Trachelomonas (Euglenophyta) from a eutrophic reservoir in Central Mexico

Abstract

This study provides valuable information on the ultrastructure and environmental conditions of the Trachelomonas Ehr. (Euglenophyceae) genus in the Guadalupe Dam, a eutrophic reservoir located in the suburbs of Mexico City, which receives a considerable volume of wastewaters. Specimens were collected at surface level between November 2005 and May 2006. Using LM and SEM twelve taxa from phytoplankton were identified of which, 9 are new records for Mexico. The reservoir is warm monomictic, with basic pH values (7.4-10.1), a high concentration of chlorophyll a(18-101 microg l(-1), a permanent anoxic bottom, specific conductivity (K25) of 205 to 290 microS cm(-1), N-NO3, 0.19-1.2 mg l(-1) and P-PO4 0.22-1.6 mg l(-1). Water temperature was 15.6-23.0 degrees C. Most of the Trachelomonas species were found during the dry season, when concentrations of organic matter, nitrogen and phosphorus as well as the temperature were the highest. Higher species richness was also associated with the warmer months. This research contributes to increase our knowledge on Trachelomonas in Mexico and constitutes the first detailed description of lorica ultrastructure of 12 taxa that grow in a body of water with high concentration of nutrients and a moderate amount of mineral contents.

Full text

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Trachelomonas (Euglenophyta) from a eutrophic reservoir in Central Mexico
Author Details
Gloria Garduno Solórzano Facultad de Estudios Superiores Iztacala, UNAM. Av. de los Barrios No. 1, Col. Los Reyes Iztacala,
(Corresponding author) Tlalnepantla de Baz, Estado de México, 54090
e-mail: ggs@servidor.unam.mx
Maria Guadalupe Oliva Martinez Facultad de Estudios Superiores Iztacala, UNAM. Av. de los Barrios No. 1, Col. Los Reyes Iztacala,
Tlalnepantla de Baz, Estado de México, 54090
Alfonso Lugo Vazquez Facultad de Estudios Superiores Iztacala, UNAM. Av. de los Barrios No. 1, Col. Los Reyes Iztacala,
Tlalnepantla de Baz, Estado de México, 54090
Maria Berenit Mendoza Garfias Instituto de Biología, UNAM, 3
er
. Circuito escolar s/n, Ciudad Universitaria, Del. Coyoacán, México
D.F., 04510
Rafael Emiliano Quintanar Zuniga Facultad de Estudios Superiores Iztacala, UNAM. Av. de los Barrios No. 1, Col. Los Reyes Iztacala,
Tlalnepantla de Baz, Estado de México, 54090
Visitacion Conforti Facultad de Ciencias Exactas y Naturales (UBA-CONICET), Dpto. de Biodiversidad y Biología
Experimental Buenos Aires, Argentina
Abstract
This study provides valuable information on the ultrastructure and environmental conditions of the Trachelomonas
Ehr. (Euglenophyceae) genus in the Guadalupe Dam, a eutrophic reservoir located in the suburbs of Mexico
City, which receives a considerable volume of wastewaters. Specimens were collected at surface level
between November 2005 and May 2006. Using LM and SEM twelve taxa from phytoplankton were identified
of which, 9 are new records for Mexico. The reservoir is warm monomictic, with basic pH values (7.4-10.1),
a high concentration of chlorophyll a (18-101 µg l
-1
), a permanent anoxic bottom, specific conductivity (K
25
) of 205
to 290 µS cm
-1
, N-NO
3
0.19-1.2 mg l
-1
and P-PO
4
0.22-1.6 mg l
-1
. Water temperature was 15.6-23.0
o
C. Most
of the Trachelomonas species were found during the dry season, when concentrations of organic matter, nitrogen
and phosphorus as well as the temperature were the highest. Higher species richness was also associated with
the warmer months. This research contributes to increase our knowledge on Trachelomonas in Mexico and
constitutes the first detailed description of lorica ultrastructure of 12 taxa that grow in a body of water with high
concentration of nutrients and a moderate amount of mineral contents.
Key words
Lorica, Dam, Euglenoids, Phytoplankton, Ultrastructure
Publication Data
Paper received:
28 October 2009
Revised received:
25 June 2010
Accepted:
23 September 2010
Introduction
There are many records of Trachelomonas species in the
world, which are mostly based on observations carried out using
light microscopy (LM). Unfortunately, with this method it is difficult to
achieve a detailed study of the lorica structure, which is the most
reliable basis for the taxonomy of the genus (Wolowski and Hindák,
2004). Some species are endemic; others are cosmopolitan or
restricted to cold, temperate or warm regions (Couté and Tell, 2006).
More than 200 species are described, in the “Monographie
du genre Trachelomonas Ehrenberg” (Deflandre, 1926). The most
important studies on these organisms took into consideration the
size, shape and ornamentation of lorica as the main characters for
their classification (Conrad and Van Meel, 1952). Huber-Pestalozzi
(1955) compiled 256 species, 190 varieties and 46 forms. Rosowski
et al. (1975) were the first to study the details of the lorica surface
using a scanning electron microscope (SEM). Ever since various
species of Trachelomonas have been examined with this method.
Some works on the study of Trachelomonas in the world have been
carried out in South America (Tell and Couté, 1980; Couté and
Thérézien, 1985, 1994; Conforti, 1993, 1999; Conforti and Nudelman,
1994; Conforti and Perez, 2000; Conforti and Tell, 1986), in North
© 2011 Triveni Enterprises
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Full paper available on: www.jeb.co.in
J. Environ. Biol.
32, 463-471 (2011)
ISSN: 0254-8704
CODEN: JEBIDP

