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Population viability analysis of the Mato Grosso Antbird (Cercomacra melanaria) in the Pantanal of Mato Grosso, Brazil

Population viability analysis of the Mato Grosso Antbird (Cercomacra melanaria) in the Pantanal... Revista Brasileira de Ornitologia 25(3): 169–175. ARTICLE September 2017 Population viability analysis of the Mato Grosso Antbird (Cercomacra melanaria) in the Pantanal of Mato Grosso, Brazil 1,4 2 3,5 Paula Fernanda Albonette de Nóbrega , João Batista de Pinho & Charles Duca Pós-graduação em Ecologia e Conservação da Biodiversidade – Instituto de Biociências, Universidade Federal do Mato Grosso, Cuiabá, MT, Brazil Núcleo de Pesquisa Ecológica do Pantanal – Instituto de Biociências, Universidade Federal do Mato Grosso, 78075-960, Cuiabá, MT, Brazil. Laboratório de Ecologia de Populações e Conservação, Universidade Vila Velha, Campus Nossa Senhora da Penha, 29102-623, Vila Velha, ES, Brazil. Current Address: Faculdade de Ciências da Saúde, Universidade Estadual do Mato Grosso, Campus Diamantino, 784000-000 Diamantino, MT, Brazil. Corresponding author: cduca@uvv.br Received on 20 December 2016. Accepted on 02 November 2017. ABSTRACT: Understanding the interaction between habitat loss, environmental uncertainty, demographic stochasticity and genetics are the goal of population viability analysis. Minimum viable population (MVP) size is an estimate of the number of individuals of a species that will allow the continuation of the population into the foreseeable future. Here, we analyzed the population viability of Mato Grosso Antbird (Cercomacra melanaria), an endemic member of the Thamnophilidae from Pantanal, Brazil. We focus on estimating the MVP. We used the program VORTEX to generate simulations for MVP based on demographic and environmental variables, including catastrophes, carrying capacity, reproduction and survival. We also used sensitivity analysis to determine which variables are most influential in viability. We conclude that the Mato Grosso Antbird has low risk of extinction in the next 100 years under current conditions (quasi-extinction probability of 0.002). The MVP of Mato Grosso Antbir d was estimated to be 160 individuals. The most influential demographic parameter is the proportion of adult females that reproduce each year. Our results corroborate the classification of the Mato Grosso Antbir d as Least Concern by IUCN criterion. “ ” KEY-WORDS: endemic passerine, extinction probability, minimum viable population, PVA, Thamnophilidae. INTRODUCTION of persistence probability for many species (Diamond et al. 1987, Soulé et al. 1988, Miller & Lacy 2005). Small Population viability analysis (PVA) is a mathematical populations are very unstable and typically have high modeling tool that evaluates extinction risk and survival extinction probability due to demographic variability probability over time to direct conservation management (Brito & Grelle 2006). For example, the loss of more (Boyce 1992, Akçakaya & Sjögren-Gulve 2000, Keedwell than 20 bird species in the Bogor Botanical Garden (Indonesia) was due to small population sizes (Diamond 2004). This is important, because attempts to conserve small populations are often expensive, difficult, and et al. 1987). The MVP is also useful in estimating the usually offer only small chances of success (Soulé 1987, minimum reserve size that is necessary to the persistence Boyce 1992, Keedwell 2004). The minimum viable of the species (Grumbine 1990, Reed et al. 2002, Leech population (MVP) is the number of individuals that et al. 2008). Information about MVP is very important since habitat fragmentation and loss are key factors a population should contain to minimize the risk of extinction over a specified time interval (Shaffer 1981, influencing the distribution and abundance of threatened Boyce 1992). MVP is based on estimates of area occupied species (Lindenmayer & Lacy 2002). by the species of interest, its demographic trends and time More than one hundred bird species from Pantanal (Soulé 1987). Habitat loss, environmental uncertainty, are on threatened species lists (Tubelis & Tomás 2003) and will require some degree of management to secure demographic stochasticity, genetics (founder effects, genetic drift, inbreeding) and their interactions are the