Research Article |
Corresponding author: Leticia Núñez-Landa ( 1226976h@umich.mx ) Academic editor: Monika Lipińska
© 2024 Tiberio C. Monterrubio-Rico, Leticia Núñez-Landa, Juan F. Charre-Medellín.
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:
Monterrubio-Rico TC, Núñez-Landa L, Charre-Medellín JF (2024) Ocelot (Leopardus pardalis) breeding effort and productivity in seasonal tropical forests of the central Mexican Pacific. In: Lipińska M, Lopez-Selva MM, Sierra JM (Eds) Biodiversity research in Central America. Neotropical Biology and Conservation 19(2): 51-67. https://doi.org/10.3897/neotropical.19.e114194
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Camera trap studies on ocelots in Mexico have focused on abundance, habitat use, and activity patterns, mainly within Protected Areas of tropical humid forests in southern regions. However, their ecology, including breeding effort and productivity in seasonal dry Pacific forests and areas outside Protected Areas, has received less attention. The species’ low reproductive rates make their populations vulnerable to decline. In evaluating the reproduction, the species’ breeding activity was examined for two survey periods performed during 2010–2016. We estimated the percentage of females raising young in the surveys and their associated fecundity–”productivity”. The overall results revealed a population of 26 males and 46 females that included 10 females (21%) rearing 12 young, defined as kittens, cubs, or juveniles, for an effective sample area (ESA) of 200 km2. Overall, the productivity averaged 2.05/100 km2 per season. All parameters varied between the two areas, and productivity was not constant in time in either area or site. Information on species breeding activity in combination with density estimations may facilitate determining the minimum area required for a viable ocelot population in the region. Protected Areas are required in the studied region for the ocelot in order to maintain a viable local population. This is due to the increasing fragmentation caused by agriculture, cattle ranching, and forest fires, and it is hypothesized that ocelots do not tolerate disturbance or severe fragmentation.
Camera trap, felines, female generational replacement, tropical dry forests
The ocelot (Leopardus pardalis Linnaeus, 1758) is considered the most abundant feline in neotropical forests (
The ocelot is considered a key species for tropical forest dynamics, as it regulates populations at lower trophic levels through predation (
The ocelot is solitary, polygynous, and polyestrous, with male home ranges overlapping the home ranges of several females. Mating occurs any season of the year, with a higher breeding effort during the September–November period. The gestation period lasts between 78 and 82 days and occurs every two years. The first litter may occur within 18–45 months, with an average litter of one to two kittens. The observed sex ratio at birth is 50:50 (
Ocelots present low reproductive rates and slow recovery potential, as their populations cannot overcome annual reductions of ≥ 3%, making them vulnerable to reduction numbers caused by habitat degradation and human hunting (
The ocelot is listed as an endangered species in Mexico (Norma Oficial Mexicana, 2010), but the current distribution and viability of their populations are unknown, even with current studies evaluating ocelot ecology using camera traps (
Notably, estimations on ocelot breeding activity such as proportion of breeding females, productivity rates, renewal rates, and influential environmental and ecological factors have not been examined. This information is needed, considering the high forest fragmentation rates in the region, for better population assessment. Few studies in Mexico present results on breeding activity. One study in the tropical and cloud forests of Sierra Norte, Oaxaca, resulted in two females, three males, and one cub in July with an estimated population density of 7.8 individuals per 100 km2 (
Regions in Mexico where ocelot populations were studied documenting density, vegetation, annual average precipitation in mm, protection status, sex ratio, and productivity (months evidenced). RF= Rainforest, TDF= Tropical Dry Forest, STF= Semideciduous Tropical Forest, TS= Tropical Scrub.
