Plant Ecology 170: 167–184, 2004.

167
© 2004 Kluwer Academic Publishers. Printed in the Netherlands.

Population dynamics of Mammillaria magnimamma Haworth. (Cactaceae)

in a lava-field in central Mexico

Teresa Valverde*, Sandra Quijas, Manuela López-Villavicencio and Silvia Castillo

Departamento de Ecología y Recursos Naturales, Facultad de Ciencias, Universidad Nacional Autónoma de
México. Ciudad Universitaria, México, 04510, D.F., México; *Author for correspondence (e-mail:
mtvv@hp.fciencias.unam.mx)
Received 23 October 2001; accepted in revised form 23 August 2002

Key words: Cacti, Demography, Elasticity analysis, Long-lived species, Population projection matrices, Stochas-
tic simulations

Abstract

One of the habitats occupied by Mammillaria magnimamma is a 2000-year old lava-field, in Mexico City. The
great ecological interest on this lava-field and the little knowledge there is regarding cacti population ecology
have compelled us to analyse the demography of this species to evaluate its present conservation status at this
site. We studied two populations of this species within the lava-field: one in a disturbed site (i.e., recently burned)
and another one in a well preserved site. For each population we built two size-based population projection ma-
trices (1996/97 and 1997/98). Demographic data were gathered directly from observations of plant fates from
one year to the next. Additionally, seed germination and seedling establishment experiments were carried out in
the field to estimate fecundity values and seedling survival probabilities. The four matrices built were used to
perform numerical analyses simulating yearly stochastic demographic variation to project the overall popula-
tion’s long-term behaviour under these changing conditions. Three of the four matrices showed ␭ values slightly
below unity. In these cases elasticity values were highest for matrix entries corresponding to plants remaining in
their same category. The matrix that showed a ␭ value above unity (well preserved site, 1997/98) had higher
elasticity values for entries referring to seedling survival and growth. The numerical simulations of demographic
stochasticity showed that the population appears to be growing at a slow rate. According to the simulation re-
sults, the variation in overall population size over time may be accounted for by yearly variation in seed germi-
nation and seedling survival. Population persistence probability might decrease significantly if fire frequency
increases.

Introduction rather than the exception (Bierzychudek 1982; Cipol-

The analysis of demographic patterns in plants dur-
ing the last three decades has produced a large
amount of information about important aspects of the
biology and life history of a number of species (Sil-
vertown et al. 1993). This information has allowed us
to deepen our understanding of the relevance of dif-
ferent population processes, particularly among an-
nual or generally short-lived herbs and shrubs from
temperate habitats. One aspect that is now generally
accepted is that spatio-temporal variation in the de-
mographic behaviour of plant populations is the rule
lini et al. 1994; Horvitz and Schemske 1995; Valverde
and Silvertown 1998). Yet, the way in which this de-
mographic variation occurs, the life cycle phases it
affects, and its long-term population consequences are
poorly understood, particularly among long-lived
plant species. The understanding of the factors that
determine long-term population dynamics still re-
mains a central issue in ecology (Horvitz and Schem-
ske 1995).
Cacti are a plant group that has received little at-
tention from a demographic point of view. Recent de-
mographic information on cacti species is adding sig-
168
nificantly to our understanding of plant population
dynamics in nature, since they include long-lived spe-
cies that inhabit dry tropical areas, two particularities
that are poorly represented within the demographic
literature. The available demographic information on
this plant group, as well as on other long-lived plant
species, suggest that spatio-temporal fluctuations in
the success of early life-cycle stages are responsible
for long-term population fluctuations (Bowers 1997;
Pierson and Turner 1998). However, projection ma-
trix results and their corresponding elasticity analyses
have shown that, in general, matrix entries corre-
sponding to seed and seedling transitions seem to
contribute little to population growth rate (Godínez-
Alvarez et al. 1999; Esparza-Olguín et al. 2002). Al-
though long-term numerical fluctuations appear to be
common within these species, no information is avail-
able as to what kind of ecological processes may ac-
count for them. In this study we analyse the popula-
tion dynamics of the cactus Mammillaria
magnimamma Haworth. in a lava-field in Mexico
City. We incorporated information on the demo-
graphic behaviour of this species in different sites and
years into a long-term stochastic numerical simula-
tion which allowed us to project the potential fate of
the population. The aim of this exercise was to con-
tribute to the understanding of the way in which spa-
tio-temporal changes in the demography of plant spe-
cies give rise to fluctuating population numbers.
An additional interest to address the population
dynamics of Mammillaria magnimamma is related to
the ecosystem in which it was studied: a 2000 year
old lava-field with a particularly high plant diversity
(Carrillo 1995). This lava-field was created after the
eruption of ⬙El Xitle⬙ volcano and comprises an im-
portant part of what is now Mexico City, one of the
most densely populated areas in the world. Unfortu-
nately, the lava-field is slowly being turned into ur-
ban areas and much of its biological value is being
lost. In fact, some of its endemic species (e.g., Mam-
millaria sanangelensis, Bletia urbana) are virtually
extinct. In recent years a small fraction of this lava-
field was protected in the form of a reserve, in which
this study was carried out. It has been suggested that
the analysis of the persistence probability of species
that are particularly vulnerable to disturbances within
an ecosystem may throw light on the conservation
status of the ecosystem itself by providing informa-
tion on the probability of native species turnover
(Menges 1990). Since it has been suggested that cacti
are particularly vulnerable to disturbances (Hernán-
dez and Godínez 1994), the demographic analysis of
M. magnimamma is expected to offer empirical tools
that could aid in the evaluation of the conservation
status of the reserve where the study was performed.
The high demographic vulnerability of cacti is pre-
sumed to be a result of their slow individual growth
rates and frequently high mortality rates during the
juvenile phases; apparently these features contribute
to limit their capacity for population growth (Hernán-
dez and Godínez 1994; Contreras and Valverde 2002;
Esparza-Olguín et al. 2002). Additionally, many cacti
species are highly restricted in their distribution and
occupy very specific habitats, which makes them
prone to extinction by habitat destruction. In fact, the
Cactaceae is a plant family with a particularly high
number of threatened species (Hunt 1992; Anderson
et al. 1994). However, our knowledge of the popula-
tion behaviour of most of these species is very lim-
ited.
In this paper we address the demographic analysis
of two populations of Mammillaria magnimamma,
one in a well-preserved and one in a disturbed (i.e.,
recently burnt) site within the ⬙El Xitle⬙ lava-field,
through the construction of population projection ma-
trices for two growth periods (1996/97 and 1997/98).
With the information obtained from the demographic
analysis we carried out numerical simulations, using
the stochastic approach developed by Bierzychudek
(1982), incorporating the effect of the spatio-tempo-
ral variation in the demography of the species in or-
der to understand the way in which this variation may
result in population fluctuations, as well as to evalu-
ate the long-term persistence probability of this spe-
cies in the area. The stochastic analysis was used to
simulate different disturbance regimes and evaluate
population persistence probability under different
probability of occurrence of fires, which are the main
cause of disturbance at the site.
Methods
The study site
This study was carried out at the ⬙El Xitle⬙ (meaning
⬙navel⬙ in Nahuatl) lava-field, located within Mexico
City (19°40⬘ N, 99°00⬘ W) (Figure 1). This lava-field
is the result of the eruption of the ⬙El Xitle⬙ volcano,
which took place over two thousand years ago (Car-
rillo 1995). The original area occupied by this lava-
field was 80 km 2, covering an altitudinal gradient