their persistence (Lindenmayer & Lacy 2002). The main important parameters of a PVA (Shaffer 1981). factors associated with bird extinctions are habitat loss, Population size is the most important determinant degradation, fragmentation and excessive harvesting Population viability analysis of the Cercomacra melanaria Nóbrega et al. (Marini & Garcia 2005), and more than one of these factors maximum-recorded age of a reproducing individual was may be affect the population viability (Temple 1986). Fire eight years and mean annual survival was 83% (Nóbrega and prolonged dry season are also factors that can impact 2009). Pairs may attempt more than one nest during each the breeding activities of birds (Duca et al. 2009, Duca breeding season and 50% of females produced at least & Marini 2011). The Mato Grosso Antbir d (Cercomacra one offspring each year (Nóbrega 2009). Nest predation melanaria, Thamnophilidae) is the Pantanal's onl y endemic is the most important cause of nest failure (68% of failed nests) (Pinho et al. 2006). Considering that the breeding bird (Silva et al. 2001). It was formerly considered as “Vulnerable” and a high priority species for conservation season of Mato Grosso Antbird start at the beginning by some authors (e.g., Brown-Jr. 1986, Olson et al. 1998, of the rainy season (Pinho et al. 2006), we expected Silva et al. 2001), but this species has been evaluated as that prolonged dry seasons should impact the breeding “Least Concern” since 1988 (BirdLife International 2016). activities (e.g., Duca et al. 2009, Duca & Marini 2011). Like most Neotropical bird species, only some aspects of the life history of the Mato Grosso Antbird are known Simulation model (Pinho et al. 2006, Bernardon et al. 2014) and therefore more information is required to determine its conservation Simulations were carried out using the program Vortex status in the Pantanal of the Mato Grosso. v. 9.95, which generates individual-based population models that include deterministic and stochastic processes The Mato Grosso Antbir d is a forest understory insectivore usually found near water, from the Bolivian (demographic, environmental and genetic) (Lacy 2000, Chaco and northern Paraguay to the Brazilian Pantanal Miller & Lacy 2005). The simulations included 500 (Ridgely & Tudor 1994, Sick 1997, Zimmer & Isler iterations for each scenario (Chapman et al. 2001, Brito 2003, Pinho et al. 2006). It is almost always found in & Fonseca 2006, Brito & Grelle 2006) and a 100-year time horizon for population dynamics (Walters et al. pairs in association with vines (Cissus spinosa, Cissus sicyoides) and shrubs in seasonally flooded areas ( Ridgely 2002, Miller & Lacy 2005). Initial population size (2500 individuals) was set at half that of the estimated carrying & Tudor 1994, Sick 1997, Silva et al. 2001, Zimmer & Isler 2003). In this study, we carry out a population capacity of the study area (e.g., Brito & Figueiredo 2003, viability analysis for the Mato Grosso Antbird. We used Brito & Fonseca 2006, Brito & Grelle 2006) (Table 1). We assume quasi-extinction when the population size simulations to examine persistence probability and the minimum viable population. falls below 50 individuals (e.g., Burgman et al. 1993) since the extinction probability of such population was high (13%) in the minimum viable population scenario METHODS (see section “Minimum Viable Population” below). Study area Scenarios The Mato Grosso Antbir d was studied in a 100 ha Basic scenario: Most data for this scenario come from forested area at the Retiro Novo Ranch (16 22'00''S; Nóbrega (2009) (Table 1). For survival estimates, we used 56 data from Myrmotherula fulviventris (Thamnophilidae) 17'57''W) in the Poconé Pantanal in Mato Grosso state, Brazil. The area is mostly savanna (see Nascimento (Greenberg & Gradwohl 1997) because its survivorship & Cunha 1989, Cunha & Junk 2001, Arieira & Cunha was estimated in a study conducted over 14 years, and 2006) and has a cold dry season (April – September) and this is a well-studied passerine with similar life history a warm rainy season (October–March). Average annual to the Mato Grosso Antbird. The consideration of the survivorship from M. fulviventris in the basic scenario temperature is 25.8 C, October is the hottest month o o (mean 34.0 C) and July is the coldest (mean 16.2 