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Population Fluctuations of Mourning Doves (Zenaida macroura) in the State of Jalisco, Mexico, During 2004-2017.


The Mourning Dove (Zenaida macrcnira) is one of the most abundant birds in North America, with estimates of about 475 million individuals during the 1980s (Tomlinson el al., 1994) and 279 million more recently in 2016 (Seamans, 2017). It is also the most hunted gamebird in the U.S. (Seamans, 2017), as well as in northern and central Mexico (Sadler, 1993). Mourning Doves breed from central Canada to southern Mexico and winter throughout much of their breeding range, but those in the most northern areas migrate southwards to winter mostly in the southern United States, Mexico, and Central America (Aldrich, 1993). This is reflected in the recovery of 13% of Mourning Doves banded in the Central Management Unit of the United States, 6% of those from the Western Management Unit, and 0.2% of those from the Eastern Management Unit, in Mexico (Tomlinson, 1993; see also Channing, 1979). These banded individuals were recovered mostly in the states of Jalisco, Michoacan, Guanajuato, Colima, and Guerrero (Tomlinson, 1993). As a result of the winter migration, the population in northern and central Mexico is increased considerably during those times (Leopold, 1959). The state of Jalisco, which represents 4.1% of the country's area, received 15% of all Mourning Dove hunting permits issued for the nation during the mid-2000s (Secretaria del Medio Ambiente y Recursos Naturales, 2008).

From 1966 to 1993, the U.S. Western and Central Mourning Dove Management Units experienced population reductions (Peteijohn et al, 1994; Seamans, 2017; Tomlison et al, 1994), and it has been speculated similar reductions may also have occurred in Mexico (Silvy and Lopez, 2014). However, this cannot be determined as Mexico lacks systematic Mourning Dove surveys. Not only are surveys nonexistent in Mexico, but overall there is ver) little information readily available on the species in the literature (Baskett et al., 1993; Tacha and Braun, 1994; Direccion General de Vida Silvestre, 2006; Otis et al, 2008; Rocha Gutierrez et al, 2009; Silvy and Lopez, 2014). After Leopold (1959) little new data have been generated on the Mourning Dove in Mexico, highlighting the need for better monitoring of this important species (Case, 2008; Otis et al., 2008).

Our aim was to obtain a more systematic understanding of Mourning Dove distribution and inter-annual population variation in the state of Jalisco. As far as we know, this is the first such long-term study performed in Mexico. Our specific objectives were to determine:

(1) whether Mourning Dove nvtmbers change between the start (November) and end (March) of the winter season, in Jalisco; (2) what the differences in Mourning Dove abundance may be between the different biogeographic zones of the state of Jalisco, and (3) whether there were any population trends between the 2004/05 and 2016/17 winter seasons.



The state of Jalisco is part of the transition between Nearctic and Neotropical regions, which, along with a topographic range from sea level to 4260 m (asi), creates a large array of different environments and habitats, from tropical forests to conifer forests (Instituto Nacional de Estadistica, Geografia e Informatica, 2002). This diverse array of habitats can be grouped into five biogeographic regions (Comision Nacional para el Conocimiento y Uso de la Biodiversidad, 1997): the Sierra Madre Occidental, Southern Highlands, Volcanic Belt, Pacific Coast, and Balsas Depression (Fig. 1).

The Sierra Madre (Occidental) region in Jalisco, above 2000 m at sea level (asl), is composed of mountain slopes and peaks but also of mesas and canyons. Its climate is temperate semiarid, with an annual rainfall of 750 mm and a temperature range of 4--23 C. Its vegetation varies from xerophytic habitats and grasslands to pine oak woodlands and low deciduous forests.

The southern highlands, at >1900 m asl in the northeastern part of the state, are the southernmost extent of Mexico's Central Plateau. It has a temperate dry climate, with an annual rainfall of 650 mm, temperatures between 4 and 25 C, and contains grasslands and xerophytic shrublands.

The Volcanic Belt, mostly at 1200-1800 m asl, is the western part of the central Mexican belt of volcanoes and includes the central valleys of Jalisco as well as the Lake of Chapala area. Its climate is temperate subhumid, with an annual rainfall of 950 mm and temperatures between 6 and 25 C. Its vegetation consisted of grasslands, xeric shrublands, low deciduous forests, and pine oak woodlands but is now heavily transformed to farmland.