Figure 1. Location of the study site (⬙El Xitle⬙ lava-field) within Mexico City.
from 3,100 to 2,250 m above sea level. The lower part
of the lava-field (2,500–2,250 m a.s.l.) was covered
by a plant community (described in detail by Rze-
dowski (1954)) dominated by the shrub Senecio prae-
cox. This shrubland, in which 350 plant species could
be found (Rzedowski 1954), comprised 40% of the
original area covered by the lava-field. Today the area
still occupied by this type of shrubland has been re-
duced to only 2.9 km 2, most of which is now pro-
tected by the ⬙Pedregal de San Angel⬙ Reserve, be-
longing to the National Autonomous University of
Mexico (UNAM). Within this reserve, only 226 of the
original plant species were found in 1990, plus 77
new records, most of which correspond to ruderal
species that have recently entered the ecosystem (Va-
liente-Banuet and de Luna 1990).
The M. magnimamma populations studied were lo-
cated within the Pedregal Reserve referred to above.
This area has a temperate sub-humid climate with a
seasonal rainfall pattern. Annual rainfall ranges be-
tween 700 and 950 mm and most of it falls between
June and September, while the rest of the year is
rather dry. The total yearly precipitation for 1997 and
1998 was 749.4 mm and 907.2 mm, respectively (data
from the Observatorio Meteorológico, UNAM).
Mean annual temperature is 16.3 °C, with an annual
minimum of −6 °C and an annual maximum of 34 °C.
Despite the relatively high annual precipitation, the
reserve shows xerophytic characteristics as a result of
the basaltic substrate, which has a low water reten-
tion capacity. The soils are shallow, produced by the
erosion of the basaltic rock and the decomposition of
organic matter. The vegetation of this area is domi-
169
nated by the shrubs Senecio praecox, Verbesina vir-
gata, Dodonea viscosa and Wigandia urens, the tree
Buddleia cordata, and the herbs Muhlenbergia ro-
busta, Dahlia coccinea and Echeverria gibbiflora,
among others (Rzedowski 1954).
Within the ⬙Pedregal de San Angel⬙ reserve we
chose two sites: a well-preserved (P) and a relatively
more disturbed (D) site. The main difference between
these two sites is the apparent degree of conservation.
As noted above, the most prevalent disturbance fac-
tor in this area is fire, generally of human origin, oc-
curring mainly in spring, towards the end of the pro-
longed dry season. Between 1996 and 1998 Site D
was burnt twice, while Site P was damaged only
slightly by the 1998 fire. Additionally, Site D is lo-
cated closer to the edge of the reserve, which makes
it more prone to colonisation by ruderals. As a con-
sequence, the general features of these two sites are
quite different, with P site showing a relatively more
developed tree cover, deeper soils, higher soil water
content, and lower solar radiation at ground level
compared to site D (Valverde et al. 1999). These dif-
fering conditions in turn determine a relatively differ-
ent species composition between the two sites: site P
shows a sparse tree cover dominated by Buddleia cor-
data, while site D is colonised mainly by grasses and
herbs.
The species
Mammillaria magnimamma is a small globular cac-
tus with a green-greyish coloration and with one to
several cylindric stems. The flowers (20–25 mm long)
170
are pinkish to purpleish. Fruits are clavate, 20–35 mm
long, and with a deep red colour. Each fruit contains
between 20 and 100 round, brown seeds, 0.7 mm in
diameter. M. magnimamma is endemic from Mexico;
it may be found in the central highlands at altitudes
from 1,700 to 2,600 m above sea level, in grasslands
on flat and rocky soils (Bravo-Hollis and Sánchez-
Mejorada 1991).
Field methods
Within each of the study sites (P and D) we marked
circular plots of 6 m in diameter (28.3 m 2) within
which M. magnimamma plants were located through
coordinates. In June–July 1996 each plant was tagged
with plastic flagging and given an identification num-
ber. The sample included 203 plants within 22 plots
in site D, and 206 within 25 plots in site P. We re-
corded plant size by measuring the diameter of all the
stems of each plant in summer 1996; we re-measured
all sampled plants in summer 1997 and later in sum-
mer 1998. Plant size was evaluated as ’cumulative
diameter’, which was the sum of the diameters of all
stems per plant. Additionally, we visited the sampled
plants every two weeks during the flowering and
fruiting periods (February to August 1997 and 1998)
and took records of flower and fruit production per
plant. A number of fruits was collected and the aver-
age number of seeds per fruit was calculated. Using
the data on the demographic fate of each individual
plant from one year to the next we built population
projection matrices for the periods 1996/97 and
1997/98 for each of the study sites.
As no germinating seeds or seedlings were ob-
served in the field, the matrix entries concerning these
life-cycle stages were calculated from the results of
field experiments. Seed germination experiments
were performed in order to estimate the probability
of seeds germinating immediately after dispersal.
These probabilities were later incorporated in the ma-
trix analysis as part of the fecundity values, as de-
tailed below. In June 1997 (just before the start of the
rainy season) 100 newly collected seeds were scat-
tered within each of four 15 × 15 cm plots marked on
the ground with wooden sticks in each study site (D
and P). These seeds were monitored every two weeks
until August in order to evaluate seed germination.
Average seed germination percentage (arcsine trans-
formed) was compared between sites using a student-t
test. In 1998 a more detailed experimental design was
followed. We built 15 × 15 cm plastic mesh open
boxes, 2 cm deep, which were slightly buried in the
ground in particular microsites within each study site.
In each box we scattered 50 newly collected seeds.
Boxes were designed to prevent seeds from being
transported by wind or water. Within each site we set
a total of eight boxes in the following type of micro-
sites: two in fully illuminated microsites with soil,
two in shaded microsites with soil, two in fully illu-
minated microsites with no soil and two in shaded
microsites with no soil (see below). These experiment
started at the beginning of August 1998 and seed ger-
mination was recorded daily for the first four weeks,
and then every week for two months. Total germina-
tion percentage per box was averaged for each site;
statistical differences in mean germination percentage
between sites were evaluated through a student-t test.
The effect of sowing treatments within sites was eval-
uated only superficially, since the number of repli-
cates per treatment did not allow further statistical
treatment.
Seedling survival experiments were carried out in
the field in order to estimate the probability of seed-
ling survival and growth from one year to the next.
These estimates were later used in the construction of
projection matrices, as detailed below. In June 1997
seeds were germinated in greenhouse conditions in
order to obtain seedlings. These seedlings were kept
in the greenhouse for two months and were later
planted in the study sites. In August 1997, three 40 ×
40 cm quadrats were marked on the ground with
wooden sticks in each study site and 40 seedlings
were planted evenly spaced (to allow relocation)
within them. The size and degree of development of
these seedlings was similar to that of two- or three-
week old seedlings observed in the field as part of the
seed germination experiments described above. The
survival of the introduced seedlings was recorded ev-
ery two weeks for two months. In 1998 a slightly dif-
ferent experimental design was followed also for this
part of the study. In June 1998 seeds were germinated
in greenhouse conditions in order to obtain seedlings.
When seedlings were two months old, they were
planted in the field in 20 × 20 cm quadrats marked on
the ground with wooden sticks; 30 seedlings were
planted evenly spaced within each of these quadrats.
A total of eight seedling quadrats were set in each
study site (D and P). The type of microsites chosen
to plant these seedling was the same as in the 1998
germination experiment described above (i.e., a fac-
torial design of fully illuminated and shaded micro-
sites, with and without soil; two replicates per treat-

