Research Article |
Corresponding author: Julio A. Lemos-Espinal ( lemos@unam.mx ) Academic editor: Uri García-Vázquez
© 2022 Geoffrey R. Smith, Julio A. Lemos-Espinal.
This is an open access article distributed under the terms of the Creative Commons Attribution License (CC BY 4.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited.
Citation:
Smith GR, Lemos-Espinal JA (2022) Factors related to species richness, endemism, and conservation status of the herpetofauna (Amphibia and Reptilia) of Mexican states. ZooKeys 1097: 85-101. https://doi.org/10.3897/zookeys.1097.80424
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Mexico is a megadiverse country with high endemicity in its herpetofauna. We examine how species richness, proportion of state and country endemic species, and proportion of species in a category of conservation concern using listings in the International Union for Conservation of Nature (IUCN) Red List and the Secretaría del Medio Ambiente y Recursos Naturales (SEMARNAT) in 27 of 32 Mexican states are related to environmental and human demographic and socioeconomic variables. Amphibian and reptile species richness were positively related to latitude range and number of physiographic regions and negatively related to latitude. The proportion of state endemic amphibian species in a state was negatively related to latitude whereas no variables influenced the proportion in reptiles. The proportion of country endemics in a state was positively related to human population density and the number of physiographic regions and negatively related to per capita gross domestic product (GDP) and latitude range for amphibians; it was positively related to human population density and elevation range and negatively related to latitude range for reptiles. The proportion of amphibian species in an IUCN category of concern in a state was positively related to human population density and negatively related to latitude; for reptiles, it was negatively related to human population density. The proportion of SEMARNAT-listed species in a state was positively related to human population density for both amphibians and reptiles and negatively related to latitude range for amphibians. Our analyses found that larger macroecological patterns (e.g., latitudinal species gradient, heterogeneity-richness relationships) and human population density play important roles in determining the richness and conservation status of Mexican amphibians and reptiles.
amphibians, environmental factors, human demographic factors, IUCN status, reptiles, SEMARNAT listing, socioeconomic factors
Much of Mexico consists of a transition zone (i.e., the Mexican Transition Zone) between the Nearctic and Neotropical zones (
Unfortunately, Mexico is not immune to global environmental decline. Indeed, Mexico is an area with high extinction debt and risk for amphibians and reptiles (
Human pressure on biodiversity has increased in the Neotropics (
Here we examine the distribution of amphibian and reptile species richness among 27 of 32 Mexican states. In particular, we examine relationships between species richness, proportion of state and country endemic species (i.e., the proportion of the species in a state that are state or country endemics), and proportion of a state’s amphibian or reptile species in a category of conservation concern using the Interational Union for Conservation of Nature (IUCN) Red List (i.e., vulnerable, threatened, endangered, critically endangered, near extinction), and the proportion of a state’s amphibian or reptile species listed in Secretaría del Medio Ambiente y Recursos Naturales (
We collected species lists for amphibians and reptiles of Mexican states from the available literature and updated these species lists using additional literature through November 2020 (see Suppl. material
Amphibians and reptile species richness, proportion of species in a state that are state and country endemics, proportion of species that are in an IUCN category of concern, and the proportion of species that are SEMARNAT listed for Mexican states.
