Amphibia-Reptilia 38 (2017): 225-231

Short Notes

Potential distribution of the endemic imbricate alligator lizard

(Barisia imbricata imbricata) in highlands of central Mexico

Armando Sunny1,∗ , Andrea González-Fernández2 , Maristella D’Addario1

Abstract. Barisia imbricata imbricata is an endemic anguid lizard living in the highlands of central Mexico, mainly in the
Trans-Mexican Volcanic Belt, but its current distribution is not well defined. For this reason, it is critical to make a systematic
study in order to determine the potential distribution of this subspecies. We considered 13 environmental and anthropogenic
variables recognized to be important to determine the presence of B. i. imbricata; we also used eight presence-only modeling
methods and selected three to generate an ensemble model. The Maxent algorithm was used to obtain the most important
variables. The variables with highest contribution to the model were: maximum temperature of the warmest month, altitude,
Abies forest and Pinus forest. From the estimated potential distribution (9614.1 km2 ), 8053.4 km2 (83.8%) are located in the
Trans-Mexican Volcanic Belt, 3821.4 km2 (39.7%) are located in natural protected areas and 2418.2 km2 (25.2%) are in crop
fields. The whole potential distribution is highly fragmented and characterized by the presence of human activities, which
could have long-term consequences, increasing extinction risk.

Keywords: conservation, ecological niche model, endemic species, ensemble model, habitat suitability.