ment). Seedling survival was followed daily for a
month, and every two weeks for another two months.
For the 1998 data, survival curves were obtained for
each treatment and each site; the effect of treatments
within each site was evaluated using the Peto and
Peto survival curve pair-wise comparisons (Pyke and
Thompson 1986). The four treatments used in the
seed germination and seedling establishment experi-
ments of 1998 all represent common microsites
within the study sites. Spatial heterogeneity in envi-
ronmental conditions is tremendous within lava-
fields. Thus, the idea of performing these experiments
in a variety of microsites was to obtain a more real-
istic picture of actual overall germination and estab-
lishment probabilities within the reserve.
Numerical methods
The field data on plant survival and growth were used
to construct population projection matrices for each
of the study sites (P and D) and for the two periods
studied (1996/97 and 1997/98). These four matrices
will be called P-1996/97, P-1997/98, D-1996/97 and
D-1997/98. Lefkovitch matrices were used, character-
ised by the classification of individuals according to
size classes. The general matrix model is given by
n t+1 = A n t, in which A represents a square matrix,
and n is a column vector whose elements represent
the number of individuals in each class at time t and
t+1. Matrices are composed by a ij entries, which re-
present the probability of an individual in the j-th
class contributing or becoming an individual in the
i-th class from time t to time t+1 (Caswell 1989). In
this case the time step was one year.
The population was subdivided into seven size cat-
egories, as defined in Table 1. This number of cate-
gories allowed a detailed dissection of the life cycle;
at the same time, we made sure that sample size in
each category was large enough to allow robust esti-
mates of transition probabilities (see n in Table 4). We
preferred this option rather than using the algorithms
that have been developed for choosing the number of
categories (Moloney 1986), since we aimed to build
matrices that were comparable (i.e., with the same
number of categories, defined by the same size inter-
vals). The probability that a plant in each size-class
would contribute to a different or the same size-class
from one year to the next was calculated directly from
field observations, i.e. from the proportion of plants
in each size-class that followed different fates be-
tween t and t+1. The fecundity elements were given
171
Table 1. Size categories chosen to model the size-based demogra-
phy of M. magnimamma.
Category Cumulative diameter (cm)
1 < 0.3 (seedlings)
2 0.3–4.5
3 4.6–7.5
4 7.6–10.5
5 10.6–13.5
6 13.6–20.5
7 > 20.6
by the number of seeds produced in a year by an av-
erage individual of each size class, multiplied by the
probability of seed germination. Thus, fecundity was
given in terms of the average number of seedlings
produced per plant in each size class. This implies
that seeds do not remain viable in the soil for long
periods of time; seed viability is high in newly col-
lected seeds (85–90%, Ruedas et al. (2000)), there-
fore it is reasonable to expect that seeds either germi-
nate or die within three or four months after seed
shed. With regard to germination probability, we cal-
culated it from the data of the germination experi-
ments performed in the field in both the disturbed and
the well-preserved sites. As the 1998 germination ex-
periments were carried out in four different microsites
(illuminated/shaded, and with/without soil) in both
the preserved and the disturbed sites, to obtain the
germination probability for each site we averaged the
results of the four microsites, assuming that their fre-
quency in the environment is approximately the same.
The transition from class 1 to class 2 (i.e., seed-
ling survival and growth) was calculated according to
the results of the seedling survival experiments per-
formed in the field each year. In 1997 all planted
seedlings died within the first five weeks in both the
preserved and the disturbed sites; thus, to estimate
their survival probability after a year, we built the
survival curves for these seedlings with the mortality
data obtained over the first five weeks and extrapo-
lated them to 365 days. In 1998 seedlings were fol-
lowed for three months, until survival curves ap-
peared to reach a plateau. In this case we also used
the survivorship data to fit a survivorship curve from
which the probability of survival to day 365 was cal-
culated. This probability was incorporated in the ma-
trix model as the probability of seedling survival and
growth to category 2.
172
The probability of plants in the largest size cate-
gory (7) remaining in the same category from one
year to the next was calculated directly from field
observation in the cases of matrices P-1996/97 and
P-1997/98. However, in the disturbed site no mortal-
ity was observed in this category; yet, some mortality
of large adults must be incorporated in the matrix,
otherwise it would imply that individuals are immor-
tal. Thus, the mortality of large adults for matrices
from site D was calculated as follows: 1) first we ran
the matrices of the well preserved site (P) in the pro-
gram STAGECOACH (Cochran and Ellner 1992),
from which a maximum longevity estimate was ob-
tained; 2) then we adjusted the value of the matrix
entry referring to the survival of class-7 individuals
in the matrices of the disturbed site (D) in order to
equal the longevity estimate obtained for matrices in
site P.
The dominant eigen-value (␭) and the correspon-
dent right (w) and left (v) eigen-vectors of the matrix
were obtained by the power method. The confidence
intervals for ␭ were obtained through the analytical
method described in Caswell (1989, p. 185). Addi-
tionally, elasticity matrices were calculated from the
right and left eigen-vectors as follows:
e ij ⫽ 共v iw j/具v, w典兲共a ij/␭兲. (1)
The elasticity (e ij) of each matrix entry, a ij, repre-
sents a measure of proportional sensitivity of ␭ to
proportional changes in a ij. Also, since the sum of all
the e ij in an elasticity matrix equals unity, each e ij
may be interpreted as the contribution of its respec-
tive a ij to the value of ␭ (de Kroon et al. 1986).
The deterministic time-invariant matrix model de-
scribed above is based on the assumption that demo-
graphic conditions do not change over time and that
the modelled population grows exponentially at a rate
given by ␭. This matrix model is therefore used to
project the future consequences of the present demo-
graphic scenario, should everything remain the same.
However, the demography of most plant species vary
over time and space (Horvitz and Schemske 1995;
Quintana-Ascencio and Menges 1996; Valverde and
Silvertown 1998; Mandujano et al. 2001). Therefore,
to evaluate the potential long-term demographic be-
haviour of a species it is more realistic to incorporate
demographic information gathered in different sites
and different years. In this study we used the four
projection matrices built (P-1996/97, P-1997/98,
D-1996/97 and D-1997/98) to perform numerical
simulations of the behaviour of the M. magnimamma
population as a whole under the spatio-temporally
heterogeneous environment characteristic of the study
site. The four resulting matrices represent demo-
graphic variation in space and time. Yet, in this case
these four matrices were used to project numerical
changes in time, because the variation that occurs in
space (i.e., the differences between the demographic
behaviour between D and P sites) may also occur
through time when disturbances take place in differ-
ent patches of the ecosystem.
In this section we used the method originally de-
veloped by Bierzychudek (1982), which incorporates
stochasticity in the demographic behaviour of the
studied species by using the information of more than
one matrix. An initial population vector was built by
averaging the observed population structures for each
site and year, so that the initial size structure in the
simulations was similar to the actual size structure
observed in the field. This vector was used to start the
matrix iterations. For each matrix iteration a ran-
domly selected matrix was used; the selection criteria
varied in each of the five simulation exercises (from
now on called ⬙runs⬙) performed. a) In the first run
each of the four matrix was sampled with the same
probability (0.25). b) In the second run we incorpo-
rated the relative frequency of ⬙good⬙ and ⬙bad⬙ years
according to rainfall data; we analysed the rainfall
pattern in the study site for the last 40 years (data
from the Observatorio Meteorológico, UNAM) and
calculated the probability of occurrence of ⬙good⬙
years (those with a total yearly precipitation above
900 mm, which represents the 1997–1998 period),
and ⬙bad⬙ years (with a yearly precipitation below 900
mm, represented by the 1996–97 period). These prob-
abilities were 0.308 and 0.692, respectively. Addition-
ally, we incorporated the probability of disturbances
(fires) taking place in two out of three years, on av-
erage. The resulting probabilities for each matrix are
reported in Table 6. c) In the third run we used the
same ⬙good⬙ and ⬙bad⬙ year frequencies as above, but
fire occurrence probability was one every other year,
which is close to the actual fire frequency observed
in the last 10 years at the site. d) The fourth run con-
sidered ⬙good⬙ and ⬙bad⬙ years as above, but fire fre-
quency was, on average, of one every three years. d)
Finally, in the fifth run we used the same conditions
as above, but with an average fire frequency of one
every five years (Table 6).
In each run, a total of 100 matrix iterations were
carried out, after which the numerical behaviour of