C). makes it an optimistic scenario because obligate mixed -1 Rainfall varies between 1000–1400 mm yr (mean 1250 flock following species typically have higher survivorship -1 mm.yr ), 80% of which falls between November–March than for antbird species feeding alone or in pairs (Jullien (Allem & Valls 1987). & Clobert 2000). The carrying capacity was estimated at 5000 individuals and details about this estimate are Life history attributes of Mato Grosso Antbird available in Nóbrega (2009). Prolonged dry seasons were included as catastrophe (Table 1). The Mato Grosso Antbir d is a monogamous and territorial The basic scenario is the one that matches the species, with territories of about 0.32 ha (Bernardon situation of Mato Grosso Antbird population in the study area, because most of the parameters were developed 2007). Clutch size is two eggs and rarely one egg (mean 1.9 eggs), and both sexes begin reproduction at one using data collected there (see Nóbrega 2009). In all year old (Pinho et al. 2006, Bernardon et al. 2014). The scenarios that follow, specific variables were modified to Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. Table 1. Summary of input parameters of Mato Grosso Antbird, Cercomacra melanaria, used in basic scenarios using computer program VORTEX. Source Parameters Value Reproduction and survival Correlated Number/type of catastrophe1 (long dry season) Reproduction Monogamous ab Age first reproduction females and males 1 Maximum reproductive age 8 ab Maximum young per year 2 Sex ratio at birth (%) 50 Reproductive females (%) 50.1 ab Environmental variation in reproduction (%) 9 ab Adult males breeding (%) 100 -1 -1 Number of offspring female year Yes ab 1 young (%) 19 ab 2 young (%) 81 First year mortality (%) 23.5 Adult mortality (%) 25 ab Environmental variation in death rate (%) 11 Catastrophe Long dry season Frequency (%) 10 Impact on reproduction (% reduction) 25 Impact on survival No Carrying capacity, K 5000 Environmental variation in K (%) 100 (2) a b c d e Sources: Nóbrega (2009), Bernardon (2007), Duca et al. (2009), Greenberg & Gradwohl (1997), INMET (2008). assess different assumptions about uncertain parameters. assume that the severity of fire results in a 25% reduction All the other variables not specifically mentioned were in reproductive success. The two catastrophes (fire and maintained at their value listed in the basic scenario. prolonged dry seasons) may both cause habitat loss Minimum Viable Population scenario (MVP): to and influence breeding season length, reducing nesting estimate MVP, we changed the initial population size success during the catastrophes. parameter in Vortex and modeled different scenarios with MVP with two catastrophes: here we estimate MVP initial population sizes of 1250, 625, 312, 156, 100, and with two catastrophes – fire and prolonged dry season. 50 individuals. Initial population sizes as above in the MVP scenario. Mortality scenario: we reduced the mortality rate from basic scenario to the one found by Nóbrega (2009). Sensitivity analysis Therefore, the annual mortality was reduced from 25% to 17% for adults (≥ 1 year old) and from 23.5% to 15.9% Sensitivity analysis was used to test the robustness of for juveniles (< 1 year old). some parameters and to assess which parameters have the Fire scenario: in this scenario we add fire as a se cond greatest influence on MVP ( e.g., Miller & Lacy 2005, catastrophe. Due to the lack of official fire recor ds in the Brito & Fonseca 2006, Duca et al. 2009). We changed region, we spoke with residents of the area and estimated mortality rates and number of reproductive adults up and this catastrophe with an annual probability of 10%. We down by 10%, 15% and 20% (Fig. 1). Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. Figure 1. Extinction probability with variation of 10%, 15% Figure 2. Relationship between initial population size and and 20% in adult and juvenile mortality, and variation of extinction probability for the Mato Grosso Antbird (Cercomacra -10%, -15%, -20% on percentage of females breeding and melanaria) in the Pantanal of Poconé in scenarios with one on percentage of males in breeding pool for the Mato Grosso (basic scenario) and two catastrophes. Antbird (Cercomacra melanaria) population in the Pantanal of Poconé. The remaining variables are as in the basic scenario. Table 2. Results summaries of PVA of the Mato Grosso Antbird, Cercomacra melanaria, using program VORTEX. Population growth rate (r) Final population size Scenario Extinction probability ± SD Deterministic Stochastic ± SD Basic 0.002 0.064 0.051 ± 0.198 3858.61 ± 1195.41 Mortality 0.002 0.163 0.152 ± 0.179 4787.05 ± 376.12 Fire 0.002 0.055 0.042 ± 0.198 3571.59 ± 1297.42 RESULTS initial population size had a greater extinction probability than the basic scenario, and again the reproductive females have the greatest impact on extinction probability. Population viability However, in this scenario adult mortality rate was also The Mato Grosso Antbir d population has a low risk of important when reaching 10% greater mortality than in the basic setting (EP = 0.214). Additionally, the becoming quasi-extinct in the next 100 years (Extinction proportion of adult males that were reproductive as well Probability = 0.002, basic scenario) (Table 2). Sensibility as juvenile mortality was also important but their effects analysis showed that the percentage of reproductive on extinction probability were weaker than reproductive females had the greatest impact on extinction probability, and mortality was less important (Fig. 1). females and adult mortality (EP = 0.052 and EP = 0.102, respectively) (Fig. 3). Minimum Viable Population (MVP) DISCUSSION Extinction probability (EP) was higher than 0.05 when the initial population size was 50 individuals (Fig. 2). Population viability With population sizes greater than 100 individuals, the population was viable (EP = 0.036). Thus, MVP of Mato Demographic and environmental processes combined Grosso Antbird with one catastrophe was approximately with the influence of catastrophes indicate that the Mato 100 individuals. With two catastrophes, MVP was Grosso Antbird has low chance of becoming extinct around 160 individuals with low extinction probability under the prevailing conditions and a tendency towards (EP = 0.042) and positive stochastic population growth positive population growth. Even when using pessimistic rate (Table 2, Fig. 2). levels of proportions of reproductive adults and adult and Sensibility analysis with MVP (160 individuals) as Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. all populations of Mato Grosso Antbird higher than 150 individuals were viable. Other studies have similar results (e.g., Soulé et al. 1988, Thomas 1990, Hamilton & Moller 1995, Leech et al. 2008) and estimates of MVP based on pessimist scenarios are more appropriated when the goal is to do a conservative analysis. Catastrophes, such as fire and long dry seasons, are known to impact the persistence probability of populations (Cahill & Walker 2000, Bolger et al. 2005, Dawson & Bortolotti 2006). Therefore, catastrophes raise the estimated MVP over that in the basic scenario to approximately 160 individuals. Prolonged drought is important because the Mato Grosso Antbird begins reproduction with the onset of the rainy season (Pinho et al. 2006). Thus, due to its limited breeding season (Hau et al. 2008), a long dry season shortens the time interval over which reproduction may occur, thereby decreasing annual fecundity. If the Figure 3. Extinction probability with variation of 5% and 10 frequency of dry years increases, population viability % in adult and juvenile mortality, and variation of -10% and will subsequently decline (Bolger et al. 2005, Duca et al. -5 % on percentage of females breeding and on percentage 2009). of males in breeding pool for the Mato Grosso Antbird The influence of fire is more direct, as it reduces (Cercomacra melanaria) population in the Pantanal of Poconé survival probability, causes habitat loss and degradation using the alternative scenario of MVP of 160 individuals and fire catastrophe. The remaining variables are as in the basic and may cause nest failure (Cahill & Walker 2000). scenario. Fire was associated with reduced reproductive success in Falco sparverius and Aceros cassidix (Cahill & Walker juvenile mortality, extinction probability remains low. 2000, Dawson & Bortolotti 2006). This is likely to be a These results corroborate the classification of the Mato catastrophe that may increase its frequency in the Poconé Grosso Antbird as “Least Concern” by IUCN criterion area because of the common practice