Land-use change is the main factor affecting
the distribution and diversity of reptiles (Castel-
lano and Valone, 2006; Ribeiro et al., 2009).
This group is extremely sensitive to local habi-
tat changes because of its low dispersal capacity
and small home ranges (Huey, 1982; White et
al., 1997; Castellano and Valone, 2006; Ribeiro
et al., 2009). Some studies indicate that be-
tween 15 and 44% of the world’s reptile species
are threatened (Böhm et al., 2013; Ceballos et
al., 2015). Habitat loss has the greatest nega-
tive effect on biodiversity related to land-use
change (Fahrig, 2003). However, habitat frag-
1 - Centro de Investigación en Ciencias Biológicas Apli-
cadas, Universidad Autónoma del Estado de México,
Instituto literario # 100, Colonia Centro, 50000 Toluca,
Estado de México, México
2 - Laboratorio de Biología Evolutiva, Facultad de Cien-
cias, Universidad Autónoma del Estado de México, In-
stituto literario # 100, Colonia Centro, 50000 Toluca,
Estado de México, México
∗ Corresponding author; e-mail:
sunny.biologia@gmail.com
© Koninklijke Brill NV, Leiden, 2017.
mentation could also have long-term conse-
quences, including increased inbreeding prob-
ability (Frankham, Ballou and Briscoe, 2005;
Allendorf and Luikart, 2007) and loss of ge-
netic diversity, thereby increasing extinction
risk (Newman and Tallmon, 2001). Natural pro-
tected areas can help to preserve habitat patches
of threatened reptile species and minimize the
factors that are causing their decline. Therefore,
it is necessary to know the potential distribution
of a species in order to make informed man-
agement decisions for its conservation. Unfor-
tunately, basic ecological information is insuffi-
cient and the actual distribution of many reptiles
is unknown or imprecise.
It is essential to determine accurately the dis-
tribution of B. i. imbricata to ensure its long-
term conservation, especially because this en-
demic anguid lizard lives in the highlands of
central Mexico, mainly in the Trans-Mexican
Volcanic Belt (TMVB), one of the areas most
disturbed by grazing, agriculture and the pres-
ence of the biggest metropolitan areas of the
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DOI:10.1163/15685381-00003092
226
Figure 1. Potential distribution of B. i. imbricata showing ANPs and biogeographic provinces used to build the distribution
model with the training points (circles) and testing points (triangles).
country (CONAPO, 2010; Sunny et al., 2015).
The TMVB is also one of the biogeographic
zones with the highest species richness and en-
demism across the country; for example, it is
the most important region in terms of endemic
amphibian and reptile species and the second
in terms of species number (Flores-Villela and
Canseco-Márquez, 2007). B. i. imbricata inhab-
its mixed Abies-Pinus woodlands, its altitudinal
range is from 1800 to 3000 m a.s.l. and it is
listed under special protection by Mexican law
(SEMARNAT, 2010), but its threats and the ac-
tual distribution of its populations are unknown
(Canseco-Márquez et al., 2007).
The hypothesis of this study is that anthro-
pogenic activities have fragmented the distri-
bution range of B. i. imbricata, leaving iso-
lated populations in patches of Pinus-Abies for-
est. The objectives of this study were: (1) to
determine the potential current distribution of
B. i. imbricata, using ecological niche model-
ing; (2) to determine the most important habitat
features associated with the presence of B. i. im-
bricata; and (3) to compare the predicted distri-
Short Notes
bution with the current system of natural pro-
tected areas in Mexico.
To address these issues, we used information from
databases and data collected in fieldwork. Firstly, we de-
fined the study area, considering the biogeographic zones
that include records of B. i. imbricata and the surrounding
biogeographic zones where migration could take place. The
biogeographic zones are: Altiplano Sur (As), Sierra Madre
Oriental (SMor), Golfo de México (GM), TMVB, Oaxaca
(Ox), Depresión del Balsas (DB) and Sierra Madre del Sur
(SMs), with a total area of 673 690.9 km2 (fig. 1). These
biogeographic zones have a variety of habitats, such as
high mountain forests, deciduous forests and cloud forests,
among others (INEGI, 2013).
Secondly, we obtained historical information about
records of B. i. imbricata from scientific literature (Ramírez-
Pérez, 2008; Bryson and Riddle, 2012; Cruz-Elizalde
and Ramírez-Bautista, 2012; Sánchez-Jasso et al., 2013;
Muñoz-Brito, 2014), databases of herpetological collec-
tions (REMIB, UNIBIO, GBIF, HERPNET, VERTNET and
IREKANI) and field work carried out from January 2013 to
September 2014. The sampling localities were in the moun-
tains and natural parks of the TMVB, in areas where small
alpine grassland species were found (Muhlenbergia sp.).
Thirdly, the field surveys were not conducted in the
same sites as those reported in literature or databases, nor
near those sites, to avoid overfitting. When we found an
imbricate alligator lizard, we recorded the geographical
coordinates. Between 1964 and 1990 significant changes in
land use and vegetation cover occurred in Mexico (FAO,
2001; Domínguez-Vega et al., 2012), so we used records
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Short Notes
only from 1990 to 2014. We filtered these data to obtain only
one record per km2 to reduce the spatial correlation between
them (Boria et al., 2014). The records obtained were divided
into two groups. The first group was comprised of the oldest
data (75%) and was used to calibrate the models, the second
group was comprised of the most recent data (25%) and was
used to validate them (fig. 1).
Finally, we considered 13 environmental and anthro-
pogenic variables from the Vegetation and Land Use layer
(series V, 1:250 000) generated by the National Institute
of Statistics and Geography, based on satellite images with
a pixel resolution of 120 meters (Landsat TM5) during
the period 2011 to 2013 (INEGI, 2013). We reclassified
the vegetation map into eight mutually exclusive classes