State (Number in map) | Density /100 km2 | Vegetation (rain in mm) | Area status | M/F | Juveniles or cubs (Months recorded) | Source |
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Chiapas (1) | 12.9/5.3 | RF (3000) | Protected | 4/10 | 0 |
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Oaxaca (2) | 38.0 | RF (2500) | Unprotected | 5/3 | 0 |
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Oaxaca (3) | 7.8 | RF (2500) | Protected | 2/2 | 1 (July) |
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San Luis Potosí-Oaxaca (4) | NA | TDF (1800) | Protected | 6/2 | 2 (Nov-Dec.) | Cacelin Castillo, L. A. 2015. |
Quintana Roo (5) | 2.5/13.8 | STF (1511) | Protected | 6/3 | 0 | Avíla-Nájera et al. 2015. |
Tamaulipas (6) | 131.0 | TDF (1441) | Protected | 19/19 | 2 (NA) | Ocañas García, 2019. |
Michoacán (7) | 18.8-24.8 | TDF (1300) | Unporotected | 26/46 | 12 (Nov.-March) |
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Quintana Roo (8) | 13.98 | STF (1200) | Protected | 5/6 | 0 |
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Jalisco (9) | NA | TDF (1150) | Protected | 0 | 1 cub (Jul/Ag.) | Aranda et al. 2012. |
San Luis Potosí (10) | NA | TDF (1017) | Protected | 10/1 | 0 |
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Tamaulipas (11) | 17.5- 59 | TDF (1000) | Protected | 23/24 | NA |
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Sonora (12) | 0.63-0.65 | TS (400) | Unprotected | 16/8 | 0 |
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Although some information is available on ocelot densities (
The long-term conservation of ocelots resides in maintaining viable populations in their original ranges, identifying areas with high densities and adequate productivity-recruitment levels to function as “source populations” for Protected Area designation (
In the absence of countrywide population assessments and productivity data and considering the broad environmental variation in the species distribution (
The study was conducted in two localities from the Sierra Madre del Sur and Jalisco dry forests (coastal lowlands) ecoregions (
Triangles indicate study regions of camera trap ocelot studies in Mexico, and circles correspond to the areas with evidenced breeding. Numbering corresponds to Table
The data came from two survey designs (2010–2011; 2014–2016) over five years. Sampling in each year was concentrated during the dry season and after the rainy season, with a total sampling of 27 months. Data were analyzed in three-month periods considering the closed population assumption in capture-recapture studies (
The first survey included three sampling seasons: March to June 2010; November 2010 to February 2011; and March to May 2011; involving 10 to 15 camera stations (a site where one or two cameras were placed), with an Effective Sampled Area (ESA) averaging 119 km2 in the community of El Naranjal, Aquila. Detailed descriptions of the localities and specific sampling efforts by locality are available in
In both surveys, the camera-trap stations were placed between 1 and 3 km apart. Camera traps were revised every 30 days to replace batteries and memory cards. To ensure that only the natural movements of the animals were registered, no baits or olfactory attractants were used (
Ocelots were individually identified using their unique combination of rosettes, spots, stripes, scars, relative size (adult, juvenile, cub-kitten), sex (
We define the breeding effort parameter as the percentage of females evidenced rearing young (kittens or juveniles). Breeding efforts were estimated for each of the two surveys and for the overall study. In the absence of precise demographic analysis (cohort, life table, or productivity table analysis) for ocelots, we defined ocelot productivity as analogous to fecundity. Both terms constitute the same parameter in reproduction, but the term fecundity is also used for humans, whereas productivity is acceptable in wildlife (
Ocelot productivity in this study was considered as the number of kittens/juveniles per reproductive female, defined as the females observed in the presence of offspring. The per capita rate (
We examined breeding activity and productivity in combination with the sex ratio and densities obtained in previous studies (
Density models used in the previous studies (
The cumulative survey effort analyzed was 11,356, averaging 1,622 camera traps/night of effort per sampling season (interval: 844–2,151 camera nights). Individual camera efforts averaged 85.6 ± 26.3 SD. The total number of ocelot records was 436 (186 from the first survey and 250 from the second survey). The overall capture rate was 4.3, in an interval from 2.4 to 6.8 ocelot records/100 camera-trap nights. Only 350 records were adequate for individual identification.
The total number of ocelots identified by combining the two surveys was 26 males and 46 females. Ten females (21%) were accompanied by 12 young (kittens, cubs, or juveniles) in an effective sample area (ESA) of 200 km2. Breeding activity and productivity were registered every year but not for all the sampled seasons (Tables
Ocelot population parameters in the central Pacific seasonal dry forests, (* when the statistic correspond to an average instead of a subtotal.