the population (and of each of the size categories)
could be plotted against time. To calculate the popu-
lation growth rate under these stochastic conditions
(␭ s), we carried out 30 series of 100 matrix iterations.
For each series of 100 iterations, we plotted the natu-
ral logarithm of average population size against time
(iterations) to represent the numerical long-term pop-
ulation behaviour expected under stochastic demo-
graphic variation and we fitted a linear regression to
the data. The slope of the fitted line was considered a
measure of the intrinsic rate of population increase
(r), from which ␭ s could be calculated (␭ = e r)
(Bierzychudek 1982). Since we calculated 30 ␭ val-
ues in each case, we were able to report an average ␭
value for each run (plus standard deviation) as well
as to estimate the ‘persistence probability’ of the over-
all population according to its projected long term
numerical behaviour over a period of 100 years. The
latter were defined as the relative frequency with
which ␭ values below unity were obtained.
Results
Early stages of plant development
In 1997 the number of seeds that were observed ger-
minating in the field experiment carried out was very
low (Table 2). Four out of 400 seeds germinated in
site P, while only one out of 400 germinated in site
D. This difference was not statistically significant (t =
1.0, d.f. = 3, p = 0.39). In 1998 mean germination
percentage across the four sowing treatments was
15.8% in site P and 5.8% in site D (Table 2); the dif-
ference between these values was statistically signif-
icant (t = 2.64, d.f. = 7, p = 0.03). The higher germi-
nation percentage obtained this year compared to the
one observed in 1997 determined much higher fecun-
dity values for the 1997–98 period (see results be-
low). The more detailed experimental design fol-
lowed in this second experiment also allowed us to
evaluate, at least in a general way, the microenviron-
mental conditions that are more appropriate for the
germination of M. magnimamma seeds. In both sites
the shaded microsites with no soil resulted in higher
germination percentages, while the fully illuminated
site with no soil did not show any germination at all
(Table 2). The low number of replicates used for each
treatment did not allow further statistical analysis of
these results.
173
Table 2. Mean germination percentages (± standard deviation) ob-
tained in the field experiments performed in the Preserved and the
Disturbed sites for the two study periods. For the 1998 results ini-
tials refer to sowing treatments: IWS = illuminated, with soil; INS
= illuminated, no soil; SWS = shaded, with soil; SNS = shaded, no
soil. In each of the 1998 sowing treatments (1998) the averages
were obtained from only two replicates per site. In 1997 averages
are from four replicates per site.
% Germination
Well-Preserved site Disturbed site
1997 1.00 ± 2.00 0.25 ± 0.50
Germination Probability 0.01 0.0025
1998
IWS 22.00 ± 14.14 0
INS 0 0
SWS 5.00 ± 4.24 6.00 ± 5.65
SNS 36.00 ± 25.45 17.00 ± 9.89
Germination probability 0.157 0.058
Seedling survival in 1997 is described in Figure
2a. All seedlings died within the first 45 days after
planting. Seedlings in site P showed a higher initial
survival compared to those in site D, but the final re-
sult was the same for both sites. According to the ex-
trapolation of the 1997 survival curves to day 365, the
probability of seedling survival was 0.001 in site D,
and 0.003 in site P. In 1998 higher seedling survival
rates were observed than in the previous year, and
seedling survival was in general higher in site P than
in site D (Figure 2, 2b and 2c). In site P seedlings
planted in shaded microsites with soil had a signifi-
cantly higher survival rate than the other three treat-
ments (Figure 2b), while in site D the most successful
treatment were the fully illuminated microsites with
soil, which showed a significantly higher survival rate
than the other treatments. In this site, the fully illu-
minated microsites with no soil resulted in total seed-
ling mortality within the first nine days after planting.
Combining the results of all treatments within each
site, the survival probability to day 365 was 0.0015
for site D, and 0.134 for site P. These data were in-
terpreted as the transition probability from class 1 to
class 2 in the Lefkovitch matrices (Table 4).
Reproduction
M. magnimamma showed a relatively long yearly re-
productive season (Figure 3). Flower buds started in
February and ended in June–July in both years and
sites. Flowers were found between March and June.
174

