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The Impact of Global Warming on the Range Distribution of Different Climatic Groups of Aspidoscelis costata costata

Author(s): Martha Anahí Güizado-Rodríguez , Claudia Ballesteros-Barrera , Gustavo Casas-Andreu , Victor Luis Barradas-Miranda , Oswaldo Téllez-Valdés and Isaías Hazarmabeth Salgado-Ugarte Source: Zoological Science, 29(12):834-843. 2012. Published By: Zoological Society of Japan DOI: http://dx.doi.org/10.2108/zsj.29.834 URL: http://www.bioone.org/doi/full/10.2108/zsj.29.834

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unam. Ext. with potential upward shifts toward higher elevation in response to climate warming. 2010). We used the maximum entropy species distribution model (MaxEnt) to project the potential distributions expected for the years 2020. Universidad Nacional Autónoma de México. This warming has induced biological and ecological responses in animals and plants.. The Earth has experienced climatic perturbations promoting an environmental temperature increase of 0. and .. The aims of our study were (1) to understand whether predicted warmer climatic conditions affect the geographic potential distribution of different climatic groups of Aspidoscelis costata costata and (2) to identify potential altitudinal changes of these groups under global warming. life history traits (Chammaillé-Jammes et al. Instituto de Biología. Consistent responses are apparent in phenology (McCarty.F. 09230 1 The ectothermic nature of reptiles makes them especially sensitive to global warming. 04510 2 Departamento de Biología. Mexico is considered a megadiverse country. Different climatic groups were revealed in our analysis that subsequently showed heterogeneous responses to climatic change illustrating the complex nature of species geographic responses to environmental change and the importance of modeling climatic or geographic groups and/or populations instead of the entire species’ range treated as a homogeneous entity. Tlalnepantla. Avenida de los Barrios 1. 04510 4 Laboratorio de Recursos Naturales Unidad de Biología. 2004). Sinervo et al. Universidad Nacional Autónoma de México. 2050.gob. Universidad Autónoma Metropolitana Iztapalapa. 2006. Key words: Aspidoscelis costata costata.ibiologia. Fax : +52(55)55-50-01-64.74°C over the last 100 years (IPCC. Batalla 5 de mayo s/n esq. Universidad Nacional Autónoma de México. 2000. most of these studies are carried out without making a distinction between populations. Avenida San Rafael Atlixco #186 Colonia Vicentina. Instituto de Ecología. and global climate change (Gibbons et al. unsustainable use.834 larly in terms of reptilian diversity as it is home to 804 (9.F. particu* Corresponding author.F. altitudinal distribution INTRODUCTION The geographic distribution of animals and plants is determined by ecological and historical factors (Begon et al. Ejército de Oriente. Oswaldo Téllez-Valdés4. 2001. environmental pollution. 2010). 2003). Delegación Iztapalapa. México D. Hawkings and Porter.29. E-mail: anahigr@ibunam2. Universidad Nacional Autónoma de México. However. and Isaías Hazarmabeth Salgado-Ugarte2. Víctor Luis Barradas-Miranda3. Tecnología y Prototipos. although even today this issue is relatively unexplored (Sinervo et al. AP 70-153.5 Departamento de Zoología. disease and parasitism. Circuito Exterior s/n.. MaxEnt. Facultad de Estudios Superiores Iztacala. climatic groups.mx doi:10.F. Estado de México.2108/zsj.mx). Here we present the first study of an Aspidoscelis species that evaluates the effects of global warming on its distribution using ecological niche modeling. Among the factors of greater threat to Mexican reptilian biodiversity is global warming. Parmesan. 47820. introduced invasive species. 1996.ZOOLOGICAL SCIENCE 29: 834–843 (2012) ¤ 2012 Zoological Society of Japan The Impact of Global Warming on the Range Distribution of Different Climatic Groups of Aspidoscelis costata costata Martha Anahí Güizado-Rodríguez1*.P 55-35. 2009). AP 70-153. global warming. México D. Circuito Exterior s/n. Los Reyes Iztacala. more than 50% of these reptilian species are considered to be at some level of risk by Mexican Official Norm (NOM-059-SEMARNAT2010 see www. Although climate change and its implications are a frequent topic of detailed studies. 2007). : +52(55)56-22-91-47. 09340 3 Departamento de Ecología Funcional.76%) of the 8. México D. Fuerte de Loreto Col. Lepetz et al. A.. Facultad de Estudios Superiores Zaragoza Campus II. Tel.ine. and 2080 under a single simulated climatic scenario. A growing body of data indicates that reptilian populations have seen elevated declines worldwide mainly due to habitat loss and degradation.. México D. 2006). Our analysis suggests that some climatic groups of Aspidoscelis costata costata will exhibit reductions and in others expansions in their distribution. Gustavo Casas-Andreu1.238 described species worldwide (FloresVillela and Canseco-Márquez.. Claudia Ballesteros-Barrera2. 54090 5 Laboratorio de Biometría y Biología Pesquera.