Surveyed season | First survey El Naranjal | Sub total/average* | Second survey San Jose de los Pinos | Sub total/average* | ||||||
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Feb-Jun 2010 | Nov-2010-Feb 2011 | Mar-May 2011 | Feb 2010-Jun 2011 | Mar-May 2014 | Nov 2014-Feb 2015 | Mar-May 2015 | Nov 2015-Feb 2016 | Nov 2014-Feb 2016 | Total/ overall+ | |
Survey effort | 1,134 | 1,188 | 1,458 | 3,780 | 2,151 | 2,752 | 844 | 1,829 | 7,576 | 11,356 |
Average effort by camera | 70.8 | 59.4 | 50.2 | 60.1* | 86 | 105.8 | 105.5 | 121.9 | 104.8* | 85.6+ |
Ocelot records | 66 | 56 | 64 | 186 | 69 | 65 | 57 | 59 | 250 | 436 |
Ocelot records for id | 52 | 47 | 64 | 163 | 41 | 49 | 43 | 54 | 187 | 350 |
Capture rate | 5.8 | 4.7 | 4.4 | 5* | 3.2 | 2.4 | 6.8 | 3.2 | 3.9* | 4.3+ |
Male/ female records | 21/31 | 12/36 | 10/54 | 43/121 | 16/25 | 22/27 | 11/32 | 32/20 | 81/104 | 124/225 |
Identified males/females | 6/9 | 7/16 | 4/11 | 17/36 | 7/16 | 8/12 | 5/10 | 9/21 | 29/50 | 46/86 |
New-unique males/females | 6/9 | 5/8 | 2/2 | 13/19 | 7/16 | 3/6 | 0/2 | 3/3 | 13/27 | 26/46 |
Females rearing young (cub-juvenile) | 3 | 2 | 1 | 6 | 0 | 1 | 0 | 3 | 4 | 10 |
Percentage of females rearing young | 33% | 25% | 0.9% | 15.4* | 0 | 8.3% | 0 | 25% | 16.5%* | 21.7+ |
No. cubs-juvenile | 4 | 2 | 1 | 7 | 0 | 1 | 0 | 4 | 5 | 12 |
No. female cub-juvenile (50%) | 2 | 1 | 0.5 | 3.5 | 0 | 0.5 | 0 | 2 | 2.5 | 6 |
Percentage of female young raised (female recruitment) | 0.22 | 0.125 | 0.045 | 0.13* | 0 | 0.04 | 0 | 0.09 | 0.03* | 0.07* |
Productivity, and densities (
First survey | Second survey | ||||||||
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Feb-Jun 2010 | Nov-2010-Feb 2011 | Mar-May 2011 | Average-subtotal* | Mar-May 2014 | Nov. 2014-Feb 2015 | Mar.-May 2015 | Nov 2015-Feb 2016 | Average- subtotal* | |
Average number of records for identified female | 3.4 | 3.75 | 4.9 | 4.0 | 2.27 | 2.16 | 3.2 | 1.6 | 2.3 |
Productivity by breeding female | 1.3 | 1.0 | 1.0 | 1.16 | 0 | 1.0 | 0 | 2.0 | 1.25 |
Female per capita productivity | 0.44 | .125 | .09 | .218 | 0 | 0.08 | 0 | 0.33 | 0.102 |
Population density | 13.8 | 28.0 | 14.8 | 18.8 | 22.0 | 22.0 | 27.8 | 27.6 | 24.8 |
Estimated female density | 9.5 | 19.4 | 10.9 | 13.2 | 15.3 | 13.2 | 16.8 | 18.9 | 16.05 |
Density of females rearing young | 2.6 | 4.7 | 1.0 | 2.7 | 0 | 1.1 | 0 | 2.7 | 1.9 |
Young productivity /100km2 | 4.18 | 2.3 | 1.0 | 2.49 | 0 | 1.0 | 0 | 5.9 | 1.72 |
Female young density/100km2 | 2.09 | 1.15 | 0.05 | 1.24 | 0 | 0.50 | 0 | 2.9 | 0.86 |
Two juvenile ocelots were observed accompanied by an adult female in December and January, respectively, whereas the kittens/cubs captured (N= 5) and recaptured (N= 14) occurred in a relatively even distribution with 3–4 records from December to March.