Figure 2. Seedling survivorship curves for a) the 1996/97 growth period (both study sites), b) the 1997/98 period in the well preserved site
and c) the 1997/98 period in the disturbed site. The different lines in b) and c) correspond to planting treatments; different letters above
survivorship curves imply significant differences between treatments according to Peto and Peto pair-wise comparisons.

Once flowers are pollinated, fruits take a long time to
develop. In the two years of study the fruiting period
lasted for around five months, from April to August,
with maximum fruit production occurring between
June and July (Figure 3). In 1997 fruit production was
slightly higher than in 1998. Yet, the reproductive
phenology of the studied populations was remarkably
similar both between sites and between years.
The total number of fruits produced per plant were
counted each year and multiplied by the average num-
ber of seeds per fruit (for 1996/97: 93.5 ± 36.8 for
both sites, n = 25; and for 1997/98: 93.5 ± 42.0 for
site P, n = 69; and 75.4 ± 45.0 for site D, n = 49).
This gave the estimated number of seeds produced
per plant, which was in turn multiplied by the seed
germination probability (see Table 2) to estimate fe-

Figure 3. Reproductive phenology of M. magnimamma plants for two years at the ⬙El Xitle⬙ lava-field. a) Well preserved site and b) dis-
turbed site.
cundity. Thus, here fecundity is given in terms of the
number of seedlings produced per plant in a repro-
ductive season. Average fecundity for each size cat-
egory is reported in Table 3; these results show that,
in general, fecundity was larger as plant size in-
creased. In 1996/97 fecundities were lower than in
1997/98 (mainly due to the higher germination per-
centages obtained in 1998), and plants in site P
showed consistently higher fecundity values than
plants in site D (Table 3). These fecundities were in-
corporated in the transition matrices as the contribu-
tion of each size category to the seedling category
from one year to the next (i.e., first row of the ma-
trix).
175
Size-based demography
We built four population projection matrices, two for
each site (1996/97 and 1997/98) (Table 4). Most cat-
egories showed positive transitions towards several
potential fates in all matrices, i.e. growing one, two,
or even three categories, decreasing in size one to
several categories, or staying in the same category.
Most matrices showed the highest values in those en-
tries referring to the persistence of plants in the same
category. The survival of plants in the 7 th category
(which was estimated for matrices from site D – see
Methods) was similar in the four matrices. Plant mor-
tality was highest in the first category and decreased
with plant size; in general, both fecundity and mor-
tality were higher in the second compared to the first
growth period for both sites. Both ␭ values obtained
176

Table 3. Fecundity values (mean number of seedlings produced per plant) for each size category in a) the preserved and b) the disturbed sites
for each of the study periods.

Size Category 1996–97 No. fruits/plant No. Seeds/plant Fecundity 1997–98 No. fruits/plant No. Seeds/plant Fecundity

a) Preserved Site
1 0 0 0 0 0
2 0 0 0 0.150 14.025 2.22
3 1.057 98.301 0.98 0.941 87.983 13.90
4 3.380 314.34 3.14 1.313 122.766 19.40
5 4.300 399.9 4.00 1.471 137.539 21.73
6 1.632 151.776 1.51 1.731 161.849 25.57
7 6.263 582.459 5.82 2.808 262.548 41.48
b) Disturbed Site
1 0 0 0 0 0 0
2 0.306 28.458 0.07 0 0 0
3 1.627 151.311 0.38 0.396 29.858 1.73
4 2.364 219.852 0.55 1.018 76.757 4.45
5 1.462 135.966 0.34 0.810 61.074 3.54
6 2.692 250.356 0.63 1.817 137.001 8.18
7 4.857 451.701 1.13 1.545 116.493 6.76

in site D were below unity (although not significantly
so); the same occurred for P-1996/97. However, for
P-1997/98 we obtained a population growth rate sig-
nificantly higher than unity. This was mainly a result
of the high fecundity values (resulting from high ger-
mination percentages) incorporated in this matrix, as
well as a high seedling survival probability for this
year in this site.
For the three matrices with ␭ values below unity
the expected population structure at equilibrium (w)
has a high proportion of individuals in the first cat-
egory (seedlings), followed by those in the last cat-
egory (Table 4). However, for the matrix with a pos-
itive ␭ value (P-1997/98) the vector of the stable size-
category distribution describes a population
represented mainly by seedlings, juveniles and young
adults. Expected size-specific reproductive values at
equilibrium (v) show a very small contribution of
seedlings, particularly in those matrices in which
seedling mortality is higher; the reproductive value of
other categories is relatively uniform, with the excep-
tion of matrix P-1997/98 in which there is a clear
tendency towards an increase in reproductive value
with increasing size.
The elasticity matrices (Table 5) show two clearly
distinct patterns: for the three matrices referring to
populations with ␭ values below unity (P-1996/97,
D-1996/97 and D-1997/98) the highest elasticity val-
ues are those corresponding to the persistence of large
adults (7 th category) in their same category, while
elasticities for fecundity and growth entries are very
low. On the other hand, for the matrix with a positive
population growth rate (P-1997/98) the highest elas-
ticity value refers to seedling survival and growth,
also showing relatively high values for the entries
corresponding to fecundity and growth (Table 5).
Numerical simulations of demographic stochasticity
When matrix iterations were performed choosing a
different (randomly allocated) matrix for each itera-
tion event, each series of 100 iterations resulted in a
different projection of numerical population behav-
iour, depending on the particular order and frequency
with which different matrices were utilised. In Fig-
ure 4 only three of such series are represented, plot-
ting the numerical behaviour of a growing and a de-
clining population (Figure 4a and 4b, respectively), as
well as the numerical behaviour of each size category
independently for a growing population (Figure 4c).
In these plots we used a logarithmic scale on the y
axis to allow visual appreciation of the overall popu-
lation trend and to fit a linear regression from which
the slope (r = intrinsic rate of population increase)
could be calculated. Since r = ln ␭, then overall pop-
ulation growth rate was calculated as ␭ s = e r. In all
series performed the numerical fluctuations in overall
population size were given mainly by fluctuations in
seedling numbers (see Figure 4c). High peaks in the
seedling category were generally followed by peaks
 177

Table 4. Population transition matrices for sites P and D and for periods 1996/97 and 1997/98. The ␭ values (± 95% confidence intervals) are
given above each matrix. n x = number of individuals per size-category from which transitions were calculated; q x = size-specific mortality;
w = stable size-category distribution; v = size-specific reproductive values. Entries in the main diagonal are underlined to facilitate reading.