The Pacific coast region is formed by the coastal lowlands < 400 m asl, between the highlands and the sea. It has semidry and subhumid climates, with an annual rainfall of 850 mm, minimum temperature of 16 C and maximum of 32 C, and has a vegetation composed mostly of tropical deciduous and subdeciduous forests.

The Balsas Depression, an East West drainage south of the Volcanic Belt limited to the south by the Sierra Madre del Sur, has only a small portion of it in Jalisco, in which it occupies also but a small portion of the state. Because of its small size, this region was not included in our study.


In November of 2004, Mourning Doves surveys began along eight routes, two in each of the four major biogeographic units in the state; two additional routes were added in February 2008 (Table 1). The survey routes differed in altitude and natural vegetation in agreement with the biogeographic region in which they occurred (Table 1). Each route was surveyed during the last week of October or first of November (autumn) and between the second week of February and first week of March (late winter) of each wintering season, although logistic limitations occasionally prevented the realization of all surveys (Table 2).

Survey routes had the same layout as the call count routes used in the U.S. They were composed of 20 survey stations at 1.6 km intervals along little traveled dirt or gravel roads that were drivable at survey times. Routes were separated from each other by at least 50 km, and those within a biogeographic region were surveyed on consecutive days or, at most, with one day in between.

As surveys were to be earned out during the nonbreeding season, when doves do not vocalize (see Otis el al., 2008), call counts were not feasible and a specific procedure had to be developed. To this end exploratory surveys were carried out in 2003. In these surveys doves were counted for 6 d, from 0600 until 1900 h, simultaneously, at six locations in the Southern Highlands, and their number, species, distance, flight height, and grouping were recorded (Rafael de J. Hernandez-Garcia, unpubl. data). These data were used to design the procedure that was followed throughout the remainder of the study. It consisted of counts of individuals flying during a 3 min period will)in a 200 m radius at each survey station. Surveys began at sunrise and continued for 1:57 h (travel time between stations was 3 min), but if wind exceeded 20 km/h, it rained, or risibility was reduced by fog counts were postponed until conditions improved (following Sayre et al., 1978). Each route was surveyed by a two-person team, one person counting and one recording. Before beginning the formal surveys in 2004, the observers were trained for standardization. Throughout the study routes were surveyed primarily by the same teams since the beginning.


Mourning Doves count data were scrutinized visually through route by route scatterplots before statistical analysis. Late winter counts did not seem different from autumn counts on a route by route basis. A correlation analysis, using the 111 cases in which there were counts for both dates, proved this to be the case (P [less than or equal to] 0.001, [r.sup.2] = 0.41), and a paired t-test indicated the means of the two survey periods were not different. Hence, autumn and late winter counts for each wintering season were averaged for all subsequent analyses. In the few cases in which only one count had been performed, this was used in place of the average. Mourning Doves were nearly absent from the Pacific Coast routes; therefore, these were not considered further.

We explored whether the estimated abundance in the U.S. Central and Western Management Units and rainfall from January until October of the survey year explained the Mourning Dove counts in Jalisco using an information-theoretic modeling approach (Burnham and Anderson, 2002). All possible models relating the Jalisco counts to (a) the summer population estimates for the Central and Western Management Units and (b) rainfall were constructed and the Akaike Information Criterion for small samples (AICc) was used to select the best model (Burnham and Anderson, 2002). U.S. Mourning Dove abundance was obtained from Seamans (2017) while rainfall data were from the El Laurel (Calvillo, Aguascalientes) weather station, the only station in our general area of interest from which data are publicly available covering the entire span of our study. As weather is governed by large scale phenomena, inter-year differences in rainfall for the entire region would be reflected at any station in the region.

The former analysis was complemented with visual scrutiny of a graph that included the Jalisco counts and the U.S. estimates, which suggested the changes in the U.S. estimates of the previous calendar year could provide a better explanation for the changes in our counts than changes in the estimates of the summer immediately prior. Therefore, year to year changes in our data from the 2006/2007 to the 2014/2015 winter seasons (natural logarithm of a year's count--natural logarithm of previous year's count; Boonstra and Krebs, 2012), were modelled in the same way as above against: (a) the changes in the U.S. estimates two summers prior (15 mo before; using the difference between natural logarithms) and (b) the rainfall from January until October of the survey year. We excluded the 2003 U.S. estimates, because not all U.S. states participated, and, being at the onset of the national Mourning Dove banding program, surveys were more prone to errors due to inexperience of the surveyors. We were unable to control for the effect of observer ability. All statistical analysis were performed using the pgirmess package (developed by P. Giraudoux) in R (R Development Core Team, 2016)