The percentage of females in the sample (the two surveys) accompanied by cubs or juveniles was 21.7% (n = 10; N 46), varying the percentage among seasons from 8.3 to 33%. During March–May 2014 and 2015, no cubs, kittens, or juveniles were registered. In the first survey, 13 male and 19 female ocelots were identified, including six females (31%) rearing seven young. During the second survey, the ocelots identified included 13 males and 27 females, with four females (14.8%) rearing five young (Table
The overall productivity estimation was 1.25 young per active breeding female, and the per capita rate was 0.26 for the study (12 young/46 females) (Table
Female density ranged from 13.2 to 16.5 2 ind./100km2 as adjusted by the observed sex ratio (
The information generated in this analysis complements the knowledge of the ocelot population ecology for tropical dry forests and constitutes a basis for examining the viability of populations and related Protected Area requirements in the Pacific. The number of females rearing young in a population is a key influencing factor for recovery or viable populations in the wild (
Most ocelot studies in general do not analyze breeding effort and productivity rates with camera traps; probably due to the limited number of identified ocelots and/or the design of the study (survey period’s length and timing, etc.). Although the breeding effort is obtainable from camera trap data, it requires a large sample size of ocelots to obtain results with ecological significance (Appendix
Before this analysis, the highest number of ocelots surveyed in Mexico corresponded to 51 ocelots, 19 males, 19 females, and two cubs for the protected wetlands of Tamaulipas (
Breeding effort in ocelot populations has been documented for humid tropical forests, varying from 10 to 26% (
The productivity observed was low and varied among survey periods and between areas; for instance, there was a slow potential population renewal. The ocelot population in this region exhibited a broad interval in productivity rates across years and between areas, from a high female annual per capita rate of 0.109 female cubs (10 females/100 adults) to as low as 0.034 (3.4 females/100 adults) female cubs. Considering the rate for the overall study, that constitutes a 0.13 for five years, and annually, a 0.026 rate implies a production of 5.6 female cubs for a hypothetical population of 100 adult female ocelots for a complete female renewal in 18.7 years. In contrast, the low rates of the second survey (0.034) would require 33 years for total female replacement. In comparison, annual recruitment rates for the ocelot populations of Belize ranged between 0.09 and 0.12 ocelots, for a female annual recruitment of 10 ocelots in a hypothetical population of 100 females (
Variation may be explained as a response to environmental variation and ecological interactions. Severe seasonal droughts may increase ocelot dispersal, hunger, and mortality caused by prey scarcity and predation by larger felines as local and regional habitat quality is affected (
The presence of cougars and jaguars and prey abundance influenced the presence and abundance levels of ocelot males and females differently in the region, as female ocelot activity was negatively correlated with higher activity of large felines such as cougars and jaguars, whereas males were more associated with prey abundance (
In general, it has been recognized that factors influencing ocelot abundance may be classified as environmental, ecological, and human impacts (Massara, Paschoal et al. 2018). For the seasonally dry forests in Michoacán, we hypothesize that an operating mechanism regulating the per capita recruitment level in the population is the percentage of local females breeding during a specific period, which fluctuates from reproduction suppression to 33% of females rearing cubs (Table
No conclusive evidence of intraspecific competition was mentioned before, but in our data, indirect evidence is noticed by examining the breeding effort levels occurring at different density levels of adults (Table