1 2 3 4 5 6 7 (w) (v)

Site P 1996/97
␭ = 0.956 ± 0.086
1 0 0 0.983 3.143 4 1.518 5.825 0.849 6.3E-05
2 0.003 0.613 0.019 0 0 0 0 0.001 0.201
3 0 0.355 0.509 0.064 0 0 0 0.002 0.195
4 0 0 0.415 0.762 0.200 0.053 0 0.009 0.180
5 0 0 0.038 0.095 0.350 0 0 0.002 0.167
6 0 0 0.019 0.032 0.400 0.737 0 0.006 0.149
7 0 0 0 0.016 0.050 0.211 0.947 0.131 0.108
nx 120 32 54 64 20 18 18
qx 0.997 0.032 0 0.0318 0 0.0001 0.0526
Site P 1997/98
␭ = 1.333 ± 0.174
1 0 2.216 13.901 19.397 21.731 25.572 41.483 0.819 0.003
2 0.134 0.450 0.073 0 0.059 0 0 0.128 0.034
3 0 0.200 0.667 0.156 0 0.039 0 0.041 0.111
4 0 0 0.177 0.590 0.235 0.039 0 0.010 0.137
5 0 0 0 0.108 0.471 0.115 0 0.001 0.152
6 0 0 0 0.012 0.118 0.692 0.039 0.000 0.199
7 0 0 0 0 0 0.039 0.923 4.1E-05 0.362
nx 240 41 51 83 17 26 26
qx 0.866 0.3500 0.0784 0.1326 0.1177 0.0768 0.0385
Site D 1996–97
␭ = 0.967 ± 0.095
1 0 0.070 0.380 0.550 0.340 0.630 1.130 0.539 1.7E-05
2 0.001 0.270 0.010 0 0 0 0 7.8E-05 0.164
3 0 0.550 0.290 0.030 0 0 0 6.7E-05 0.177
4 0 0.100 0.560 0.240 0 0 0 6.3E-05 0.169
5 0 0 0.090 0.360 0.150 0 0 3.6E-05 0.172
6 0 0 0.040 0.270 0.850 0.540 0 1.2E-04 0.165
7 0 0 0 0.060 0 0.460 0.967 0.461 0.153
nx 120 50 76 34 14 14 15
qx 0.999 0.0800 0.0100 0.0400 0 0 0.0333
Site D 1997/98
␭ = 0.945 ± 0.099
1 0 0 1.732 4.452 3.542 8.182 6.757 0.832 2.0E-04
2 0.002 0.579 0.038 0.036 0 0 0 0.010 0.129
3 0 0.211 0.623 0.250 0.048 0 0 0.035 0.148
4 0 0.105 0.226 0.554 0.143 0.065 0 0.033 0.155
5 0 0 0.019 0.089 0.571 0.162 0 0.018 0.205
6 0 0 0.019 0 0.238 0.548 0.046 0.019 0.195
7 0 0 0 0 0 0.194 0.881 0.055 0.168
nx 240 19 53 56 21 31 22
qx 0.9985 0.1053 0.0755 0.0714 0 0.0323 0.0735

in category 2 and so on; larger size categories were The results of the stochastic simulations were a
numerically relatively stable. function of the particular probability of occurrence
178

Table 5. Elasticities matrices for sites P and D and for periods 1996/97 and 1997/98. Entries in the main diagonal are underlined to facilitate
reading. The three highest elasticity values in each matrix are typed in bold letters.

1 2 3 4 5 6 7

Site P 1996/97
1 0 0 7.8E-06 1.1E-04 2.5E-04 3.1E-05 0.003
2 0.003 0.006 4.8E-04 0 0 0 0
3 0 0.004 0.013 0.007 0 0 0
4 0 0 0.009 0.076 0.004 0.003 0
5 0 0 8.0E-04 0.009 0.006 0 0
6 0 0 3.6E-04 0.003 0.006 0.036 0
7 0 0 0 9.5E-04 5.4E-04 0.007 0.797
Site P 1997/98
1 0 0.054 0.108 0.038 0.005 0.002 3.3E-04
2 0.208 0.109 0.006 0 1.5E-04 0 0
3 0 0.160 0.170 0.010 0 1.1E-04 0
4 0 0 0.056 0.046 0.002 1.3E-04 0
5 0 0 0 0.009 0.005 4.3E-04 0
6 0 0 0 0.001 0.002 0.003 1.8E-05
7 0 0 0 0 0 3.4E-04 7.8E-04
Site D 1996/97
1 0 1.4E-09 6.2E-09 8.5E-09 3.0E-09 1.8E-08 1.3E-04
2 1.3E-04 4.9E-05 2.1E-06 0 0 0 0
3 0 1.1E-04 5.1E-05 4.9E-06 0 0 0
4 0 1.9E-05 9.2E-05 3.7E-05 0 0 0
5 0 0 1.6E-05 5.7E-05 1.4E-05 0 0
6 0 0 6.4E-06 4.1E-05 7.3E-05 1.5E-04 0
7 0 0 0 8.5E-06 0 1.2E-04 0.999
Site D 1997/98
1 0 0 4.7E-04 0.001 4.8E-04 0.001 0.003
2 0.006 0.029 0.006 0.006 0 0 0
3 0 0.012 0.121 0.045 0.005 0 0
4 0 0.006 0.046 0.105 0.015 0.007 0
5 0 0 0.005 0.022 0.078 0.023 0
6 0 0 0.005 0 0.031 0.075 0.018
7 0 0 0 0 0 0.023 0.305

allocated to each of the four matrices used (see Ta-
ble 6). The highest ␭ value was obtained in the run in
which all four matrices were given the same proba-
bility of occurrence (Table 7). When the probability
of ⬙good⬙ and ⬙bad⬙ years was incorporated, along
with different disturbance regimes (given by different
fire frequencies), interesting patterns emerged. Low
fire probabilities (i.e., 0.2 and 0.3) resulted in ␭ val-
ues well above unity. When the probability of occur-
rence of a fire event for each particular year was 0.5
(which is close to the actual disturbance regime at the
site), the overall ␭ value was also above unity; how-
ever, from the 30 series performed with these condi-
tions, one described a crashing population with a
long-term ␭ value below unity. From this we esti-
mated the persistence probability of the overall popu-
lation as P = 1 − (1/30) = 0.97 (Table 7). Finally,
when the probability of fire events was set at 0.66,
the ␭ value was just above unity, while the persistence
probability of the population was 0.66 (i.e., 10 out of
30 series described a crashing population) (Table 7).
Discussion
From the four population matrices obtained in the
present study, three of them showed ␭ values slightly
below unity. Yet, in these cases the confidence inter-
 179

