ooo Clumped

o o o o o

o o o o o

o o o o o

o o o o o

Random

Uniform

-cologists are often interested in the spatial distribution of populations because it provides information about the social behavior and.or ecological requirements of the species. ,or e)ample, some plants occur in clumped distributions because they propagate by rhizomes (underground shoots) or because seed dispersal is limited. /lumped distributions in plants may also occur because of slight variations in soil chemistry or moisture content. 0any animals e)hibit rather uniform distributions because they are territorial (especially birds), e)pelling all intruders from their territories. 1andom distributions are also common, but their precise cause is more difficult to e)plain. 2nfortunately, it is often difficult to visually assess the precise spatial distribution of a population. ,urthermore, it is often useful to obtain some number (quantitative measure) that describes spatial distribution in order to compare different populations. ,or this reason, there are a variety of statistical procedures that are used to describe spatial distributions. /ommunities are assemblages of many species living in a common environment. Interactions between species can have profound influences of their distributions and abundances.

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/omprehensive understanding of how species interact can contribute to understanding how the community is organized. One way to loo# at species interactions is to evaluate the level of association between them. Two species are said to be positively associated if they are found together more often than e)pected by chance. *ositive associations can be e)pected if the species share similar microhabitat needs or if the association provides some benefit to one (commensualisms) or both (mutualism) of the species involved. Two species are negatively associated if they are found together less frequently than e)pected by chance. uch a situation

can arise if the species have very different microhabitat requirements, or if one species, in some way, inhibits the other. ,or e)ample, some plants practice allelopathy, the production and release of chemicals that inhibit the growth of other plant species. !llelopathy results in a negative association between the allelopathic species and those species whose growth is inhibited.

E#ercise In this lab we will use the quadrat technique to estimate population size, spatial distribution, association among three species of plants occurring in a heavily grazed pasture (see map of how to get to the study site). 3e will try to use /ommon 'roomweed ( Gutierrezia dracunculoides), 3estern 1agweed (Ambrosia psilostachya) and 4arrow+leaf umpweed (Iva angustifolia). 5owever, due to variations in weather (droughts, floods, etc.) we may have to use other species. 6ou will be notified of any changes at the time of the lab e)ercise. !bout the study species 7 Information about the ta)onomy and ecology of these study species can be found in hinners and 0ahler8s 9Illustrated ,lora of 4orth+central Te)as: online at; http;..artemis.austincollege.edu.acad.bio.gdiggs.shinners.html. In particular, details on the biology of our three study species can be found by searching for the appropriate genus in the pdf file at; http;..artemis.austincollege.edu.acad.bio.gdiggs.4/T<=$>pdf.,4/T=$>>$">+>?"@.pdf. !dditonal information on each species is also readily available on the web or by consulting boo#s on the Ath floor of the library.

0ethod; The Buadrat Technique. I$P%&TA'T '%TE( In case of rain + ma#e sure you bring appropriate clothing. 'ring a

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plastic bag to #eep your data sheets in and ma#e sure you record all your data in pencil. )o to great lengths to ma!e sure that your paper stays dry* A'%T+E& I$P%&TA'T '%TE( ,ield wor# necessarily entails certain hazards. If you have allergies, ta#e appropriate precautions before hand. 3atch for fire ants, stinging insects, and venomous reptiles (3e have not encountered a live venomous reptile on this study site to date 7 there once was a dead cottonmouth on the road, however). The quadrat method is used primarily in studies of plant populations, or where animals are immobile. The principal assumptions of this technique are that the quadrats are chosen randomly, the organisms do not move from one quadrat to another during the census period, and that the samples ta#en are representative of the population as a whole. It is often conducted by dividing the census area into a grid. -ach square within the grid is #nown as a quadrat and represents the sample unit. Buadrats are chosen at random by using a random number generator or a random number table to select coordinates. The number of individuals of the target species is then counted in each of the chosen quadrats. %ur technique will be to randomly choose quadrats by throwing metal quadrats in random directions. 3here the quadrat lands will mar# the location of an individual sample point. The number of individuals of each of the three species within the quadrat will then be counted and recorded. 'e sure to count all individuals whose stems originate from within the quadrat and whose stem originate from under the edge of the quadrat. If you are using the small metal quadrats, census ?> quadrats. If you are using the large quadrats, census C> quadrats. 3hen you are finished hand in your data to me, so that I can tabulate them for you. The tabulated data will be distributed to you in class.