The ocelot in the region is also sympatric with cougars (Puma concolor) and, to a lesser degree, jaguars (Panthera onca), which ecologically constitute competitors and predators (
Notably, direct comparison between the reproductive parameters documented between both surveys is limited since they do not correspond to a balanced design, and some local factors such as waterhole availability may be influential. Water is a scarce resource in the region, limited during the 7-month dry season (
Although the studied region presents few human settlements and density, as mentioned earlier, the low and variable per capita productivity of the ocelot population increases its vulnerability to hunting by local villagers, which occurs in retribution for domestic cattle losses (personal observation). The poaching of one ocelot by each human settlement in the region may represent the total or a substantial proportion of the annual productivity in 100 km2. Poaching control, habitat restoration, and population monitoring are required in the region to preserve all the feline populations and other endangered species (
It is fundamental to the continuation of the monitoring effort for productivity in the region. Ideally, balanced designs measure the variation, consistency, and frequency of the results. Additional surveys in the landscape should extend into the Coahuayana and western Aquila municipalities to increase the sample of reproductive parameters, especially in areas of the region near Jalisco State. In Michoacan, the ocelot is also present in the Balsas basin, but the survey in the region that included the Zicuiran-Infiernillo Biosphere Reserve (ZIBR) revealed low abundance (
The forest composition and topography of the studied region present similarities to the neighboring southern Guerrero and Oaxaca states along the Pacific slope mountain ranges, as these constitute the southern extension of the same ecoregion, the Sierra Madre del Sur (
This study provides the first estimation of ocelot population fecundity and productivity for the western seasonal dry tropical forests. The population parameters vary seasonally and between surveyed areas. The potential population renewal scenarios are low, confirming the species’ vulnerability if drought, additional habitat loss, and poaching negatively affect the population size. The long-term viability of the species is uncertain, as drought and increasing forest fires experienced during the latest years are reducing habitat availability and quality, in addition to the ongoing human-caused fragmentation.
We appreciate the assistance from a wide range of people and institutions. TCMR appreciates the continuous financial support provided by the Coordinación de Investigación Cientifica at UMSNH. JFCM thanks CONAHCYT for the postdoctoral fellowship awarded. LNL thanks the master’s scholarship 1106517 awarded by CONAHCYT. We appreciate the field support given by students of Lab Vertebrate Priority Terrestrial and Faculty Biology for the facilities granted and UMSNH for the preparation of the manuscript. We appreciate the helpful comments of anonymous reviewers.
The authors have declared that no competing interests exist.
During the study, ocelots were not captured, harmed or stressed, as all data was
obtained by camera trapp surveys.
This study was funded by the Coordination of Scientific Research of the Universidad Michoacana de San Nicolas de Hidalgo and the master’s scholarship 1106517 awarded by CONAHCYT to Núñez-Landa.
Conceptualization: TCMR. Data curation: LNL. Formal analysis: TCMR. Funding acquisition: TCMR. Investigation: JFCM, LNL, TCMR. Methodology: LNL, TCMR. Resources: TCMR. Supervision: JFCM, TCMR. Writing - original draft: TCMR. Writing - review and editing: JFCM, TCMR.
Tiberio C. Monterrubio-Rico https://orcid.org/0000-0001-5786-9696
Leticia Núñez-Landa https://orcid.org/0009-0002-2639-6253
Juan F. Charre-Medellín https://orcid.org/0000-0002-2192-1680
All of the data that support the findings of this study are available in the main text.
Characteristics of 35 ocelot population studies using camera traps. RF= Rainforest, TDF= Tropical Dry Forest, STF= Semideciduous Tropical Forest, TS= Tropical Scrub.