c. MaxEnt calculates the distribution of maximum entropy for a species within a given area based on observed climate values at locations where the species has been reported. 2006. 2009). Therefore. A. Foden et al.3. c.biogeomancer. 2010) to investigate how different environmental variables co-varied in order to reduce variable redundancy in the model (Beaumont et al. 2008). 2003). c.Lizard Distribution and Global Warming 835 geographic distributions (Blaustein et al. http://edcdaac. 2004. and modeled their projected distributions under our chosen climatic scenario.. we aimed to explain the potential impacts of climate change by modeling potential distributions and the altitudinal variation of different climatic groups of A. on the basis of its observed presence and (sometimes) absence at different localities. The Aspidoscelis costata costata locality records constitute all that is known about its distribution. Finally. 2003.. We used the maximum entropy species distribution model (Philips et al. nigrigularis (Reeder et al. Climate variable selection Distribution models were generated using different environmental surfaces: 1) topographic data including aspect (i. We generated a bioclimatic profile that was classified by the hierarchical association of the PATN program. 2006. The files produced were mapped as raster grids of 1 km2 of spatial resolution to produce ARCINFO/ . S.worldclim. 1986). slope (i. the soil capacity for water retention) (U.. c. 2006) as implemented in the software MaxEnt version 3. The effects of these climatic changes on the genus Aspidoscelis remain unexplored even though it is among the most diverse lizard groups inhabiting Mexico. or is likely to occur in other areas in which it has not yet been detected at present. 2005). Maslin and Walker.. Selection of climatic groups We choose climatic groups (CG) using a Bioclim extension in the GIS (Geographic Information System) Arc View 3. Aspidoscelis costata belongs to the sexlineata group.e. the amount of inclination of a surface to the horizontal plane). A.. Therefore we assigned 50% of our data for training and 50% for testing to generate potential distribution maps for those CG with more than 50 records of point localities.2 (ESRI) based on the principle that geographic limits can be predicted for a species according to its bioclimatic preferences (Nix. org) and the Global Gazetteer V2. This niche modeling has been used in several studies to predict species distributions (Zonneveld et al.. 2004) at range margins both in latitude (Hickling et al. However. 2005). c. Shoo and Williams. 2006).1 (freely available at http://www.. zweifeli. 2005).. 2006). Raxworthy et al. A. Aspidoscelis costata costata (Cope. 2005. Essentially. costata. we thought that the impact of global warming is in progress and the sampling period of locality records that we used to forecast current potential distribution is according to the period of time that was used to model construction as current climate.edu/ wschapire/maxent/). c. These climate data are derived from meteorological monthly temperature and rainfall values from 1950 to 2000 and represent biologically meaningful variables for characterizing species distributions (Nix. A. In the case of CG with less than 50 points..usgs.. Parmesan. With the previous analysis. 2006). In this paper. it is a technique used to estimate actual or potential areas of distribution. griseocephalus.. http://www. mazatlanensis and A. improvement of model performance takes place at 20 data points. 2010).. or sets of favorable habitats for a given species. because these percentages are related to the number of species records. costata across its full distributional range. climate change scenarios show there to be a tendency to dryness with a temperature increase of 2°C and a decrease of 10% in precipitation due to global warming (Villers-Ruiz and Trejo-Vázquez. MATERIALS AND METHODS Data sources Species records consist of individual point-locality information that is presence-only data obtained from fieldwork and 24 national and international museum collections (see Acknowledgments) and some related references (Maslin and Walker. Walker.gov/gtopo30/hydro/). which takes into account the limiting effects of biological interactions (Hutchinson. we verified whether the CG were climatologically different. There is no formal procedure to assign a certain percentage to training and testing data set.. and topographic index (i. 2002). 2001. RodríguezRomero et al. Wang et al. c. The sampling period of the locality records was from 1937 to 2007. As a result it is exposed to broad array of selection pressures. which is the most diverse clade within the genus Aspidoscelis.. 2010). 