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America (Conforti and Joo, 1994; Wolowski and Walne, 2007), in
Europe (Kocárková et al., 2004; Wolowski and Hindák, 2004;
Wolowski and Grabowska, 2007), in Asia (Kim et al., 1999; Conforti
and Ruiz, 2001), and in Africa (Couté and Iltis, 1981; Da et al.,
2009).
It is important to note that there is not very much information
available on the taxonomical, ecological and geographical distribution
of the Trachelomonas genus in Mexico. A list of 27 taxa has been
recorded in Mexico (Ortega, 1984; Díaz-Pardo et al., 1998; García-
Rodríguez and Tavera, 2002; Schmitter-Soto et al., 2002; García-
Rodríguez et al., 2003; Moreno-Ruiz, 2005; Quiroz-Castelán et al.,
2007; Moreno-Ruiz et al., 2008). Two species were found to have
been reported previously in Guadalupe Dam by Lugo et al. (1998,
2007). The present paper contributes to the knowledge of the
Trachelomonas species composition in Mexico. For the first time our
country, details of the lorica ultrastructure of the species are described,
using LM and SEM. Likewise, the main physicochemical conditions
found during the study are also provided.
Materials and Methods
Study area: The Guadalupe Dam is located outside Mexico City in
the suburbs, in the State of Mexico (19
o
48’ 30’’ N 99
o
15’ W, 2350
m.a.s.l., maximum volume 60 X 10
6
m
3
, maximum depth 20 m). The
dam was built to control and store the waters of the Cuautitlan River
for irrigation (Fig. 1). Currently, a high percentage of the total annual
inflow comes from sewage discharges from a highly populated area
around the dam (Lugo et al., 2007).
The climate is temperate subhumid with a rainy summer.
Mean annual temperature is 16
o
C, and annual precipitation is of 706
mm (Hidalgo-Wong and Pulido-Navarro, 2006). The reservoir is
eutrophic with high concentrations of chlorophyll a (Lugo et al.,
1998).
At the beginning of the 80’s the dam was invaded by water
hyacinth Eichhornia crassipes. The Department of Agriculture (SARH)
together with the Mexican Institute of Water Technology (IMTA) in
1993 successfully implemented the first chemical control program,
through the application of herbicides such as 2, 4 D and diquat. In
1995, the weed reappeared, and in 1997 it was eliminated again by
chemical and mechanical weed control. Ever since, hyacinth has not
invaded the water surface again, however other ecological problems
such as blooms of algae and the death of fish during the winter
seasons in 2004 and 2005 have arisen (Lugo et al., 2007).
Sample collection: Samples were collected with 20 µm mesh
plankton net at surface level in November 2005, April and May
2006. The material was fixed in 4% formaldehyde. Detailed
examination of the material was carried out with a Zeiss phase-
contrast microscope (LM). On the other hand, for scanning electron
microscopy (SEM) analysis, a concentrated subsample was filtered
using Millipore® filters (0.45 µm pore size), and air-dried. The pieces
of filters were adhered to aluminum stubs and coated with gold
(Zalocar de Domitrovic and Conforti, 2005). A Hitachi S-2460N
electron microscope, operating at 15kV at the Institute of Biology,
UNAM and a JEOL JSM6380LV at the Electron Microscopy Service
(FES Iztacala, UNAM) were used. The samples were deposited at
the IZTA herbarium with reference numbers 1753-1764 (Holmgren
et al., 1990).
Environmental conditions were recorded at the same time
the samples of Trachelomonas were collected. Water temperature,
dissolved oxygen and conductivity (K
25
) were measured using a
YSI multisonde mod. 85 (Yellow Spring Instruments Co. Ohio, USA).
And transparency was measured using a Secchi disk.
The content of chlorophyll a was evaluated in vivo with a
portable Aquafluor fluorometer (Turner Designs Co. California USA).
N-NO
3,
P-PO
4
and BOD
5
samples were obtained from five sites at
the surface and sent to the laboratory to be analyzed. N-NO
3
as well
as total dissolved phosphorous was measured using the cadmium -
reduction method for the first and the ascorbic acid method for the
latter. BOD
5
was evaluated with the bottle dilution method (APHA,
1989).
The identification and distribution of the species was based
on the research works by Conrad and Van Meel (1952); Hüber-
Pestalozzi (1955); Couté and Iltis (1981); Tell and Zalocar de
Domitrovic (1985); Tell and Conforti (1986); Conforti and Tell (1986,
1989); De la Rosa and Sanchez-Castillo (1991); Conforti (1993,
1999); Conforti and Joo (1994); Conforti and Nudelman (1994);
Couté and Thérézien (1994); Kim et al. (1999, 2000); Dillard (2000);
Conforti and Ruiz (2001); Kocárková et al. (2004); Wolowski and
Hindák (2004); Da et al. (2009) and Algaebase.org. The terminology
used in this paper follows Conforti and Tell’s criterion (1986). In
average, 35 organisms were used for measurements. As far as
scientific names and synonyms are concerned, same were verified
in the Integrated Taxonomic Information System and Index Nominum
Algarum.
Results and Discussion
Environmental conditions: The Guadalupe Dam is a warm
monomictic water body. In November, at the beginning of the
cold dry season, the water column was thermally homogeneous
at around 17
o
C (16.9-18.8
o
C) which means that mixing
conditions were prevalent. In March, and particularly during
April (15.6-21
o
C) and May (17-23
o
C), there was a thermal
stratification.
The high organic load at the bottom of the dam causes
permanent anoxia in the deep water layer. In November, dissolved
oxygen ranged from non detectable (n.d.) to 12 mg l
-1
in the top 10
m of the water column; however during April and May oxygen was
detected only in the top five meters. K
25
values in November were
lower (205-241 µS cm
-1
) than in April and May (250-290 µS cm
-1
).
At the surface level pH was always basic, in November ranging from
7.4 to 9.3 and during April and May the values increased to 7.9-
10.1. Water transparency varied between 0.2 and 0.5 m indicating
a shallow euphotic zone.