expressed as proportions (agriculture, cloud forest, Abies
forest, Quercus forest, Quercus-Pinus forest, Pinus forest,
Pinus-Quercus forest and grassland). We also included dis-
tance to water sources. Climatic variables, such as annual
precipitation, maximum temperature of the warmest month
and minimum temperature of the coldest month, were ob-
tained from WorldClim (Hijmans et al., 2005). Finally, we
obtained altitude from the USGS/NASA Shuttle Radar To-
pography Mission (USGS, 2007).
No algorithm has been demonstrated to be the best to
define potential distributions in all cases (Hernandez et al.,
2008). When the dataset is relatively small, models could
overfit and predict very different suitabilities, even with sim-
ilar performances. In these cases, the use of an ensemble
model is a good method to avoid relying too much upon
any single model. Therefore, we applied eight algorithms.
We used six algorithms of the software OPENMODELLER
1.0.8 (Sutton et al., 2007): Bioclim (Nix, 1986), climate
space model (Robertson, Caithness and Villet, 2001), en-
velope score (Nix, 1986), environmental distance (Hirzel
and Arlettaz, 2003), GARP best subsets new OPENMOD-
ELLER (Stockwell and Peters, 1999), and support vec-
tor machines (Vapnik, 1995). We used the Maxent algo-
rithm in the MAXENT 3.3.3 software (Phillips, Ander-
son and Schapire, 2006) with a logistic output, and cre-
ated response curves. Finally, we implemented the ecolog-
ical niche factor analysis (ENFA) using BIOMAPPER 4.0
(Hirzel, 2008). We evaluated the performance of each model
with the Area Under the Curve AUC/ROC (Receiver Op-
erating Characteristic, Hanley and McNeil, 1982). In or-
der to calculate AUC/ROC values, we used IDRISI Selva
17.0 software (Clark Labs, 2012). We used the weighted
average formula in the IDRISI calculator to elaborate the
final ensemble model, as it has been demonstrated that it
has the highest predictive performance of all the ensemble
methods (Marmion et al., 2009a, b; Zarco-González et al.,
2013). We included the highest three models with the high-
est AUC/ROC values; AUC/ROC values > 0.85 are consid-
ered to have moderate to high performance (Zarco-González
et al., 2013). To simplify the interpretation of the ensem-
ble model, we reclassified the original prediction values (0-
100) using the mean probability value of the records used to
calibrate the models, so that zones with a prediction value
greater than or equal to the mean were considered to be of
high suitability and those with lower values of low suitabil-
ity (Liu et al., 2005). Finally, to evaluate whether the current
227
NPAs contribute to the protection and conservation of the
subspecies, we assessed the percentage of high suitability
area that overlaps with NPAs.
We found a total of 1446 records and
worked with 126 records after filtering them:
70 records from databases of herpetological
collections acquired between 1990 and 2013,
40 records from scientific literature between
2007 and 2011, and 16 records from field
surveys carried out between 2013 and 2014
distributed in 12 states (Jalisco, Guanajuato,
Queretaro, Hidalgo, Estado de México, Ciu-
dad de México, Michoacán, Morelos, Puebla,
Tlaxcala, Oaxaca and Veracruz) and in four
biogeographic provinces. Our results indicated
that 8053.4 km2 (83.8%) of the total poten-
tial distribution were located in the TMVB,
544.9 km2 (5.7%) in the SMor, 413.2 km2
(4.3%) in the SMs, 366.1 km2 (3.8%) in the
As, 222.4 km2 (2.3%) in the Ox and 13.9 km2
(0.1%) in the DB. The algorithms with AUC
values below 0.850 were: ENFA (AUC/ROC =
0.697), BIOCLIM (AUC/ROC = 0.708), en-
velope scores (AUC/ROC = 0.777), environ-
mental distance (AUC/ROC = 0.786), climate
space model (AUC/ROC = 0.821). The algo-
rithms that performed best were: GARP best
subsets new OPENMODELLER (AUC/ROC =
0.854), support vector machines (AUC/ROC =
0.904), Maxent (AUC/ROC = 0.916), and the
ensemble model presented an AUC/ROC value
of 0.916. In this case, the high suitability area
(probability > 0.79) encompasses 9614.1 km2
(fig. 1); 3821.4 km2 (39.7%) of this area was lo-
cated in natural protected areas and 2418.2 km2
(25.2%) in crop fields (fig. 1). According to
ENFA, we obtained a global marginality value
of 1.8, which indicates that the observed con-
ditions in the occurrence sites are different from
the mean environmental conditions. We also ob-
tained a global tolerance value of 0.1, which
means that this subspecies has low tolerance to
changes in environmental variable values. The
variables with the highest contribution, accord-
ing to Maxent, were: the maximum temperature
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Figure 2. Variables with the highest contribution to the potential distribution of B. i. imbricata according to Maxent.
of the warmest month (52.3% contribution), al-
titude (21.9% contribution), the proportion of
Abies forest (17.2% contribution) and the pro-
portion of Pinus forests (3.9% contribution).
The probability of detection related to the max-
imum temperature of the warmest month was
high in temperatures below 23°C, the probabil-
ity decreased rapidly between 23 and 34°C, and
above 34°C the probability was 0 (fig. 2A). The
probability of detection related to altitude was
very low under 2000 m a.s.l. and high above
3200 m a.s.l. Between 2000 and 3200 m a.s.l.
there was a rapid increase in the probability of
detection (fig. 2B). There was a positive linear
correlation between the proportion of Abies for-
est and probability of detection (fig. 2C). There
was also a positive correlation between the pro-
portion of Pinus forest and the probability of de-
tection, but at the highest values of proportion
of Pinus forest there was a drastic decline in the
probability of detection (fig. 2D).
Short Notes
The AUC/ROC results indicate that the en-
semble model has a very accurate predictive
ability that could be better than the individ-
ual models. According to these results we pro-
pose the potential distribution of B. i. imbricata
as the plausible actual distribution of the sub-