2004). making it a suitable model for testing a population-dependent response due to global warming. Modeling method There is a suite of methods to model the ecological niche of a species (Elith et al. we obtained a cluster of localities by contrasting climates that allowed us to identify different CG.e. and even in depth for marine fishes (Perry et al. This species is comprised of eight subspecies: A. 1965. huico.e. We then performed a discriminant function analysis and obtained Mahalanobis distances between groups to validate the non-random occurrence of the constituted groups and to test for similarity between them using the Statistica 9 Software (StatSoft. 1962. using the Gower Index and the UPGMA association option. According to Stockwell and Peterson (2002). Geological Survey. Walker. 1986). testing the hypothesis that climate warming is causing species declines is challenging given that many interacting processes operate at different spatial scales and the responses are clearly species and population-dependent (Araújo et al. 1877). We performed principal component analysis (PCA) using the Statistica 9 Software (StatSoft. 1957). we 1) compared current and future potential distributions to assess whether suitable climate space is projected to increase or decrease with climate change. this species is likely to have been distributed in other areas in which it was not detected in the past.. 2003. and 2) evaluated the potential altitudinal variation under a simulated climatic scenario. In Mexico. When necessary. which represent 158 different distributional points for A. 2007). occidentalis. 1965. 2010. Weinsheimer et al. the focal species of our study. However. c. Specifically. A. c. costata.. The data set consisted of 4565 records.. Most studies addressing this issue focus only on a portion of the species range and fewer still analyze the response of more than one population (Chamaillé-Jammes et al. A. we assigned 100% of point localities to the training data set. the direction to which a mountain slope faces). The Worldclim data layers included 11 temperature and eight precipitation metrics representing spatial variation in annual means and extremes. BioGeoMancer (http://classic..cs. The niche modeling approach outlines the potential ecological distributions onto Hutchinson’s fundamental niche concept as opposed to the realized niche space (Soberón and Peterson. and 2) environmental data obtained from Worldclim (Hijmans et al. org) at a spatial resolution of ~1 km2. jointly with digital maps (GIS raster layers) of environmental variables (Soberón and Nakamura. 2009. The output model is a cumulative probability of occurrence expressed as a percentage (Phillips et al. 2003).. RodríguezRomero et al. c.princeton. 2005. 2006) and altitude (Gian-Reto et al. is an endemic lizard of Mexico with a wide altitudinal range of 528–2468 m above sea level (Duellman and Zweifel.2 were used for georeferencing. barrancorum.

Salgado-Ugarte et al.5 imply discrimination that is no better than random (Elith et al. we converted each grid of every current and future potential distribution map into a point. AUC values greater than 0.. and frequency or proportion of the sample due to each component—from the angles and intercepts described by the negatively sloped line segments. these differences are plotted and the components are selected by eye. P < 0. CG-C inhabits in the highest altitude in contrast CG-A and CG-B. which were edited with the GIS ArcView 3.. 1 2π exp ⎜ − ⎛ z2 ⎞ ⎟ ⎝ 2⎠ Z= ( x − Xi) h Optimal bandwidth was chosen based on Silverman’s rule (Silverman. This single plot structure makes comparisons of distributional factors of several variables much easier (Hintze and Nelson. Scott. to reveal the structure found within the data (Hintze and Nelson.0 (Stata Corporation. 1986.2 C for 2100 y (IPCC. costata (Wilki’s Lambda = 0. Violin plots include the median of the data and a box indicating the interquartile range. ASCII/GRID files. F = 21. costata due to global warming. Third.2 (ESRI). (1994. Güizado-Rodríguez et al. It is based on strengthening regional cultural identities. models were classified into binary (1 or 0) or presenceabsence model. Table 1). 2050 and 2080 from the CGCM2SRES model under the A2 emission scenario which is based on projections produced by the Canadian Climate Centre (http://www. 1993). i..01° (~1 km2) (Flato et al. This contribution intends to introduce an alternative way to analyze one of the consistent responses of animals and plants in the face of global warming. 