Figure 4. Examples of the numerical behaviour of the overall M. magnimamma population under stochastic demographic variation. a) A
growing population, obtained by allocating the same probability of occurrence to each of the four matrices. b) A declining population, in-
corporating the probability of occurrence of ⬙good⬙ and ⬙bad⬙ years and a relative frequency of fire events of 0.66. c) A growing population
showing the variation in numbers of each size category, for a fire frequency of 0.5. Line codes for size categories are as follows: 1 = dark
dotted; 2 = Dark dashed; 3 = Dark continuous; 4 = Dark dashed-dotted; 5 = Fine continuous; 6 = Fine dotted; 7 = Fine dashed.
 180

Table 6. Probabilities used for the occurrence of different matrices in the stochastic simulations. ⬙Good⬙ years refer to periods with high rainfall (above 900 mm – probability 0.308) and
⬙bad⬙ years to those with low rainfall (below 900 mm – probability 0.692). In addition to the probability of occurrence of ⬙good⬙ and ⬙bad⬙ years, different fire frequencies were simulated.
The larger numbers in each column are the resulting probabilities of occurrence of each matrix for each particular run.

Matrix ␭ 1 st run Equal prob. 2 nd run Good and bad years; 3 rd run Good and bad years; 4 th run Good and bad years; 5 th run Good and bad years;
fire 2 out of 3 years fire every other year fire every 3 years fire every 5 years

P-96/97 0.956 0.250 (0.692 × 0.33) = 0.230 (0.692 × 0.5) = 0.346 (0.692 × 0.66) = 0.461 (0.692 × 0.8) = 0.554
P-97/98 1.333 0.250 (0.308 × 0.33) = 0.102 (0.308 × 0.5) = 0.154 (0.308 × 0.66) = 0.205 (0.308 × 0.8) = 0.246
D-96/97 0.967 0.250 (0.692 × 0.66) = 0.457 (0.692 × 0.5) = 0.346 (0.692 × 0.33) = 0.231 (0.692 × 0.2) = 0.138
D-97/98 0.945 0.250 (0.308 × 0.66) = 0.203 (0.308 × 0.5) = 0.154 (0.308 × 0.33) = 0.103 (0.308 × 0.2) = 0.062
 181