Study and Country | Vegetation (precipitation mm)- | Area status | No. camera stations | Trapping effort (camera-trap/days) | Individuals identified | Density | Juveniles and cubs | Source |
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1 Ecuador | RF (3200) | Protected | 26 | 2340 | 35-36 | 31-34 | – | Salvador and Espinoza 2015. |
2 Peru | RF (3000) | Protected | 23 | 983-2085 | 22-27 | 49-64 | 2 | Kolowski and Alonso 2010. |
3 Brasil | RF (2373) | Protected | 50 | 7020 | 19 | 12-25 | – | Gómez da Rocha et al. 2016. |
3 Brasil | RF (2373) | Protected | 50 | 7020 | 30 | 27-40 | – | Gómez da Rocha et al. 2016. |
3 Brasil | RF (2373) | Protected | 50 | 7020 | 17 | 19-28 | – | Gómez da Rocha et al. 2016. |
4 Colombia | RF (2236) | Protected | 21 | 1283 | 6 | 5-11 | – | Diaz-Pulido and Payán-Garrido 2011. |
5 Panama | RF (6000) | Protected | 14 | 490 | 11 | 22-93 | – |
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6 Costa Rica | RF (4000) | Protected | 10 | 600 | 5 | 8-11 | – | González-Maya and Cardenal-Porras 2011. |
7 Bolivia | RF (2440) | Protected | 69 | 1018 | 17 | 40-66 | – | Ayala et al. 2010. |
8 Brasil | STF (1800) | Protected | 52 | 504 | 9 | 9-30 | – | Trolle and Kelly 2005 |
9 Mexico | RF (3000) | Protected | 29 | 1920 | 14 | 12.9 | – |
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10 Belize | RF (1500) | Protected | 7-19 | 239-1577 | 4-19 | 25.85 | – | Dillon and Kelly 2007. |
11 Belize | RF (2000) | Protected | 7-17 | 238-1513 | 9 | 8-26 | – | Dillon and Kelly 2008. |
12 Guatemala | STF (1350) | Protected | 33 | 1455 | 38 | 10 | – | Moreira et al. 2007. |
13 Mexico | RF (2500) | Unprotected | 29 | 8529 | 9 | 22-38 | – |
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14 Guatemala | RF (1700) | Protected | 25 | 1150 | 17 | 10.83 | – |
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15 Mexico | STF (1200) | Protected | 27 | 2160 | 12 | 13.98 | – |
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16 Bolivia | TDF (1200) | Protected | 16-31 | 960-2280 | 18-42 | 24-66 | – | Maffei et al. 2005. |
17 Brasil | STF (1300) | Protected | 30 | 450 | 9 | 62.11 | 1 | Trolle and Kéry 2003. |
18 Mexico | TDF (1300) | Unprotected | 9-9 | 2378-2270 | 6-11 | 23.7 | – |
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19 Mexico | STF (1511) | Protected | 22-27 | 1936-1755 | 9-10 | 2-26 | – | Avíla-Nájera et al. 2015. |
20 Mexico | TDF (1017) | Protected | 34-45 | 7786 | 15 | 4-46 | – |
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21 Brasil | STF (3014) | Protected | 14-19 | 585 | 6 | 21 | – | Fusco-Costa et al. 2010. |
22 Argentina | RF (2200) | Protected | 34-39 | 1409-1631 | 17-34 | 7-20 | 2 |
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23 USA | TS (680) | Protected | 18 | 658 | 3 | 14.8 | A pregnant female |
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24 Argentina | RF (1919) | Protected | 42-47 | 1871-2059 | 10-33 | 4-9 | – |
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25 México | RF (2500) | Protected | 44 | 12800 | 5 | 7.82 | 1 |
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26 México | TDF (1441) | Protected | 34 | 5168 | 51 | 131.2 | 2 | Ocañas García, 2019. |
27 Guatemala | RF (2252) | Protected | 40 | 784 | 8 | - | – | Hermes, 2004. |
28 México | TDF (1800) | Protected | 63 | 2381 | 13 | - | 2 | Cacelin Castillo, L. A. 2015. |
29 Panamá | RF (2600) | Protected | - | 1824 | - | 163-181 | 4 | Rodgers 2014. |
30 Belize | RF (3000) | Protected | 20-50 | 29137 | - | 6-22.5 | – |
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31 Belize | RF (3000) | Protected | 20-50 | 74854 | 51 | 6-14 | – |
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32 Ecuador | TDF (2800) | Protected | 22-36 | 7000-1700 | 45 | 102-149 | – | Mosquera 2011. |
33 Colombia | TDF (2176) | Protected | 9 | 360 | 2 | 1.5-1.8 | – | Garrote et al. 2019. |
34 Colombia | TDF (-) | Unprotected | 41 | 1502 | 2 | - | – | Valderrama-Vásquez 2013. |
35 México | TS (400) | Unprotected | 270 | 88508 | 33 | 1.53 | – |
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