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Estimating Population ,i-e and Distri ution Part . - Data Analysis

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Three things are to be accomplished in this lab; /0 estimate the population sizes of /ommon 'roomweed, 3estern 1agweed and 4arrow+leaf umpweed using large and small

quadrats and using A> and $>> quadrats (D estimates), 10 statistically analyze the dispersion pattern of /ommon 'roomweed, 3estern 1agweed and 4arrow+leaf umpweed using large and small quadrats and using A> and $>> quadrats (D analyses), and 20 determine the degree of association between 3estern 1agweed and /ommon 'roomweed, between 3estern 1agweed and 4arrow+leaf umpweed, and between /ommon 'roomweed and 4arrow+leaf umpweed (% measures of association)

/0 Estimating population si-e

To determine population size, first determine the average number of individuals of each species per quadrat (mean). (o this for each of the three species and for sample sizes of A> and $>> quadrats. ,or estimates using A> quadrats simply use quadrats " to A> from your data sheets. The data should be typed into an -)cel 3or#boo#. Once this is done it is a simple matter of clic#ing on the paste function #ey and selecting A"erage from the statistical function menu. The size of the study area was "$"?> m$. The large quadrats were ".> m$ in size, the small quadrats were only >." m$ in size. Therefore, different calculations are used to estimate population size (4T) depending on what quadrat size was used. 3or small quadrats( 3or large quadrats( 'T 4 $ean 5 /.1/6 5 /78 'T 4 $ean 5 /.1/6 5 /79

/alculate the total populations of /ommon 'roomweed, 3estern 1agweed, and 4arrow+ leaf umpweed in the study area using small quadrats and large quadrats and using A> quadrats and $>> quadrats. 6ou should be able to fill in the following tables;

Ta le /. Estimates of population si-e for Common .roomweed using sample si-es of 87 and 177 and using small and large quadrats.

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ample ize (4umber of Buadrats) A> $>>

Buadrat ize >." m$ ".> m$

Ta le 1. Estimates of population si-e for :estern &agweed using sample si-es of 27 and /77 and using small and large quadrats. ample ize (4umber of Buadrats) A> $>> Buadrat ize >." m$ ".> m$

Ta le 2. Estimates of population si-e for 'arrow-leaf ,umpweed using sample si-es of 27 and /77 and using small and large quadrats. ample ize (4umber of Buadrats) A> $>> Buadrat ize >." m$ ".> m$

10 Estimating population dispersion ;Te#t Pages <82 = <890 3e will use the variance (s$) to mean ratio to measure population dispersion. The

variance to mean ratio is based on the following line of logic. If a population is uniformly distributed, one should find about the same number of individuals in every quadrat measured. In this case there will be little or no variation between quadrats in the number of individuals counted (s$ E >). Thus, for uniform distributions the variation will be much smaller than the mean and the ratio of s$.mean will approach zero. If a population has a clumped distribution you should find some quadrats containing a large number of individuals and many quadrats that are empty. In this case the variation between

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quadrats will be very high, and the s$.mean ratio will be large. The values of s$.mean and their interpretation are as follows;

s1>mean ? /.7 4 clumped distri ution s1>mean 4 /.7 4 random distri ution s1>mean @ /.7 4 uniform distri ution

The variance to mean ratio allows us to test the statistical significance of the dispersion pattern. ,or e)ample, given that our measurements are made with some error, if s $.mean F ".$, would this really indicate a clumped dispersion patternG To test whether this is significantly different from a random dispersion (i.e. s$.mean F ".>) we calculate a test statistic, in this case t, where;

tF

H (s$.mean) 7 ".> H $.(n 7 ")

3here n F sample size (number of quadrats sampled). ,or our purposes, any absolute value of t I ".J? indicates a distribution that differs from random with a certainty of JA=. ,or your lab reports you must /0 calculate the variance to mean ratio and identify the dispersion pattern indicated, 10 calculate the value of t, and 20 determine whether the dispersion is significantly different from random. 6ou must do this for both sizes of quadrats and for sample sizes of A> and $>> quadrats. 6ou should already have your means calculated from the first part of this lab e)ercise. 4ow you need only calculate s$. These easiest way to do this is to select Aar from the statistical function menu in -)cel. /alculate the s$.mean ratio for each species of plant and for A> and $>> quadrats. 4e)t determine the significance of the variance to mean ratios by calculating t. Kust insert the values into the formula;

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tF

H (s$.mean) 7 ".> H $.(n 7 ")

If t I ".J? then the dispersion is different from random. following tables;

ummarize your data in the

Ta le 9. Estimates of dispersion for Common .roomweed using sample si-es of 87 and 177 and using small and large quadrats. ample (etails A> small Buadrats $>> small Buadrats A> large Buadrats $>> large Buadrats s$.mean ratio Indicated dispersion t ignificance (yes or no)

Ta le 8. Estimates of dispersion for :estern &agweed using sample si-es of 87 and 177 and using small and large quadrats. ample (etails A> small s$.mean ratio Indicated dispersion t ignificance (yes or no)

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Buadrats $>> small Buadrats A> large Buadrats $>> large Buadrats

Ta le 6. Estimates of dispersion for 'arrow-leaf ,umpweed using sample si-es of 87 and 177 and using small and large quadrats. ample (etails A> small Buadrats $>> small Buadrats A> large Buadrats $>> large Buadrats s$.mean ratio Indicated dispersion t ignificance (yes or no)