1). an estimated frequency polygon is generated by adding the estimates of each of the Gaussian components (Salgado-Ugarte et al. In order to create a binary map of presence/absence of the species we used the minimum cumulative value of training sample points as a threshold value (Phillips et al. According with PCA results. First.7 are considered to be potentially significant.37 m (range: 1101–2927 m). This scenario predicts an environmental temperature increase of 3. 2006). f (x) = density estimation for variable x n = inspection number . while scores of 0. it could be challenging to live under these climate conditions. Salgado-Ugarte. it starts with a box plot and then it adds a rotated kernel density plot to each side of the box plot. Next.e. therefore they already live close to their optimal body temperature.org/sres/gcm_data. This area has a cold and dry climate. c.. Altitudinal variation To evaluate the potential altitudinal variation of each CG. There are four steps in Bhattacharya’s method.349 ⎠ ⎥ ⎦ Given that altitudinal data distributions were multimodal. Bhattacharya develops formulas for estimating the parameters of each component—the mean. A violin plot is a combination of a box plot and a kernel density plot. standard deviation. we made a non-parametric analysis of variance Kruskal-Wallis with a post-hoc Z test to compare the medians of the Gaussian components to determine differences in altitude between CG of A. We next used a violin plot as an exploratory tool of the altitudinal data distribution. The synergistic combination of the box plot and the density trace allows much of the information from each to be displayed in one plot. Salgado-Ugarte et al. 2005). Using the cut-off value. high population growth. 2005) for the statistical package Stata software 9. h = bandwidth Xi = altitude value of each grid point K = Kernel function A kernel Gaussian function was used: K(z) = Where.836 M. 1998).e. By themselves. Finally. Specifically. each map was overlaid onto an elevation digital model with a resolution of ~1 km2 to obtain current and future altitudinal grid values. 1995. moving to higher altitudes. Future climate data was derived from GCM (General Circulation Models) projections for the years 2020. Future potential distribution We used the same topographic and climatic data that were chosen for modeling current potential distribution to project future potential distributions. 2000). the logarithmic differences of successive frequencies are calculated. 1997): h= Where.01° × 0. mainly based on different climatic preferences (Fig. and C). For Aspidoscelis. This statistical method was proposed by Rosenblatt (1956) and is defined as: Where. RESULTS Selection of climatic groups With the classification of the bioclimatic profile we found three groups (A.. 2004). After that.0 to 5. we used a kernel density estimator (KDE) for exploring and analyzing altitudinal data distributions of CG. 40°C.. B. The A2 scenario (liberal model) is described as a very heterogeneous world.0001. 1986. 2006).240. we determined the dominant Gaussian components of each CG with Bhattacharya’s semi-graphical method (1967) by using the programs presented by Salgado-Ugarte et al. The mean altitude is 2291 ± 396. 2005).9 A n 1/ 5 A = min ⎢⎜ ⎡⎛ Σ( Xi − x )2 ⎞1/ 2 ⎛ Dispertion ⎞ ⎤ ⎟⎥ ⎟. Moreover this scenario has been used in other studies investigating the effect of climate change on lizard distributions (Ballesteros-Barrera et al. ⎜ ⎢ ⎣⎝ n − 1 ⎠ ⎝ 1. We chose the A2 emission scenario because it is one of the most extreme climate scenarios to predict a high increase in environmental temperature. c. 1992. Vitt and Pianka.ipcc-data. 2007). box plots provide succinct summaries of data. Optimal bandwidth was chosen based on Silverman’s rule (Silverman. the line segment corresponding to each segment is estimated. density traces reveal important information about the distribution of data. 2007). The discriminant function analysis and the Mahalanobis distances corroborated with these CG across the geographical range of A. 0 .html) at a spatial resolution of 0. Modeling method The number of variables selected for modeling the impact of climate changes on CG distributions was reduced from the original 22 variables (19 bioclimatic and three topographic data) to 14 variables (Table 2). Second. as in standard box plots. Once we obtained the dominant Gaussian components. 1994. 1998).0457.. Individually. This step is the graphical part of the method. A.. and less concern for rapid economic development. AUC ranges from 0 to 1 where a score of 1 indicates perfect discrimination generally. This genus has one of the highest body temperatures among reptiles (i. We used the receiver operating characteristic curve (ROC) to examine model performance calculating the area under the receiver operating curve (AUC) which measure the ability of a model to discriminate between sites where a species is present versus those where it is absent.