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Trachelomonas from a eutrophic dam
Species richness of Trachelomonas was correlated with
nutrient concentration. In November, when low N and P
concentrations were measured (mean value: N-NO
3
0.186 mg l
-1
;
P-PO
4
0.216 mg l
-1
), only four species were observed. On the other
hand, during April and May with an increased N and P concentration,
(mean value: N-NO
3
0.7-1.2 mg l
-1
, P-PO
4
1.6 mg l
-1
), the number of
Trachelomonas species rose to six and ten respectively.
The concentration of Chlorophyll a was also related with
nutrients. In November the measured values were low, within a
range from 18 to 25 µg l
-1
. April showed the highest values (72-101
µg l
-1
) decreasing in May (30-45 µg l
-1
). These average values are
high and clearly show the eutrophic conditions of the dam (Margalef,
1983).
BOD
5
values measured occasionally in the dam, ranged
between n.d. to 65 mg O
2
l
-1
, indicating a moderate organic load in
the surface level. Wolowski and Hindák (2004) found that
Trachelomonas are generally resistant to organic pollution. Sládecek
(1973) considered Trachelomonas as a typical indicator of medium
to high organic matter concentrations in water, especially associated
with high ammonium concentrations (Alves-da-Silva et al., 2008;
Conforti and Tell, 1986; Conforti, 1986,1993; De la Rosa and
Sanchez, 1991; Da et al., 2009).
In temperate climates euglenoids are observed in the warmer
months (spring, summer) and only a few species can be found in
cold water or even under the ice (Starmach, 1983; Conforti and Tell,
1988). In this work we observed that the highest richness
Table - 1: List of taxa observed in Guadalupe Dam, the marked with an asterisk are new record for Mexico and P (presence).
Taxa November 2005 April 2006 May 2006
*T. globularis var. gigas Drezepolski 1923 P
T. hispida var. hispida (Perty) Stein 1878 P P P
*T. hispida var. coronata Lemmermann 1913 P P
*T. nexilis Palmer 1925 P
*T. rugulosa var. rugulosa Stein 1878 P
*T. rugulosa var. meandrina (Conrad) Conrad 1952 P
*T. rugulosa var. steinii Deflandre 1927 P
*T. similis var. spinosa Hüber-Pestalozzi 1955 P P
*T. sydneyensis Playfair 1915 P P P
*T. verrucosa f. irregularis Deflandre 1926 P
T. volvocina var. volvocina Ehrenberg 1833 P P
T. volvocina var. punctata Playfair 1915 P P
Total 4 6 10
Fig. 1: Location of Guadalupe Dam, Mexico