State | Amphibians | Reptiles | ||||||||
---|---|---|---|---|---|---|---|---|---|---|
Species Richness | Prop. State Endemic | Prop. Country Endemic | Prop. IUCN | Prop. SEMARNAT | Species Richness | Prop. State Endemic | Prop. Country Endemic | Prop. IUCN | Prop. SEMARNAT | |
Aguascalientes | 19 | 0 | 0.68 | 0.16 | 0.16 | 65 | 0 | 0.54 | 0.05 | 0.15 |
Baja California | 17 | 0 | 0 | 0.29 | 0.06 | 103 | 0.23 | 0.32 | 0.21 | 0.23 |
Baja California Sur | 3 | 0 | 0 | 0 | 0 | 84 | 0.44 | 0.57 | 0.13 | 0.32 |
Campeche | 24 | 0 | 0.04 | 0 | 0 | 103 | 0 | 0.12 | 0.13 | 0.16 |
Chiapas | 108 | 0.10 | 0.21 | 0.53 | 0.04 | 219 | 0.06 | 0.18 | 0.11 | 0.16 |
Chihuahua | 37 | 0.03 | 0.35 | 0.11 | 0.03 | 140 | 0.01 | 0.33 | 0.06 | 0.15 |
Coahuila | 24 | 0 | 0.25 | 0.17 | 0.04 | 117 | 0.06 | 0.30 | 0.15 | 0.22 |
Colima | 39 | 0.03 | 0.68 | 0.13 | 0.08 | 117 | 0.04 | 0.66 | 0.11 | 0.18 |
Durango | 36 | 0 | 0.6 | 0.11 | 0.03 | 119 | 0.02 | 0.45 | 0.05 | 0.18 |
Guerrero | 78 | 0.32 | 0.76 | 0.45 | 0.11 | 181 | 0.15 | 0.68 | 0.08 | 0.16 |
Hidalgo | 52 | 0.08 | 0.65 | 0.46 | 0.15 | 126 | 0.01 | 0.49 | 0.10 | 0.18 |
Jalisco | 55 | 0.06 | 0.72 | 0.22 | 0.07 | 173 | 0.01 | 0.66 | 0.10 | 0.17 |
Mexico | 49 | 0.08 | 0.77 | 0.42 | 0.21 | 99 | 0.01 | 0.71 | 0.08 | 0.17 |
Mexico City | 18 | 0.11 | 0.83 | 0.5 | 0.39 | 45 | 0 | 0.76 | 0.04 | 0.22 |
Michoacán | 58 | 0.09 | 0.74 | 0.26 | 0.09 | 161 | 0 | 0.7 | 0.09 | 0.18 |
Morelos | 38 | 0 | 0.70 | 0.27 | 0.14 | 97 | 0 | 0.68 | 0.04 | 0.17 |
Nayarit | 37 | 0 | 0.6 | 0.08 | 0.03 | 117 | 0.01 | 0.60 | 0.07 | 0.14 |
Nuevo León | 25 | 0 | 0.32 | 0.2 | 0.04 | 119 | 0.02 | 0.34 | 0.09 | 0.20 |
Oaxaca | 152 | 0.39 | 0.69 | 0.60 | 0.10 | 299 | 0.11 | 0.55 | 0.11 | 0.18 |
Puebla | 92 | 0.03 | 0.7 | 0.49 | 0.16 | 176 | 0.02 | 0.61 | 0.07 | 0.14 |
Querétaro | 34 | 0.60 | 0.56 | 0.32 | 0.12 | 104 | 0.02 | 0.49 | 0.07 | 0.18 |
Quintano Roo | 23 | 0 | 0.09 | 0.04 | 0 | 108 | 0.01 | 0.12 | 0.12 | 0.16 |
San Luis Potosí | 42 | 0 | 0.41 | 0.34 | 0.12 | 138 | 0 | 0.41 | 0.09 | 0.17 |
Sinaloa | 39 | 0 | 0.55 | 0.13 | 0 | 119 | 0.01 | 0.51 | 0.10 | 0.14 |
Sonora | 36 | 0 | 0.39 | 0.11 | 0.03 | 159 | 0.10 | 0.37 | 0.13 | 0.18 |
Tamaulipas | 44 | 0.11 | 0.41 | 0.27 | 0.14 | 137 | 0.04 | 0.34 | 0.11 | 0.21 |
Yucatán | 17 | 0 | 0.12 | 0.06 | 0 | 85 | 0 | 0.12 | 0.12 | 0.18 |
Human demographic and socioeconomic variables and environmental variables for Mexican states.