costata as a consequence of the climate changes represented by the CGCM2SRES A2 sce- .4°C). points of CG-A and 38 of CG-B to construct an ecological niche model.1 A 1. df = 13. The mean altitude of the habitat of CG-A is of 1362. slope faces). but in CG-C.1 38.46 B 14. CG-B inhabits a wetter climate.31 1.143 p-levels the rest were used to test the model. 11.6°C) as distinguished from CG-B and CG-C. [18] precipitation of warmest quarter. Eigen values PCA 1 PCA 2 PCA 3 PCA 4 9. The CG-A population lives in the warmest zone with an environmental temperature of 23°C (15. [9] mean temperature of driest quarter. 5.95 10. 1.53 m (range: 1993–621 m).74 A 14.78 ± 288.9 implying potentially significant results: CG-A (AUC = 0. 9 12. [5] max temperobserved in the zero omission error ature of warmest month.93 0 43. and the mean annual precipitation is of 890 mm (range: 578–1294 mm). [6] min temperature of coldest month.75 m (range 113–2484 m).990). Mahalanobis Distances between climatic groups of Aspidoscelis costata costata.03 ± 466.2–26. third.84 63. Results generated for each C 41.81 2. This CG occupies a region with a lowest mean altitude: 1217. and aspect (direction to which a mountain pared to known distribution. we assigned 100% Group Group Group of the point localities for the training A 0 5. PCA 1–4 refers to the contribution of the first. quarter. Climatic Climatic Climatic A B C A B C A B C On the other hand. 10. [10] mean temperature of warmest quarter. second. c. and Squared Mahalanobis Distances F-values. 6. Circles represent Climatic Table 3 we described a brief summary of group A.992) and CG-C (AUC = explaining the cumulative percentage variation in climate variables on Aspidoscelis costata 0.63 Selected variables 1.86 81. 16 18 aspect Future potential distribution Our modeling using MaxEnt projected potential shifts in the distribution areas of A. The selected variables are: [1] annual mean temperature. respectively. The mean annual precipitation is of 1006 mm (range: 696– 1442 mm) and the mean annual temperature is of 22°C (16. [16] precipitation of wetnot show high overprediction. the scenario is very different. [8] mean temperature of wettest registered in the models for three CG. Points indicate species records that were gathered from national and the other groups (personal observation).75E-20 0 records. reproductive performance and behavior of lizards: for CG-A and CG-B the daily activity period is from 0800 h to 1800 h and the reproductive period is extended across seven months.78 73.Lizard Distribution and Global Warming 837 the annual mean temperature is around 15°C and the mean annual precipitation is 700 mm (range: 390–1052 mm) unlike CGA and CG-B. [13] precipitation of wettest obtained for the current distribution do month. In international museum collections and some related references. These temperature and precipitation differentiations affected daily activity periods. 14.65 38. Finally. rectangles correspond to Climatic group B and triangles represent Climatic PCA results showing the difference in cligroup C.6–27. CG-B Table 2.84 20.77 Cumulative Percentage 42. costata had AUC values > 0.65 36. Climatic groups of Aspidoscelis costata costata selected based on different clitive period is shorter (four months) than in matic preferences.78 Percentage 42. [15] precipitation seasonality.75E-20 0 data set for CG-C as it had only 16 B 5. and the reproducFig. which inhabit warmer and wetter climates. This predictive ability is costata distribution. as comtest quarter. [14] precipitation of driest month.85 4.992). 15.74 43. mate preference among the three CG populations. 8. [12] annual precipitation. [11] as well as the fact that the areas mean temperature of coldest quarter. the daily activity period is from 1200 h to 1600 h.55 C 0 0 CG of A.93 41. c. 13.55 B 1. CG-A and CG-B had 66 and 76 records.07 7.46 0 C 36. We used 33 Table 1. and four PCA axes in (AUC = 0.