20 $easuring ,pecies Associations ;177 large quadrats only0 ;Te#t Pages <89 = <860

If two species are positively associated then we should find them together more frequently than would be e)pected simply based on their overall abundances. 'y corollary, if two species are negatively associated then we should find them together less frequently than would be e)pected based simply on their overall abundances. 3e can test these predictions by evaluating the number

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of quadrats in which two species were observed together. ,or this analysis you must include all $>> sample points collected from " m$ quadrats. 6ou will want to conduct three separate analyses; 3estern 1agweed vs. /ommon 'roomweedL 3estern 1agweed vs. 4arrow+leaf

umpweedL /ommon 'roomweed vs. 4arrow+leaf umpweed. ,or each analyses the data should initially be organized into a table li#e the one below; pecies ! M pecies ' M + a c aMc + b d bMd aMb cMd aMbMcMd

,or each pair of species enter the number of quadrats containing both species in cell 9a:, the number of quadrats containing only species ! in cell 9c:, the number of quadrats containing only species ' in cell 9b:, and the number of quadrats containing neither species in cell 9d:. 3e can then use these data to calculate /, the Coefficient of Association. The coefficient of

association varies from M".> for a ma)imum of positive association to 7".> for a ma)imum negative association. ! value of >.> indicates that the degree of association observed is that which would be e)pected by chance (i.e. the species are randomly distributed with respect to each other). The following formulas are used to calculate the coefficient of association; If bc I ad and d I a, then / F (ad 7 bc).((aMb) (aMc)) If bc I ad and a I d, then / F (ad 7 bc).((bMd)(cMd)) If ad I bc and c I b, then / F (ad 7 bc).((aMb)(bMd)) If ad I bc and b I c, then / F (ad 7 bc).((aMc)(cMd)) 5aving calculated the indices, we now need to determine whether these indices deviate significantly from random (>.>). ! /hi+square test is used for this purpose (are you having a genetics dN&O vu e)perienceG). /hi+square tests first require that you calculate e)pected values for each cell in the table above. The following formulas are used to calculate the e)pected values for cells a, b, c, and d; -)p a F (aMb)(aMc).n

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-)p b F (aMb)(bMd).n -)p c F (cMd)(aMc).n -)p d F (cMd)(bMd).n *ut your observed and e)pected frequencies into a table li#e the one below (one for each species pair); pecies ! M Obs pecies ' M + -)p Obs + -)p

3e could calculate the test statistic, X2, as the sum of (obs+e)p)$.e)p, but it is easier to use the single formula below (which includes 6ate8s continuity correction for a $<$ table);

X2 F

((Had+bcH + >.An)$ (n)) ((aMb) (aMc) (bMd) (cMd))

!s before, n F the number of quadrats sampled ($>>).

This chi+square statistic is

associated with " df. Thus, if the value of X2 is greater than %.DC, then the value of / significantly deviates from random. (etermine X2 for each of the measures of association calculated and assess their significance.

:riting Bour &eports 3rite your report with the usual headings (!bstract, Introduction, 0ethods, etc.). 'riefly describe the methods used to collect the data. 0ention that the data were used to calculate population size, variance to mean ratio, the significance of the variance to mean ratio, the coefficients of association, and the significance of the coefficients of association. (o not include a long+winded description of how you performed your calculations. 6our results should include four tables (tables " 7 ? if you are following the format of this

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handout) plus three tables (tables @ 7J) showing observed and e)pected values of association for each analysis of interspecific association. In the written section identify and briefly describe the differences in population sizes among species and e)plain how the results vary according to quadrat size and sample size. 1efer to your tables and briefly mention which measures of dispersion are significant and whether these significant measures indicate uniform or random dispersions. 'riefly describe the differences in results obtained when measuring dispersion using small and large quadrats, and when using A> or $>> quadrats. 1efer to your tables and briefly describe the patterns of association determined and which measures are significant. 6our discussion should be relatively brief. !ddress the following questions; ") 3hich measures of 4T are more li#ely to be accurate + those obtained using $>> small quadrats or those obtained using $>> large quadratsG 3hyG $) 3hich measures of 4T are more li#ely to be accurate + those obtained using A> quadrats or those obtained using $>> quadratsG 3hyG %) 5ow does quadrat size affect the values obtained for dispersionG 3hy do you suppose this affect occursG C) 5ow does the number of quadrats used affect the values obtained for dispersionG 3hich sample size is more li#ely to be most accurateG 3ould using a large number of quadrats &ustify using a smaller size of quadratG A) 5ow does the abundance of a species (i.e. common versus rare) affect the reliability of the measurementsG ?) 'ased on your confidence in these measures what do you thin# are the real population sizes and dispersions of each species and what ecological factors do you thin# cause themG @) Is there any evidence that any of the species sampled are allelopathicG 3hat ecological factors might e)plain the patterns of association measuredG

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