In total 250 individual surveys were carried out, 124 in autumn and 126 in late winter. During these surveys 13,866 Mourning Doves were recorded. The Sierra Madre region supported the highest densities of Mourning Doves (118 doves /route /date on average), followed by the Southern Highlands (64), and the Volcanic Belt (60); whereas, the Pacific Coast routes had only two doves /survey. Mourning Dove counts in late winter were correlated significantly with counts from the previous autumn, and both were not significantly different. Additionally, Mourning Dove counts exhibited an overall decrease from the 2004/2005 to the 2014/2015 nonbreeding season (Fig. 2).

Despite differences in individual routes, there were general trends, particularly accentuated during the first half of the study (Fig. 2). In modelling our counts, neither rainfall nor the U.S. estimates, either from the summer immediately prior or from 1 y earlier, were included in the best models. However, the season to season changes exhibited by our data were explained by the changes in the U.S. estimates 15 mo earlier, and the model that best explained the counts of all our routes together, included the estimates of both the Central and the Western Management Units, although the models with only the central or only the western units included were also just as good (Table 3).

The changes in Mourning Dove counts in the three biographic units analyzed were explained differently by changes in the U.S. management units' population estimates (Table 3). The model best explaining the changes in the Sierra Madre region included only the changes in the Western Management Unit. Those in the Southern Highlands were best explained by both the Western and the Central Management Units. However, by the rule of parsimony, the one including only the Western Unit is to be considered better (sensu Burnham and Anderson, 2002). The changes in the Volcanic Belt region were best explained by the model including only the Central Management Unit. Rainfall was not a component of any best model.


The two routes with tropical habitats clearly had much lower numbers of Mourning Doves than those in highland habitats, and this conforms to known patterns of the species' distribution (Leopold, 1959). In the highlands (Volcanic Belt, Sierra Madre, and Southern Highlands), there were some routes with many doves, and others with few, but numbers were not related to any particular biogeographic region (Fig. 2).

Contrary to the band returns in the late 1960s and early 1970s, which were mostly from the Central Management Unit (Tomlinson, 1993), our data exhibited a relationship with the population changes in both the Central and Western Management Units. This might suggest the reduction in the U.S. central dove populations has caused a decrease in the population in Jalisco, and, at the same time, a larger influence of the population from the western U.S. The later, probably facilitated by changes in the agricultural patterns of northwestern (Sonora, Sinaloa) and western (Nayarit) Mexico, which might have led to improved connectivity between Jalisco and the Western Management Unit. Lack of data prevents further discussion of this issue. Interestingly, the variables that fit our count data best are the U.S. population changes two summers (15 mo) before. Again, the paucity of additional information prevents the exploration of potential ecological explanations of this, and this relationship must await further research to be confirmed and explained, or rejected.

Wintering Mourning Doves in our survey routes experienced a decrease of >60% from 2004/2005 to 2014/2015. This is much larger than the overall reduction of 14% reported for the species in North America (Partners in Flight, 2017). The large difference between the reductions in the U.S. and in our counts suggests that Mourning Dove abundance in Jalisco, in addition to the reduction in the U.S. central management unit's population, is apparently driven by other factors, either at the local, landscape, or at a much larger regional or extra-regional scale. These factors cannot currently be explored for lack of Mourning Dove data from other areas in Mexico. Notwithstanding this, the population reduction in our study should raise concerns among those involved with the species' management and points to the need for surveys to be continued and broadened until a better overall picture of the dove's population trends and the factors that drive them are obtained.

In conclusion, our data on Mourning Dove numbers in Jalisco during the nonbreeding season indicate: (1) the counts recorded might reflect changes in the western U.S. and central U.S., albeit with a 1 y delay, and this should be the subject of further research; and (2) the numbers have decreased significantly between the autumn of 2004 and the 2014/ 2015 winter season, in part due to population decreases in the central U.S., but also due to local, landscape, regional, or extra-regional processes that are yet to be understood. This decrease in Mourning Dove numbers should raise a warning flag for those involved in the species' management and points to the urgency of further surveys.