State | State Area (km2)1 | Human Population (2018)1 | Human Population Density (N/km2)1 | Per capita GDP (US$)2 | Proportion Protected Territory3 | GPS Coordinates1 (°) | Elevation Range (m)1 | Number of Physiographic Regions4 |
---|---|---|---|---|---|---|---|---|
Aguascalientes | 5618 | 1.337,792 | 238.1 | 9975 | 26.0 | 22.1243, 1.0042 | 1666 | 3 |
Baja California | 71.450 | 3.633,772 | 50.9 | 9449 | 19.0 | 30.3593, 4.7186 | 3100 | 2 |
Baja California Sur | 73.909 | 832.827 | 11.3 | 11.060 | 42.0 | 25.4360, 5.1280 | 2080 | 1 |
Campeche | 57.507 | 948.459 | 16.5 | 51.460 | 39.6 | 19.6167, 0.7667 | 390 | 2 |
Chiapas | 73.311 | 5.445,233 | 74.3 | 3592 | 18.0 | 17.2588, 5.4530 | 4080 | 3 |
Chihuahua | 247.460 | 3.816,865 | 15.4 | 8833 | 8.1 | 28.6843, 6.4175 | 3050 | 2 |
Coahuila | 151.595 | 3.063,662 | 20.2 | 12.838 | 19.0 | 27.2114, 5.3372 | 3380 | 3 |
Colima | 5627 | 759.686 | 135 | 9177 | 6.6 | 19.0983, 0.8283 | 3820 | 2 |
Durango | 123.317 | 1.815,966 | 14.5 | 7888 | 22.1 | 24.5950, 4.5000 | 3240 | 4 |
Guerrero | 63.596 | 3.625,040 | 5.7 | 4586 | 0.15 | 17.6018, 2.5719 | 3550 | 2 |
Hidalgo | 20.813 | 2.980,532 | 143.2 | 6508 | 6.9 | 20.4982, 1.8008 | 3251 | 3 |
Jalisco | 78.588 | 8.197,483 | 104.3 | 9239 | 11.2 | 20.8380, 3.8244 | 4339 | 4 |
Mexico | 22.500 | 17.604,619 | 782.4 | 6199 | 43.8 | 19.3264, 1.9189 | 5268 | 2 |
Mexico City | 1495 | 8.788,141 | 5878.4 | 21.079 | 14.1 | 19.3206, 0.5444 | 1702 | 1 |
Michoacán | 58.599 | 4.687,211 | 80 | 5522 | 5.9 | 19.1547, 2.4794 | 4100 | 2 |
Morelos | 4879 | 1.987,596 | 407.4 | 6961 | 26.8 | 18.7319, 0.7994 | 4580 | 2 |
Nayarit | 27.857 | 1.290,519 | 46.3 | 6220 | 30.8 | 21.8439, 2.4811 | 2760 | 4 |
Nuevo León | 64.156 | 5.300,619 | 82.6 | 16.228 | 8.9 | 25.4810, 4.6364 | 3660 | 3 |
Oaxaca | 93.757 | 4.084,674 | 43.6 | 4446 | 7.1 | 17.1635, 3.0125 | 3720 | 5 |
Puebla | 34.306 | 6.371,381 | 185.7 | 5890 | 19.5 | 19.3500, 2.9667 | 5530 | 4 |
Querétaro | 11.699 | 2.091,823 | 178.8 | 12.502 | 33.6 | 20.8425, 1.655 | 2600 | 3 |
Quintano Roo | 50.212 | 1.709,479 | 34 | 11.381 | 32.6 | 19.7000, 3.7667 | 230 | 1 |
San Luis Potosí | 61.137 | 2.824,976 | 46.2 | 8118 | 6.6 | 22.8258, 3.3311 | 3160 | 3 |
Sinaloa | 58.328 | 3.059,322 | 52.5 | 8108 | 7.6 | 24.7547, 4.5750 | 2520 | 2 |
Sonora | 179.355 | 3.050,473 | 17 | 11.543 | 10.3 | 29.3954, 6.1969 | 2620 | 4 |
Tamaulipas | 80.249 | 3.661,162 | 45.5 | 9347 | 13.7 | 24.9430, 5.4722 | 3280 | 3 |
Yucatán | 39.524 | 2.199,618 | 55.7 | 8.011 | 25.6 | 20.5667, 2.0667 | 210 | 1 |
We used generalized linear models (Normal distribution, Identity link) for amphibians and reptiles separately to examine the relationships of the human demographic and socioeconomic variables and the geographic and climatic variables and species richness, proportion of country endemics, proportion of state endemics, proportion of species in an IUCN category of concern, and proportion SEMARNAT-listed species. We used JMP Pro 15.1 (SAS Institute, Cary, NC). for statistical analyses.
Amphibian species richness was positively related to latitude range and number of physiographic regions and negatively related to latitude (Table
Results of generalized linear models examining the relationship between human demographic and socioeconomic variables and environmental variables and species richness, proportion of species in a state that are state and country endemics, proportion of species that are in an IUCN category of concern, and the proportion of species that are SEMARNAT listed for amphibians in 27 Mexican states. Values on first line are coefficients, values on second line are P-values. Bolded entries are significant at α = 0.05.