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Figs. 2-7. Figs. 2-3: Trachelomonas globularis var.gigas: 2- general view, 3- apical view, showing detail of the pore; Figs. 4-5: T. hispida var. hispida, 4-
general view, 5- apical view, showing detail of the pore surrounded by a short neck with spines Figs. 6-7: T. hispida var. coronata, 6- general view, 7- apical
view, showing detail of the neck. Scale bars = 10 µm (Figs. 2, 6), 5 µm (Figs. 3, 4), 2µm (Fig. 7), 1µm (Fig. 5).

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Figs. 8-13. Fig. 8: T. nexilis Fig. 9:T. rugulosa var. rugulosa Fig. 10: T. rugulosa var. meandrina Fig. 11: T. rugulosa var. steinii Figs. 12-13: T. similis var.
spinosa, 12- general view, 13- detail of the neck. Scale bars 5 µm (Figs. 8, 9, 10, 11, 12), 1µm (Fig. 13)

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Figs. 14-19. Figs. 14-15: T. sydneyensis, 14- general view, 15- detail of the neck Figs. 16-17: T. verrucosa fo. irregularis, 16- general view, 17- apical view,
showing detail of the pore Fig. 18: T. volvocina var. volvocina Fig. 19: T. volvocina var. punctata. Scale bars 5 µm (Figs. 14, 18), 2µm (Figs. 15, 16, 17, 19).
Garduno et al.