2–13.7 ± 2. A.4 17.2 1006. Potential distributions models of Aspidoscelis costata costata for liberal climate change scenario (SRES A2).7 ± 10.838 M.4 ± 2.3 ± 7.3 ± 1. .4 ± 39.9 ± 2.1 24–13.7 15132 ± 11639.3 26. Colors indicate the following: red = areas preserved in the future. (B) 2050.3–10.5 ± 3.8 26.7 ± 186.1–12. Mean and standard deviation values of the main bioclimatic parameters selected from the PCA analysis that were the most important variables predictors of Aspidoscelis costata costata habitat distribution.4 13.8 Bio14 = Precipitation of Driest Month (mm) 3.2 586–166 244.9 Bio13 = Precipitation of Wettest Month (mm) 194.2 3.9 ± 178.6–4.3 ± 2.3 ± 2.5 216.1 ± 4.7 ± 48.4 22.4 Bio16 = Precipitation of Wettest Quarter (mm) 528. blue = distributional areas lost.4 ± 2.6 335–145 27.9 ± 77.2 95–69 389. and (C) 2080.7 197–76 7.0 Bio12 = Annual Precipitation (mm) 890.8 ± 8.2 35948– –1 Climatic Group C 15.6 21.1 233.3 11–2. Climatic group C for the years: (G) 2020.1 ± 23.2 23.5 Bio11 = Mean Temperature of Coldest Quarter (°C) 20.5–18. Table 3.1 726.4 26–18.4–16.4 Bio18 = Precipitation of Warmest Quarter (mm) 272.6–17. Climatic group A for the years: (A) 2020.2 Bio10 = Mean Temperature of Warmest Quarter (°C) 24.1 23.6 ± 111.2 29.3 19.1 15769 ± 7041 34662–7469 Bio1 = Annual Mean Temperature (°C) 22.3 618.2 22–4 84.7 16.4 24.1 ± 2.6 25.7 ± 20.8 103.4 Bio15 = Precipitation Seasonality (Coefficient of variation) 102.4 ± 2. and green = new predicted distributional areas in the future.5 ± 2.2 ± 0.2–16.3 22.2–19.2 1442–696 333–149 10–1 113–91 926–422 378–55 54–15 16.9 27.9 ± 77.8 1052–390 138.7 27.5 ± 2.1 ± 2.7 ± 118.1–4.9 ± 33.2 ± 2.01 Bio6 = Minimum Temperature of Coldest Month (°C) 23.3 ± 7057 Fig.1 22.4 25.0 ± 1.3 22.4 83–17 4.4–19.8 33343– –1 Climatic Group B 27.2–13.2 ± 2.4 ± 15.8 Bio19 = Precipitation of Coldest Quarter (mm) 26.6–14 12.3 12.4–16.6 30.6–15.6 Bio8 = Mean Temperature of Wettest Quarter (°C) 23.8 Bio5 = Maximum Temperature of Warmest Month (°C) 10.4 ± 2.0 ± 2. Bioclimatic parameters Mean ± SD Max-Min Mean ± SD 23.1 ± 112.1 ± 2.9 23.2 ± 1. 2.5 20.2–14.1 ± 4.9 1294–578 268–150 11–1 112–87 715–302 361–187 93–12 14.2 Bio9 = Mean Temperature of Driest Quarter (°C) 21.4–13.1 ± 2.6 Max-Min Mean ± SD Max-Min Climatic Group A 26.7 28.4 28.5 ± 2. Climatic group B for the years: (D) 2020. (E) 2050 and (F) 2080.6–10.5 Aspect = Direction to which a mountain slope faces (meters) 19272.8 ± 2. Güizado-Rodríguez et al. (H) 2050 and (I) 2080.

00 .00 163.00 0.05 1 2.075 km2 by 2080.00 Current Potential Distribution 2020 223.63 m (Fig. 3. 2E. 2020 C 2050 2080 4.00 223.00 155.93 0.00 0. 2G.00 6.000) (Table 4).289 km2) by 2008 (Fig.39 0. respectively) (Fig.00 194. c.525 km2) of their potential current range and it could gain only 3% (3. the kernel density estimators (EDK). N = 188694.42 0. CG-B (H3 = 37452.28 0.05 1 0.00 1.33 1 3. lines across middles of boxes are medians.24 0. Violin plots that show altitudinal data distributions for climatic groups of Aspidoscelis costata costata: (A) Climatic Group A. Fox.01 0. In our view the frequency of occurrence of individuals at certain altitudes reflects their tolerance limits.91 0.04 1 0.00 6. P = 0.06 0.00 4. (B) Climatic Group B and (C) Climatic Group C. which can resolve some of the problems of the traditional histogram (specially the origin and discontinuity) (Silverman. 2H and 2I). Results of the multiple comparison post-hoc Z test for the three climatic groups of Aspidoscelis costata costata.00 0. and 2C).39 0.63 0.49 0.38 0.000) and CG-C (H3 = 122590.65 0. 332 km2. whereas for CG-C the expansion was of 215. interquartile range is the difference between quartiles. which will be reduced to 254.Lizard Distribution and Global Warming 839 nario in the years 2020.00 3. 2050. while the line at each side of the box plot is the kernel density plot.48 0.65 0. P = 0.00 155. The picture looks even more severe for CG-C since the range will be reduced by 52% (from 113. which are the indirect relations of histograms with bio-ecological processes.04 1 2080 6. which means that these reptiles may be able to move to higher altitudes. 1990).119 km2) of its area by the same year (Fig.55 0.55 0. One approach to resolve such Table 4. N = 199257. The results of the Gaussian components showed changes in the mean altitude of the three CG (Fig. Ends of boxes are the 25th and 75th quartiles. N = 81237.63 0.923 km2. 1986. For CG-B. 2).899 km2 to 54. We used the most efficient methods to study the frequency.28 162. 2B. conserving more than 80% of its modeled range (Fig. P = 0.93 283.49 0.00 194. that is.06 167.41. and 2080 (Fig. 4). but it could gain more than 31% (50.00 163.63 0. the forecast current potential distributional area was 158.00 0.24 167.6. costata show a tendency toward an upward shift in response to climate warming toward 2080. The Kruskal-Wallis analysis showed that these differences were statistically significant: CG-A (H3 = 49980.000).38 300. The results of the violin plots were supported by the extract of dominant Gaussian components of altitudinal data determined by the Bhattacharya’s method.00 178.55 0. 