Acknowledgments.--Javier Ochoa Covarrubias, Ricardo Millan, Cristina Ascendo, Araceli Valverde, and Rafael Zermeno assisted during field work. Jaime Luevano provided logistic support during the preparation of this article and Patrick Oiraudoux provided advice on analysis. Sula Vanderplank and three anonymous reviewers provided important comments to improve this article substantially. We thank them all for their kindness and support. Funding was provided by lhe Mexican government. This work was prepared while Kric Mellink was on sabbatical at the Institut Universitaire de France, in Besancon, supported by the Consejo National de Ciencia y Tecnologia (Mexico).


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RAFAEL DE JESUS HERNANDEZ-GARCIA (1) Departamento de Cienrias Ambientales, CUCBA, t 'niversidad de Citada/ajara. Camino Ramon Padilla Sanchez #2100 45110 Nextipac, Zapopan, Jalisco


ERIC MELLINK (2) Departamento de Biologia de ta Conservacion, Cientro de Investigacion Cientifica y de Educacion Superior de Ensenado. Carretera Ensenada-Tijuana So. 3918, Zona Plfiyitas, 22860 Ensenada, Baja California, Mexico

(1) E-mail:

(2) Corresponding author: e-mail:

Caption: FIG. 1.--Biogeographic regions of Jalisco, Mexico, and Mourning Dove survey routes. 2004-2017. Survey routes: 1- La Laguna, 2- San Juan, 3- Pegueros, 4- Villa Hidalgo, 5- Teocaltiche, 6- Ameca, 7Jocotepec, 8- Amatitan, 9- La Gloria, and 10- La Huerta

Caption: FIG. 2.--Average counts of wintering Mourning Doves along eight survey routes in the state of Jalisco, Mexico, from 2004 to 2017, and Mourning Dove population estimates in the Central and Western Management Units, U.S., from 2004 to 2017 the summer immediately before our survey season and two summers (i.e., 15 mo) prior. Counts are indicated as kite graphs, in which the width of a kite indicates the average count in any particular season. A scale bar = 100 individuals is inserted between Pegueros and Amatitan on the right side. Autumn and winter counts were significantly correlated (P [less than or equal to] 0.001, [r.sup.2] = 0.41) and not significantly different (paired t-test). Average Mourning Dove abundance on survey routes was positively related with the percentage of farmland along lhe route (P < 0.01), and their overall numbers decreased from the 2004-2005 lo the 2014-2015 season (P < 0.01)
TABLE 1.--Mourning Dove survey routes in the state of Jalisco,
Mexico, by biogeographic region, their altitude, and number
of stations per vegetation type. 2004-2017

                                   Coodinates                Elevation
  Region/route                  (lag N, long W)               (m asl)

Sierra Madre
  San Juan            21.6668[degrees], 103.5372[degrees]    2000-2250
  La Laguna           22.1127[degrees], 103.3162[degrees]    2000-2250
Southern Highlands
  Teocaltiche         21.4515[degrees], 102.5723[degrees]    1772-2343
  Villa Hidalgo       21.5659[degrees], 102.4157[degrees]    2000-2200
  Pegueros            20.8619[degrees], 102.40304[degrees]   1950-2250
Volcanic Belt
  Amatitan            20.8403[degrees], 103.7342[degrees]     684-1876
  Jocotepec           20.3014[degrees],103.4681[degrees]     1600-1750
  Ameca               20.5319[degrees], 104.0392[degrees]    1350-1950
Pacific Coast
  La Gloria           19.9610[degrees], 105.4372[degrees]      <100
  La Huerta           19.4945[degrees], 104.6724[degrees]     250-400

                           Medium        caducifolius
                      subcaducifolius     forest and    Oak-pine
  Region/route        troppical forest    shrublands    woodland

Sierra Madre
  San Juan                   --               --           7
  La Laguna                  --               --           --
Southern Highlands
  Teocaltiche                                 5            5
  Villa Hidalgo              --               5            --
  Pegueros                   --               4            --
Volcanic Belt
  Amatitan                                    8
  Jocotepec                  --               5            --
  Ameca                      --               3            4
Pacific Coast
  La Gloria                  2                --           --
  La Huerta                  8                --           --

                      or induced
  Region/route        grassland    Farmland   Village

Sierra Madre
  San Juan                3           9          1
  La Laguna               --          18         2
Southern Highlands
  Teocaltiche             2           8         --
  Villa Hidalgo           1           11         3
  Pegueros                8           8         --
Volcanic Belt
  Amatitan                1           8          1
  Jocotepec               --          15        --
  Ameca                   2           10         1
Pacific Coast
  La Gloria               8           10        --
  La Huerta               12          --        --