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Trachelomonas species was associated with the warmer months.
Only T. hispida and T. sydneyensis were observed during the entire
period that was (Table 1).
Taxonomical descriptions: During the study period 12 taxa of
Trachelomonas were identified, including species, varieties and forms;
nine of these taxa are new records for Mexico. Lugo et al. (1998)
found T. hispida and T. volvocinopsis in a previous survey in the
dam. This study confirms the presence of T. hispida and adds 10
taxa not observed previously. Trachelomonas globularis var. gigas
Drezepolski 1923 Fig. 2, 3.
Lorica spherical with 19 µm in diameter, wall with 80 100
-1
µm
2
punctae covered sparsely with 0.5-0.95 µm short conical spines.
Apical pore with a 2.5-3.7 µm diameter, surrounded by an annular
thickening (IZTA-1753).
Distribution: Argentina, Poland and the US.
The specimens studied here are smaller than those
described by Conforti (1999) with a 31-32 µm diameter and Dillard
(2000) 34 µm diameter. Trachelomonas hispida var. hispida (perty)
Stein 1878 Fig. 4, 5.
Lorica elliptical; 22-25 µm long, 16.5-21 µm wide, with a
wall covered with 72-100 100 µm
2
punctae, 0.5-1.3 µm short conical
spines, uniformly distributed 8-20 100 µm
2
. Apical pore with a 3.2-
4.2 µm diameter, surrounded by an annular ring-like thickening
(IZTA-1754).
Distribution: Cosmopolitan. It was previously reported in the
Chapultepec and Xochimilco Lakes in Mexico City; the Guadalupe
Dam in the State of Mexico; El Rodeo Lagoon in the State Morelos;
in Tulancingo, in the State of Hidalgo; Tonatihua and Zempoala
Lagoons in the State of Morelos; González River and Mandinga
respectively in the States of Tabasco and Veracruz in Mexico.
Trachelomonas hispida var coronata Lemmermann 1913 Fig. 6,7.
Lorica elliptical; 28-30 µm long, 21-21.5 µm wide, with a
rounded or acuminate posterior end, wall 88 100 µm
2
punctae
covered with conical spines 1.5-3 µm high with a diameter 0.75-1.5
µm at base, uniformly distributed 16 100 µm
2
. Apical pore surrounded
by a short 1-2.5 µm high and 5 µm wide neck, with sharp 1.5 µm
long spines along its margin (IZTA-1755).
Distribution: Britain, Romania, Spain, Africa, Australia,
Argentina, Portugal, New Zealand and US.
In the specimens described by Da et al. (2009) the punctae
are 270-354 µm/100 µm
2
. Our specimens showed 88/100 µm
2
.
Trachelomonas nexilis Palmer 1925 Fig.8.
Lorica spherical with 9-17 µm in diameter, wall with irregularly
0.15-0.17 µm wide vermicular lines and depressions. Apical pore 2
µm wide surrounded by an annular thickening (IZTA-1756).
Distribution: Argentina, Brazil, Spain and the US.
Trachelomonas rugulosa var. rugulosa Stein 2878 Fig.9.
Lorica spherical with 14-15 µm in diameter, wall covered
with distinct anastomosing ridges arranged longitudinally or spirally
0.65 µm thick and separated by an average of 0.7 µm, ten ribs by
10 µm, twisted in the anterior end. Apical pore 2 µm in diameter
surrounded by an annular thickening (IZTA-1759).
Distribution: Britain, Romania, Spain, Argentina and the US.
Trachelomonas rugulosa var. meandrina (Conrad) Conrad 1952
Fig.10.
Lorica spherical with 24 µm in diameter, wall covered with
anastomosing, arranged folds. Apical pore with 2 µm in diameter
surrounded by a ring-like thickening (IZTA-1764).
Distribution: Britain, Slovakia. Trachelomonas rugulosa var.
steinii Deflandre 1927 Fig.11.
Lorica spherical with 15-18.5 µm in diameter, and densely
or lightly ornamented wall with anastomosing ridges coming radially
from the pore. Apical pore 1.5-2 µm diameter, surrounded by an 0.8
µm annular thickening (IZTA-1760).
Distribution: Colombia, Korea, France and Austria.
Trachelomonas similis var. spinosa Hubar-Pestalozzi 1955 Fig.12,
13.
Lorica elliptical; 23.5-25 µm long and 19-20.5 µm wide,
with a rounded or slightly acuminate posterior end, 45-80/100 µm
2
punctae covered with a few short 1.6-2 µm long conical spines with
a diameter of 0.50-0.80 µm at the base, sparsely distributed 10-13/
100 µm
2
. Apical pore surrounded by a cylindrical neck usually
curve towards one side, 2-4.5 µm wide and 2-5.0 µm high, 0.20-
0.35 µm thick wall with spines irregularly distributed spines at the
end of the neck which has a thick rim where there are conical spines
up to 2 µm long (IZTA-1757).
Distribution: Africa, Argentina, Bolivia, Brazil, Colombia, the
US and Venezuela. Trachelomonas sydneyensis Play fair 1915
Fig. 14, 15.
Lorica elliptical; 28-33 µm long, 20-24 ``µm wide, the
rounded poles and covered with long and sharp conical spines in
the poles and smaller ones 1-3 µm long, 12-24/100 µm
2
in the
center; 56-94/100 µm
2
punctae. Apical pore surrounded by a
conspicuously divergent neck 1-3.5µm high and 5-7.5 µm wide in
its opening and spines in the rim (IZTA-1758).
Distribution: Africa, Asia, Argentina, Australia, Brazil, Britain,
New Zealand, Romania and Spain.
In the specimens described by Da et al. (2009) the punctae
are 145-181/100 µm
2
. Our specimens showed 56-94/100 µm
2
.
Trachelomonas verrucosa F. irregularis Deflandre 1926 Fig. 16,
17.
Lorica spherical with 11-11.5 µm in diameter, ornamented
wall with small uniformly distributed 300/100 µm
2
warts. Apical pore
Trachelomonas from a eutrophic dam