species. Results indicate that the most impor-
tant variables are the warmest temperature of
the warmest month and altitude. This subspecies
occurs in areas with high elevation and cannot
tolerate temperatures higher than 34°C. B. i. im-
bricata is adapted to high elevations and low
temperatures because it evolved with the for-
mation of the TMVB in the Neogene (Bryson
and Riddle, 2012); in fact, the uplifting of the
TMVB is considered one of the most impor-
tant forces driving the evolutionary history of
several taxa (León-Paniagua et al., 2007; Mc-
Cormack et al., 2008; Navarro-Sigüenza et al.,
2008; Bryson et al., 2011; Bryson and Riddle,
2012), including B. i. imbricata (Bryson and
Riddle, 2012). The results obtained suggest that
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Short Notes
this subspecies could be very vulnerable to cli-
mate change because it is limited by high tem-
peratures. Moreover, the fact that its distribu-
tion already includes the existing areas with the
highest altitude implies that, as climate change
takes hold, this subspecies will be limited in its
ability to shift its distribution upward, increas-
ing the possibility of becoming extinct.
Although we usually find B. i. imbricata in
grasslands, this vegetation class was not se-
lected as an important variable to explain the
distribution of the sub-species. This may be be-
cause there are different kinds of grasslands; the
subspecies occurs in Muhlenbergia sp. grass-
lands but not in other kinds of managed grass-
lands where the grass is shorter. However, it is
not possible to distinguish between these kinds
of grasslands with satellite images.
The proportion of Abies religiosa forest
(Aguirre-Planter, Furnier and Eguiarte, 2000;
Aguirre-Planter et al., 2012) and Pinus forest
(Moreno-Letelier and Piñero, 2009) were im-
portant to define the distribution of the sub-
species since B. i. imbricata occurs only in
grasslands near these forests; this may be be-
cause coniferous forests favor moisture (annual
precipitation between 1000 and 3800 mm) and
a cold microclimate (2-24°C) (Sáenz-Romero et
al., 2012) preferred by the subspecies. There-
fore, the microclimatic conditions of grasslands
surrounded by forests and of large extents of
grasslands without forest are very different. The
fact that the probability of detection declines
drastically at the highest proportion of Pinus
forest may be because at lower proportions there
is not only Pinus forest but also other kinds of
vegetation, such as Abies forest, so at the high-
est proportion of Pinus forest there is no Abies
(it is almost totally Pinus forest), which is drier
than Abies forest or than a combination of both.
The ensemble model shows that the highest
percentage (83.8%) of the potential distribution
is located in the TMVB (fig. 1), mainly because
this biogeographic zone has mountains with an
altitude between 1800 and 4500 m a.s.l. and the
highest amount of Abies forest (91.1%) and Pi-
229
nus forest (29.6%) in Mexico. Within this bio-
geographic zone there is only 1346.9 km2 of
Abies forest (1.1%) and 6507.7 km2 of Pinus
forest (5.4%), which are highly fragmented by
urban settlements (3.4%). In fact, the TMVB
contains the biggest metropolitan areas of Mex-
ico, the Valle de Mexico that has 2557.4 peo-
ple per km2 and it is one of the world’s largest
metropolitan areas (CONAPO, 2010) with a
high percentage of roads (25.3%) and agricul-
tural fields (44.7%). Therefore, it is essential
to establish conservation plans in order to pre-
serve the highly endangered patches of Abies-
Pinus forest (Ponce-Reyes et al., 2012) and con-
sequently the connectivity between B. i. imbri-
cata populations in order to avoid genetic drift,
bottlenecks, inbreeding, and loss of genetic and
allelic variability, which can lead to increased
susceptibility to extinction (Frankham, Ballou
and Briscoe, 2005). We found some mountain
areas with suitable conditions for populations
of B. i. imbricata in other biogeographic zones,
such as the SMor, the SMs, the As and the DB;
however, in these provinces there are other sub-
species of B. imbricata (Canseco-Márquez et
al., 2007) which can be a biotic component lim-
iting B. i. imbricata distribution.
We also found that 39.7% of the poten-
tial subspecies distribution is located in NPAs,
where land-use change is smaller. Although in
these NPAs it is not allowed to build houses and
kill wildlife, there are communities that lived
there before the government established these
areas as protected. Moreover, local people, as
well as some visitors, believe that B. i. imbri-
cata is a venomous lizard (Fernández-Badillo,
2008) and they kill it, contributing to population
decline even in NPAs. Furthermore, the federal
government proposes tripling the area of for-
est under management, by increasing the area
of commercial forest plantations (National For-
est Program 2014-2018), so grasslands are be-
ing destroyed by roller chopping. These grass-
lands are also important to the ecosystem, as
they host a large number of reptile species such
as Barisia imbricate; Barisia herrerae; Cro-
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230
talus triseriatus, C. tlaloci, Plestiodon copei,
Sceloporus torquatus, S. grammicus S. aeneus,
Thamnophis eques, T. melanogaster, T. scalaris;
amphibians such as Hyla eximia, H. plicata,
Lithobates spectabilis; and mammals such as
the endemic rodent Neotodomon alstoni. As a
measure to avoid this problem, we propose to
implement environmental education, not only to
local communities but also to the authorities.
Acknowledgements. We thank Martha Zarco-González
and Octavio Monroy-Vilchis for the analysis advice. Also,
we would like to thank Victor Muñoz-Mora for help with
field work. We thank the editor and two anonymous re-
viewers for their comments that helped to improve the
manuscript.
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Submitted: August 27, 2016. Final revision received: Janu-
ary 31, 2017. Accepted: February 13, 2017.
Associate Editor: José Brito.
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