Variable | Intercept | State area | Human population | Human population density | per capita GDP | Proportion Protected | Latitude | Latitude Range | Elevation range | Physiographic regions | Overall P |
---|---|---|---|---|---|---|---|---|---|---|---|
Species Richness | 153.03 < 0.0001 | 0.00011 0.16 | < 0.00001 0.63 | -0.0010 0.725 | -0.0002 0.18 | -44.61 0.074 | -7.37 < 0.0001 | 5.91 0.043 | 0.0025 0.39 | 10.62 0.0006 | < 0.0001 |
Prop State endemic | 0.447 0.0003 | < 0.000001 0.17 | <0.0000001 0.39 | <0.000001 0.65 | -<0.000001 0.49 | -0.247 0.057 | -0.018 0.0009 | 0.004 0.81 | -<0.00001 0.52 | 0.013 0.36 | 0.0108 |
Prop Country endemic | 0.568 0.0046 | 0.000001 0.10 | <<0.000001 0.36 | 0.000062 0.024 | -0.0000010 0.004 | -0.41 0.066 | 0.00055 0.95 | -0.116 < 0.0001 | 0.000047 0.078 | 0.069 0.008 | <0.0001 |
Prop IUCN | 0.469 0.0065 | -<0.000001 0.35 | <<0.000001 0.26 | 0.000050 0.0336 | -0.0000016 0.55 | -0.33 0.086 | -0.018 0.017 | 0.022 0.30 | 0.000029 0.20 | 0.035 0.10 | 0.0002 |
Prop SEMARNAT | 0.0045 0.93 | -0.0000001 0.56 | <<0.0000001 0.14 | 0.000050 < 0.0001 | -0.0000012 0.19 | -0.042 0.19 | 0.0034 0.18 | -0.018 0.016 | 0.000012 0.12 | 0.0096 0.19 | <0.0001 |
The relationships between amphibian species richness and a state’s latitude A latitude range B and number of physiographic regions C between the proportion of a state’s amphibian species that are state endemics and a state’s latitude D and between the proportion of a state’s amphibian species that are country endemics and the state’s human population density E per capita GDP F latitude range G and the number of physiographic regions for Mexico H.
The relationships between the proportion of a state’s amphibian species that are in an IUCN category of concern and a state’s human population density A and latitude B and between the proportion of a state’s amphibian species that are listed in
Reptile species richness was positively related to latitude range and the number of physiographic regions and negatively related to latitude and the proportion of land protected (Table
Results of generalized linear models examining the relationship between human demographic and socioeconomic variables and environmental variables and species richness, proportion of species in a state that are state and country endemics, proportion of species that are in an IUCN category of concern, and the proportion of species that are SEMARNAT listed for reptiles in 27 Mexican states. Values on first line are coefficients, values on second line are P-values. Bolded entries are significant at α = 0.05.
Variable | Intercept | State area | Human population | Human population density | per capita GDP | Proportion protected | Latitude | Latitude Range | Elevation range | Physiographic regions | Overall P |
---|---|---|---|---|---|---|---|---|---|---|---|
Species Richness | 279.65 < 0.0001 | 0.00023 0.06 | 0.0000005 0.73 | -0.0084 0.061 | 0.00026 0.62 | -109.7 0.0046 | -10.74 < 0.0001 | 13.35 0.0031 | 0.0013 0.76 | 16.57 0.0003 | <0.0001 |
Prop State endemic | -0.070 0.58 | -0.0000002 0.61 | <0.0000001 0.074 | 0.000015 0.40 | -<0.00001 0.79 | 0.272 0.072 | 0.0009 0.87 | 0.031 0.08 | 0.000025 0.17 | -0.024 0.15 | 0.22 |
Prop Country endemic | 0.245 0.18 | 0.00000035 0.61 | <0.0000001 0.39 | 0.000068 0.011 | -0.0000065 0.039 | 0.082 0.69 | 0.0094 0.26 | -0.067 0.009 | 0.000091 0.0011 | -0.0016 0.95 | 0.0002 |
Prop IUCN | 0.051 0.28 | -0.0000003 0.095 | <0.0000001 0.34 | -0.000015 0.035 | 0.0000014 0.10 | -0.064 0.24 | 0.0024 0.28 | 0.0122 0.063 | -0.000005 0.42 | -0.0060 0.34 | 0.076 |
Prop SEMARNAT | 0.079 0.087 | -0.0000002 0.23 | -<<0.00001 0.084 | 0.000015 0.028 | 0.00000025 0.74 | 0.088 0.10 | 0.0031 0.14 | 0.010 0.09 | 0.000011 0.10 | -0.010 0.083 | 0.054 |
The relationships between reptile species richness and a state’s proportion of protected land A latitude B latitude range C and number of physiographic regions D between the proportion of a state’s reptile species that are country endemics and the state’s human population density E latitude range F and elevation range G between the proportion of a state’s reptile species that are in an IUCN category of concern and a state’s human population density H and between the proportion of a state’s reptile species that are listed in
Our analyses found that species richness, endemism, and conservation status of amphibians and reptiles in Mexican states are related to both human demographic and socioeconomic variables and environmental variables. Below we discuss our observations on the factors related to species richness, endemism, and conservation status of amphibians and reptiles in Mexico.