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1-1.5 µm diameter, surrounded by an annular thickening (IZTA-
1761).
Distribution: Africa, Slovakia and US.
In the specimens described by Conforti and Joo (1994) the
warts were more dense (520/100 µm
2
) than in our material.
Trachelomonas volvocina var. volvocina Ehrenbeig 1833 Fig. 18.
Lorica spherical with 10-22 µm in diameter and smooth wall.
Apical pore with 1-2 µm in diameter, surrounded by an annular
thickening. Two lateral chloroplasts have double sheathed pyrenoids.
Flagellum three times longer than the lorica (IZTA-1762).
Distribution: Cosmopolitan. In Mexico it was recorded in the
Chapultepec and Xochimilco Lakes in Mexico City; Lerma wetland
in the State of Mexico; Tulancingo, State of Hidalgo; in the El Rodeo
Lake, Tonatihua and Zempoala Lagoons in the State of Morelos;
Labradores Lake in the State of Nuevo León; Tehuantepec River in
the State of Oaxaca; Gonzalez River in the State of Tabasco and
Apizaco in the State of Tlaxcala in Mexico. Trachelomonas volvocina
var. punctata Play fair 1915 Fig.19.
Lorica spherical with 13-15 µm in diameter, 130-135/100
µm
2
punctae Apical pore with 2.5 µm diameter, surrounded by a low
neck. Two lateral chloroplasts have double sheath pyrenoids (IZTA-
1763).
Wolowski and Hindák (2004) observed 200-300/100 µm
2
punctae, our specimens had a lower number of 66-132/100 µm
2
punctae.
Distribution: Argentina, Australia, Romania, Spain, New
Zealand, Denmark, Slovakia, Switzerland, Turkey, US, Venezuela.
In Mexico it was reported at the Tehuantepec River in the State of
Oaxaca.
All the studied species are widespread or cosmopolitan,
although T. rugulosa var. meandrina has only been observed in
Britain and Slovakia. In this work the geographical distribution is
extended in North America.
The number of taxa of Trachelomonas in the Guadalupe
Dam was intermediate, in comparison with Alves-da- Silva and
Schüler- da- Silva (2007) who found only nine taxa Trachelomonas
in 26 water bodies of the Jacuí Delta State Park of the Río Grande do
Sul State in Brazil. In several shallow lagoons of Granada in Spain
De la Rosa and Sanchez-Castillo (1991) observed only 5. Kocarková
et al. (2004) found 25 taxa Trachelomonas in ponds and puddles of
the north region of Moravia in the Czech Republic. On the other
hand, Conforti (1993) reported the presence of 90 taxa in the
Camaleao Lake in Manaos, Brazil, and Conforti and Ruiz (2001)
found 41 taxa in the Chuman reservoir in South Korea.
In Mexico, the usual number of taxa found per water body
is two with the exception of the Tehuantepec River where 8 taxa
have been found (Moreno-Ruiz et al., 2008). In the present study
the information obtained with the use of SEM allowed us to increase
our knowledge and characterization of the observed taxa. On the
other hand, in previous studies, where only LM was used, it is quite
likely that the number of Trachelomonas taxa could have been
subestimated.
The Guadalupe Dam was infested with water hyacinth
Eichhornia crassipes for more than 12 years. In 1993 a program to
remove hyacinth was carried out at reservoir including the use of
herbicides and mechanical control. Changes in the habitat conditions
and the disappearance of the hyacinth have promoted an increase
in the diversity and abundance of phytoplankton (Lugo et al., 1998).
There has been an increase in the number of species of
Trachelomonas under the new environmental conditions, from two
to twelve species. The present study shows an important increase in
the diversity of the Trachelomonas species most likely associated
with the presence of better environmental conditions for phytoplankton
growth.
Acknowledgments
This research was financially supported by project 18330-
615-11-V-06 of the municipality of Cuautitlán Izcalli. The authors
wish to thank Biol. Peter Michael Mueller Meier, Laboratory of Scientific
Photography at FES Iztacala UNAM for his skillful support in the
preparation of the illustrations. This paper greatly benefited from the
comments and critical revision of the manuscript by Dra. Margarita
Caballero from the Institute of Geophysics at UNAM and two
anonymous reviewers for their valuable observations.
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