Table 7. Population growth rate (␭) obtained in the different sto- to be determined by the dynamics of the early stages
chastic simulations. Each figure is the average of 30 series of 100 of plant development, during which a tremendous
iterations each; standard deviations are a measure of variation
within each of these 30 series in each case. See Table 6 for details
mortality occurs in this and other cacti species (Sten-
on the conditions for each run. Persistence probability refers to the nbergh and Lowe 1977; Jordan and Nobel 1981; Va-
relative frequency of ␭ values above unity in the 30 series per- liente-Banuet and Ezcurra 1991). Therefore, it is im-
formed. portant to address this particular aspect of the species’
␭ St. Dev. Persistence probability life-cycle to understand one of the most important
determinants of population dynamics (Godínez-Alva-
1 st run 1.086 0.020 1.00 rez et al. 1999). It is noticeable, for instance, that the
2 nd run 1.008 0.018 0.66 seeds of many cacti species germinate readily under
3 rd run 1.036 0.019 0.97 controlled conditions. M. magnimamma seeds reach
4 th run 1.063 0.019 1.00
85% germination in ca. 20 days (Ruedas et al. 2000)
5 th run 1.075 0.022 1.00
and other cacti (Echinocactus platyacanthus, Fero-
cactus robustus, F. recurvus, F. flavovirens, Cephalo-
cereus chrysacanthus, Pachyrereus hollianus,
vals for ␭ were relatively wide, thus, these matrices
Neobuxbaumua tetetzo, N. macrocephala) show the
appear to represent populations that are close to the
same type of germination behaviour (Rojas-Aréchiga
numerical equilibrium. The fourth matrix, P-1997/98,
et al. 1997, 1998; Godínez-Alvarez and Valiente-Ban-
had a positive ␭ value which was significantly differ-
uet 1998; Esparza-Olguín et al. 2002). However, in
ent from unity, representing a growing population. In
natural conditions the frequently low soil moisture
this case the high ␭ may be accounted for by the high
content appears to dramatically decrease the probabil-
fecundity and seedling survival observed in this site
ity of successful germination.
for the second study period. In turn, these fecundity
After seed germination, very few seedlings survive
values were given by high seed germination percent-
the harsh conditions imposed by high solar radiation
ages in the field. These results could have been re-
and low nutrient and water availability (Steenbergh
lated to two particular events. First, in 1997/98 the
and Lowe 1969; Stennbergh and Lowe 1977; Jordan
results of the field experiments on seed germination
and Nobel 1981; Valiente-Banuet and Ezcurra 1991).
and seedling survival were recorded daily; thus, seed
The results of our field experiments on seed germina-
germination (and therefore fecundity) might have
tion and seedling survival suggest that the probability
been underestimated the previous year, in which re-
that a germinated seed reaches the stage of a success-
sults were recorded every two weeks. However, note
fully established seedling is tremendously low. Sev-
that the general increase in fecundity observed in
eral factors may determine this. First, cacti seedlings
1997/98 did not result in a positive ␭ in the case of
show very low relative growth rates (Jordan and No-
the population in the disturbed site, which suggests
bel 1981; Nolasco et al. 1996), and thus the highly
that the increase in population growth rate observed
fragile phase of seedling establishment becomes ex-
in P-1997/98 was not an artefact of the differences in
traordinarily long (relative growth rate of M. magni-
the frequency of experimental observations between
mamma seedlings is ca. 0.015 g g −1 d −1 – Ruedas et
the two years. The second event that could have de-
al. (2000)). Secondly, seedlings show a relatively low
termined these results was that rainfall was higher in
water use efficiency due to C 3 metabolism during the
1998 than in 1997, as reported above. This could have
early stages of seedling development (Altesor et al.
determined the high seed germination and seedling
1993). Thirdly, the high water content of cacti tissues
survival values obtained for 1997/98. Water availabil-
determines a high incidence of seedling predation
ity is clearly one of the most important limiting fac-
(Valiente-Banuet and Ezcurra 1991; Mandujano et al.
tors for the growth and establishment of cacti species
1996). As a result, seedling survival probability vary
(Valiente-Banuet and Ezcurra 1991; Mandujano et al.
greatly from year to year or between different micro-
1996); the reduced water retention capacity of the
sites. Our results show that for M. magnimamma, this
bare basaltic substrate at ⬙El Xitle⬙ lava-field deter-
variation may be of two orders of magnitude (i.e.
mines that water availability is directly related to the
from 0.001 to 0.134), which accounts for an impor-
amount of rainfall.
tant variation in ␭ between years or between sites.
The most important differences in population
The variation in fecundity values also resulted in
growth rate between years and between sites appears
important changes in ␭. Fecundity was given by two
182
main components: seed production and seed germi-
nation. Although fruit and seed production showed
only small variations, seed germination probabilities
(as estimated from our field experiments) varied 100-
fold, ranging from 0.002 to 0.157. This variation was
the main reason for the observed changes in fecun-
dity between years and between sites.
The elasticity values of fecundity elements were
low in all cases, especially in the matrices that had a
␭ value below unity. This appears to suggest that
changes in fecundity entries would have only a minor
effect on ␭, yet, when fecundity increased we ob-
served a significant increase in ␭. However, note that
fecundity varied from 0.1 to 41.5 seedlings per plant,
which represents a ca. 1000-fold variation, whereas
the value of other matrix entries changed little be-
tween years or between sites (de Kroon et al. 2000).
For instance, the persistence of adults in category 7
ranged from 0.88 to 0.97, and although this particular
entry had high elasticity values, the changes in this
entry did not account for much variation in ␭. The
high elasticity values of this entry in the matrices with
␭ < 1 suggest that those populations, given the lack
of seedling recruitment, rely mainly on the survival
of old adults and so population numbers would de-
cline at the rate these adults disappear. A different
picture emerges when analysing matrix P-1997/98,
which showed a ␭ > 1. In this case the elasticity val-
ues for fecundity entries added up to 20% of total
elasticity, and the elasticity of seedling establishment
was also particularly high. As other authors have
stated, this implies that the dynamics of the early
stages of plant development are fundamental for a
population that is growing, and changes in such
stages dramatically affect population growth rate
(Horvitz and Schemske 1995; Silvertown et al. 1996;
Valverde and Silvertown 1998; Mandujano et al.
2001).
When incorporating the observed spatio-temporal
variation in the demography of M. magnimamma into
numerical simulations of overall population behav-
iour, allocating the same probability of occurrence to
all four matrices, we obtained a population growth
rate of ␭ s = 1.09. The average population growth rate
of the four matrices (␭ a = 1.05) did not differ much
from the value obtained from this stochastic analysis.
Yet, it is important to note that this average popula-
tion growth rate does not offer any insights into the
temporal variation in population size. On the other
hand, the results of the different stochastic simula-
tions performed in this study, which resulted in a
rather erratic variation in population numbers, offer a
more complete picture of the species’ actual demo-
graphic behaviour. In these cases, the variation in
population size which resulted from the use of differ-
ent matrices for the iteration process, was mainly a
result of changes in seedling establishment success,
which was the main difference between matrices. This
phenomenon has also been documented for other
cacti species (Ferocactus cylindraceus and Carnegia
gigantea – Bowers (1997) and Pierson and Turner
(1998)). The underlying causes of this type of numer-
ical fluctuations in populations of wild plants have
long been a concern of plant population ecologist.
Although there is a great lack of empirical evidence
on this issue, our simulation results suggest that tem-
poral variation in seedling survival (including all the
life-cycle phases involved in it) may be responsible
for the erratic numerical behaviour observed in many
plant populations.
In the first run of stochastic simulations we gave
the same probability of occurrence to each of the four
matrices. However, each matrix represent a particular
scenario that may occur at different frequencies. In
1998 the rainfall was above average, while in 1997 it
was below average. So an important aspect to incor-
porate in the simulations was the relative frequency
with which rainy and dry years occur. Additionally,
there are years with widespread fire events, while in
other years fires are kept to a minimum. Thus, the for
matrices represent rainy years with low fire impact
(P-1997/98), rainy years with high fire impact (D-
1997/98), dry years with low fire impact (P-1996/97)
and dry years with high fire impact (D-1996/97). Al-
though we could have access to the information on
the relative frequency of rainy and dry years at the
site, no information is available so far concerning the
frequency and intensity of fire events. Thus we simu-
lated the overall population growth rate under differ-
ent theoretical fire probabilities (ranging from 0.20 to
0.66). In all cases the analyses yielded ␭ s values
above unity; however, as fire frequency increased,
population persistence probability decreased. If fires
were as common as two every three years, the likeli-
hood that the M. magnimamma population would per-
sist in the long run would be unacceptably low.
Regarding the stochastic simulations, it is impor-
tant to take into account that the four matrices used
are certainly not the only possible ways in which the
demography of M. magnimamma could be character-
ised. A more complete picture would emerge if popu-
lation dynamics was evaluated over longer periods of

time and over a wider range of conditions within the
⬙Pedregal de San Angel⬙ reserve.
Cacti species are generally thought to be highly
vulnerable to disturbances. The results of our study
support this idea, since the only matrix that showed a
␭ significantly higher than unity was the one that cor-
responded to a population in a well preserved site and
during a particularly favourable year. The long-term
fate of this species at the ⬙El Xitle⬙ lava-field would
thus depend on the proportion of the reserve area that
remains ⬙well preserved⬙ and the way in which
weather patterns and disturbances affect seedling es-
tablishment success through time. Comparing the
present abundance of M. magnimamma at the study
site (600 plants/ha – Valverde et al. (1999)) with that
observed in 1954, it has been noted that its overall
density appears to be declining (Rzedowsky, personal
communication). The increasing disturbance fre-
quency at the site has determined the local extinction
of a number of native species while favouring the ar-
rival of ruderal species (Valiente-Banuet and de Luna
1990). Thus, a great effort should be made to decrease
the disturbance frequency and preserve the conditions
adequate for the establishment and growth of the na-
tive vegetation if the reserve at the ⬙El Xitle⬙ lava-
field is to maintain its original ecological interest and
biodiversity.
Acknowledgements
This project was carried out with the support of CON-
ACyT (5-3181 PN). Two of the authors (MLV and
SQ) received support from student grants (DGEP-
UNAM and CONACyT, respectively). We are grate-
ful to Ana Mendoza for comments and suggestion on
a previous version of this manuscript. We also wish
to thank Mariana Hernández, Pedro Eloy Mendoza,
Marco Antonio Romero, Lucrecia Trejo, Ligia Es-
parza, Elena Vilchis, Ariel Arias, Yvonne Vargas,
Marcela Ruedas and Cinthya Contreras for valuable
assistance in different stages of the study.
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