3A and 3B. 3C).00 0. For CG-A and CG-B.00 178.01 162.42 0. Climatic Group Current Potential Distribution 2020 A 2050 2080 Current Potential Distribution 2020 B 2050 2080 Current Potential Distribution Fig. Numbers in bold show those periods of time that are different. the environmental niche showed an increase of more than 100 m in mean altitude toward 2080 (115. For CG-A the current potential distributional area was 315.51 and 107. which will be reduced by 47% in year 2080. 2A. respectively). The resulting density estimators also exhibit complex distribution. 2D. and 2F). Altitudinal variation The violin plots indicate that the environmental niches of the three CG of A.55 0.9 m.4.91 0.33 1 2.11 1 1.00 283.63 0.11 1 0.48 0.00 300.00 2050 6.

Numbers in the right corner of each figure show the mean and the standard deviation (x ± SD) of the dominant Gaussian component. from those mixed distributions we extracted the component with the high-density (frequency) value considered as a dominant component. c. As a first step.840 M. warming in low latitudes and altitudes will be relatively small in magnitude. 4. DISCUSSION We provide here the first comprehensive study on the potential effect of global warming on the geographic distribution of a teiid lizard. The results of this study shows the potential distribution area of three CG of A. Güizado-Rodríguez et al. 2008). However. 2) Climatic Group B and 3) Climatic Group C of Aspidoscelis costata costata. (C) 2050 and (D) 2080.. A. and future altitudinal potential distribution for (B) 2020. as organisms are currently living very close to their optimal temperature (Deutsch et al. According to the literature. species at higher latitudes and altitudes have broader thermal tolerance and are living in climates that are currently cooler than their physi- Fig. (A) Current altitudinal distribution. in these regions. warming will likely have the most deleterious consequences. Graphics indicating dominant Gaussian components estimated for altitudinal data for: 1) Climatic Group A. complex frequency behavior is to consider the distribution as a mix of individual components. costata will experience in some cases significant reductions and in others important expansions by the year 2080 due to global warming. In contrast. This component contains an important fraction of the total frequency distribution and may be used as an indicator of the relation with ecological and biological characteristics of the species involved. The line with circles represents the observed frequencies and the lines with dots correspond to the dominant Gaussian component. .

our results showed an altitudinal shift among CG of A. At present Aspidoscelis species have some problems that need to be overcome if they want to reach higher altitudes. ecological habitats need also to be considered. 2006). they do not give any glimpse of multimodality. the Illinois Natural History Survey (UIMNH). Peterson et al.. 2004). In this sense. 1992) and plants (Hansen et al. 40°C. But according with our results. In other respects. We thank the California Academy of Science Herpetology Collection Catalog (CAS).. 2006). CG-A and CG-B inhabit a region dominated by tropical dry forest. therefore it is likely to disappear due to actual global warming (Villers-Ruiz and Trejo-Vázquez. 1971) to avoid the negative effects of high body temperature during gestation. ACKNOWLEDGMENTS We thank the Instituto de Biología and Posgrado en Ciencias Biológicas of Universidad Nacional Autónoma de México. the Los Angeles County Museum of Natural History (LACM).. UNAM (CNAR-IBH). 2002). their construction is only based on five values (and some derivative ones): median. we used a kernel density estimator. c. c. which is considered to be among the best distribution descriptors available. the resulting density estimators exhibit complex distribution. This group inhabits pine forest remnants. c. which helped us to determine the structure found within altitudinal data. The components of these violin plots comprise a box plot and a kernel density plot (Hintze and Nelson. the distribution of the environmental requirements of the ecological niche for the presence of A. Besides. which means that this lizard will not have problems to reach higher altitudes. c. 2008). 1998. which agrees with the results found in species of insects (Parmesan et al.e. 1999. 