TABLE 2.--Surveys along ten different routes to document
Mourning Dove abundance in the state of Jalisco, Mexico.
2004-2017. A = autumn (last week of October to first week
of November), W= late winter (second week of February to first
week of March). An "x" indicates that a survey was carried out.
The two routes on the Pacific Coast were almost devoid of
Mourning Doves and were not included in the analysis

                                2004-   2005-   2006-   2007-
                                2005    2006    2007    2008

Region          Route           A   W   A   W   A   W   A   W

Sierra Madre    San Juan        x   x   x   x   x   x   x   x
                La Laguna       x   x   x   x   x   x   x   x
Southern        Teocaltiche                                 x
  Highlands     Villa Hidalgo   x   x   x   x   x   x   x   x
                Pegueros        x   x   x   x   x   x   x   x
Volcanic Belt   Amatitlan                                   x
                Jocotepec       x   x   x   x   x   x   x   x
                Ameca           x   x   x   x   x   x   x   x
Pacific Coast   La Gloria       x   x   x   x   x   x   x   x
                La Huerta       x   x   x   x   x   x   x   x

                                2008-   2009-   2010-   2011-
                                2009    2010    2011    2012

Region          Route           A   W   A   W   A   W   A   W

Sierra Madre    San Juan        x   x   x   x   x   x   x   x
                La Laguna       x   x   x   x   x   x   x   x
Southern        Teocaltiche     x   x   x   x   x   x   x   x
  Highlands     Villa Hidalgo   x   x       x   x   x   x   x
                Pegueros        x   x       x   x   x   x   x
Volcanic Belt   Amatitlan       x   x   x   x   x   x   x   x
                Jocotepec       x   x   x   x   x   x   x   x
                Ameca           x   x   x   x   x   x   x   x
Pacific Coast   La Gloria       x   x   x   x   x   x       x
                La Huerta       x   x   x   x   x   x       x

                                2012-   2013-   2014-
                                2013    2014    2015

Region          Route           A   W   A   W   A   W

Sierra Madre    San Juan        x   x   x   x   x   x
                La Laguna       x   x   x   x   x
Southern        Teocaltiche     x   x   x   x       x
  Highlands     Villa Hidalgo   x   x   x   x   x
                Pegueros        x   x   x   x   x
Volcanic Belt   Amatitlan       x   x   x   x   x   x
                Jocotepec       x   x   x   x   x
                Ameca           x   x   x   x   x
Pacific Coast   La Gloria       x   x   x       x   x
                La Huerta       x   x   x       x   x

                                2015-   2016-
                                2016    2017

Region          Route           A   W   A   W

Sierra Madre    San Juan
                La Laguna
Southern        Teocaltiche     x   x   x   x
  Highlands     Villa Hidalgo
Volcanic Belt   Amatitlan       x   x   x   x
                Jocotepec           x   x   x
                Ameca               x   x   x
Pacific Coast   La Gloria       x   x
                La Huerta       x   x

TABLE 3.--Model that best explained the changes in Mourning
Dove abundance in Jalisco survey routes as a function of the
population estimates in lhe different U.S. Mourning Dove Management
Units (M.U.), 1 y (i.e., 15 mo) before. AlCc values are indicated
only when they were very similar between different competing models.
Rainfall during the months previous to the doves' arrival was not
part of any best model. 2004-2017

Region                    U.S. Mourning Dove M.U. in best model

All our routes together   Central + Western M.U. (AlCc = 200);
                          central M.U. only (AICc = 201.8);
                          western M.U. only (AICc = 202.1)

Sierra Madre region       Western M.U.

Southern Highlands        Western + Central M.U. (AICc = 78.5);
                          Western M.U. only (AICc = 79.6, best
                          by rule of parsimony)

Volcanic Belt region      Central M.U.
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Author:Hernandez-Garcia, Rafael De Jesus; Mellink, Eric
Publication:The American Midland Naturalist
Article Type:Report
Geographic Code:1MEX
Date:Jul 1, 2018
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