For amphibians, species richness was positively related to latitude range and the number of physiographic regions in a state and negatively related to latitude (i.e., species richness decreased with latitude), whereas reptile species richness was positively related to latitudinal range and the number of physiographic regions and negatively related to latitude and the proportion of land protected. The proportion of a state’s amphibian species that are state endemics was negatively related to latitude whereas none of the state variables we examined were related to the proportion of a state’s reptile species that are state endemics. For the proportion of a state’s species of amphibians that are country endemics there was a positive relationship with human population density and the number of physiographic regions and a negative relationship with per capita GDP and latitude range. For reptiles, this proportion was positively related to human population density and elevation range and negatively related to latitude range.
The positive relationships between species richness and latitude range and the number of physiographic regions for both amphibians and reptiles likely result from the increased variety of habitats and climates in a state leading to a greater number of niches, which can result in increased number of species occurring in a state. Our results for the herpetofauna of Mexico are similar to studies on Sceloporus lizards (
The decrease in species richness of amphibians and reptiles with increasing latitude is consistent with the latitudinal species gradient (
In addition, for reptiles, species richness decreased with increasing human population density and the proportion of the state’s area protected. This relationship suggests there might be a negative impact of human population on species richness, perhaps due to the impact on amphibian and reptile populations. For example, in Europe, climate and human related factors explained 41–42% of variation in species richness of amphibians and reptiles (
The patterns of state endemism that we observed show limited effects of the state variables we examined, with only a negative relationship between latitude and the proportion of a state’s amphibian species being state endemics. In part this general lack of relationships may reflect the artificial nature of state boundaries (see Caveats below) such that states often share such physiographic regions or habitats and so likely share species, even over a small area along borders. For country-level endemism, we found positive relationships with human population density and measures related to habitat heterogeneity (e.g., number of physiographic regions or elevation range) for both amphibians and reptiles. Such relationships likely reflect the high level of endemism found in the Transvolcanic Mexican Belt (
Both human demographic and environmental variables affected the proportion of both amphibians and reptiles of conservation concern in a state. The proportion of amphibian species in an IUCN category of concern decreased with latitude and increased with human population density. For SEMARNAT, the proportion of listed species increased with human population density and decreased with latitudinal range. For reptiles, the proportion of a state’s species in an IUCN category of concern decreased with human population density, but the proportion that is SEMARNAT listed increased with human population density. The importance of human population density in determining conservation status is consistent with the impacts that anthropogenic effects on the environment have on amphibians and reptiles. For example, a high proportion of endangered amphibians in Mexico being found in areas that have experienced transformation to agriculture or urbanization (
We recognize that our analysis is a snapshot in time of both taxonomic knowledge and conservation status. Species lists are dynamic and changing as new species are described, new localities are found, and populations are extirpated. In addition, conservation status for species, whether at the global (i.e., IUCN) or national (i.e., SEMARNAT) scale, are frequently reassessed. Thus, we realize that our analysis represents our knowledge at the time we generated our species lists and conducted our analyses. However, obtaining a snapshot now will provide a baseline that can be monitored and evaluated as our understanding of taxonomy, species distributions, and conservation status change over time.
We also recognize that by using global and national conservation status we are not taking into account variation in conservation status of species populations in each state such that species may be doing well in some states but poorly in other states. This is a challenge, but unfortunately conservation status at the state level is known for even fewer species than for the national and global measures. In addition, many regulations are focused on, or use, national or global level assessments rather than state level assessments.
Finally, we further recognize that the use of states in our analyses does not necessarily reflect ecological or biogeographical reality (i.e., they are not natural regions). However, given the nature of governmental processes, conservation efforts are usually a function of state or political boundaries and thus we argue that understanding patterns at the state level is pragmatic.
We thank L. Wilson, J. Sigala, and an anonymous reviewer for helpful comments on the manuscript. Support for this study was provided by Dirección General de Asuntos del Personal Académico – Programa de Apoyo a Proyectos de Investigación e Innovación Tecnológica (DGAPA-PAPIIT) through Project IN202021.
Table S1
Data type: Summary of sources for data used.
Explanation note: This file contains a list of the checklists from which we obtained our species lists for each Mecixan state. It also provides information on sources used to update those checklists.
Table S2
Data type: Occurrences.
Explanation note: This file contains the data matrix used to generate the data used in our analyses. Basically it is a matrix of presence and absence of herpetofaunal species in each of the considered Mexican states. This matrix was created using the sources provided in Table S1.