1999). The second challenge that A. and Escuela Nacional de Ciencias Biológicas del Instituto Politécnico Nacional (ENCBIP) for species records. Special thanks to Constantino González Salazar for his help during the development of this work. a major portion of embryonic development occurs out at the nesting site making this species especially vulnerable to global warming. As a result. University of Kansas Biodiversity Research Center Herpetology Collection (UK). this type of vegetation could expand its distributional area to higher altitudes in response to global warming. We also thank Consejo Nacional de Ciencia y Tecnología for graduate study scholarships to MAGR (CONACYT no. costata must confront involves female reproductive physiology. and biotic interactions directly influence energy gain and fitness much work remains to be done to identify the magnitude and consequences in A. This article is a requisite for MAGR to obtain a Ph. MaxEnt forecasted a loss of more than 50% of its current potential distribution. 2004). The distribution had several regions of point concentration (several levels) which means that coexist several (simpler) individual components (mixed distribution). What is usually overlooked in these modeling efforts is the fact that species can be composed of different populations or geographical groups across their ranges.D. not a simple Gaussian (normal) distribution or an asymmetric (biased) one. costata. University of California. Christiansen. University of Colorado Museum (UCM). UNAM (MZFC). and extremes and in spite of may give an adequate indication of skewness and outliers. the same authors explain that this ecosystem has had the most deleterious damage due to anthropogenic activities. The CG-C population has the bleakest prognosis. Sistema Nacional de Información Biológica-CONABIO (SNIB). since these CG inhabit one of the most degraded and populated regions of Mexico. 2000. Carnegie Museum of Natural History (CM). American Museum of Natural History (AMNH). Because the impact of warming will be both site. as it contains an important fraction (and often the main part) of the information regarding the behavior of the studied phenomenon (Salgado-Ugarte et al. These modifications can induce alterations in community structure and in the relative abundance of species. 1994). Los Angeles (MVZ). 1998). University of Arizona Museum of Natural History (UAZ). Texas Cooperative Wildlife Collection-Herpetology Collection (TCWC). We must be able to accurately predict range shifts in order to mitigate impacts on biodiversity and propose appropriate conservation strategies. indicating that warming may even enhance their fitness (Addo-Bediako et al. costata will expand to higher altitudes due to global warming. Deutsch et al. Museum of Natural Science (LSU). c. 2001).. 1966. In the present study. .. Vitt and Pianka. Museum of Southwestern Biology of the University of New Mexico (MSB). Michigan State University Museum (MSUM). This vegetation type does not tolerate a rise in environmental temperature. According to Villers-Ruiz and Trejo-Vázquez (2004). Field Museum of Natural History (FMNH). Museo de Zoología de la Facultad de Ciencias. as is the case of the distributions obtained in this work. birds (Thomas and Lennon. quartiles.. The first one is to attain and maintain a high body temperature (i. National Museum of Natural History Smithsonian Institution (USNM). Therefore. Beyond latitude and altitude considerations.. as well as increases in habitat fragmentation and population extinction (Davis et al. mammals (Hersteinsson and MacDonald. as they are improved histograms (Silverman. For this reason. Ghalambor et al. costata persistence due to global warming.. which have the potential to respond independently to changes in climate (Araújo et al. the synergetic effect of habitat destruction with global warming could be more devastating for CG-A and CG-B of A. c. 165072).Lizard Distribution and Global Warming 841 ological optima. degree within the Posgrado en Ciencias Biológicas of Universidad Nacional Autónoma de México. Brigham Young University (BYU). Arizona State University (ASU). 1986). University of Michigan Museum of Zoology (UMMZ). we developed a method to evaluate the potential altitudinal distribution under a climate change scenario for A. costata. Although the box and whiskers diagrams are very useful graphs. Therefore in order to understand the complex distribution of altitudinal data we identified the dominant component (as a single Gaussian distribution). Colección Nacional de Anfibios y Reptiles del Instituto de Biología. and a gain of only 3% by year 2080. Collection of Herpetology. costata. British Museum of Natural History (BMNH). As an exploratory tool we used a violin plot. Females of this genus typically keep their eggs inside oviducts for short periods of time from seven to 20 days (McCoy and Hoddenbach. as the environmental requirements needed for survival today will be present at higher altitudes in the future. As a result. However.. our results draw attention to the importance of modeling climatic or geographic groups and/or populations instead of the entire species’ range treated as a homogeneous entity.and species-specific.

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