of burning fields (BirdLife International 2016). every year (Harris et al. 2005). As indicated by the sensitivity analysis, the When including the catastrophic events we percentage of reproductive females is the most important can see the population sensitivity to the presence of parameter for extinction risk as in other bird studies such factors. Catastrophes affect population viability (França & Marini 2010, Duca et al. 2009) and those of because they affect other parameters determining the other vertebrates (Goldingay & Possingham 1995, Reed continuity of the population persistence. The sensitivity et al. 1998, Brito & Grelle 2006, Brito et al. 2008). Adult analysis of scenarios with two catastrophes indicate mortality was also indicated by the sensitivity analysis that reproductive parameters are most relevant in the as the second most important parameter to population extinction process and should be prioritized in the viability. Theses results suggest that management strategies definition of management strategies for Mato Grosso (e.g., nest protection - Duca et al. 2009) should be first Antbird population. Studies carried out in the Cerrado directed at female reproduction to improve persistence in Biome suggest that management strategies should be this population, and that future studies should prioritize directed towards increasing reproduction rates rather than estimates of fecundity and mortality, and their variances. to manipulate other factors, such as habitat availability We highlight that our results are indicating a (Duca et al. 2009, França & Marini 2010). Also, adult tendency of the Mato Grosso Antbird population in the survival is clearly important because the impact of study area, but the conclusions should be viewed with stochastic variation in the size of the effective population caution because there are some uncertainties in the input can be eased by the presence of adults able to replace died parameters of the model. breeders (Goldingay & Possingham 1995, Walters et al. 2002). Therefore, providing refuge or other means of Minimum Viable Population (MVP) escaping catastrophes are important management options (Duca et al. 2009). The population size of Mato Grosso Antbir d must The Mato Grosso Antbir d population in the current remain above 50 individuals to be viable (> 95% chance demographic and environmental settings, including to persist in the next 100 years). In a pessimist scenario catastrophes, is not vulnerable to extinction. 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Population viability analysis of the Mato Grosso Antbird (Cercomacra melanaria) in the Pantanal of Mato Grosso, Brazil

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Abstract

Revista Brasileira de Ornitologia 25(3): 169–175. ARTICLE September 2017 Population viability analysis of the Mato Grosso Antbird (Cercomacra melanaria) in the Pantanal of Mato Grosso, Brazil 1,4 2 3,5 Paula Fernanda Albonette de Nóbrega , João Batista de Pinho & Charles Duca Pós-graduação em Ecologia e Conservação da Biodiversidade – Instituto de Biociências, Universidade Federal do Mato Grosso, Cuiabá, MT, Brazil Núcleo de Pesquisa Ecológica do Pantanal – Instituto de Biociências, Universidade Federal do Mato Grosso, 78075-960, Cuiabá, MT, Brazil. Laboratório de Ecologia de Populações e Conservação, Universidade Vila Velha, Campus Nossa Senhora da Penha, 29102-623, Vila Velha, ES, Brazil. Current Address: Faculdade de Ciências da Saúde, Universidade Estadual do Mato Grosso, Campus Diamantino, 784000-000 Diamantino, MT, Brazil. Corresponding author: cduca@uvv.br Received on 20 December 2016. Accepted on 02 November 2017. ABSTRACT: Understanding the interaction between habitat loss, environmental uncertainty, demographic stochasticity and genetics are the goal of population viability analysis. Minimum viable population (MVP) size is an estimate of the number of individuals of a species that will allow the continuation of the population into the foreseeable future. Here, we analyzed the population viability of Mato Grosso Antbird (Cercomacra melanaria), an endemic member of the Thamnophilidae from Pantanal, Brazil. We focus on estimating the MVP. We used the program VORTEX to generate simulations for MVP based on demographic and environmental variables, including catastrophes, carrying capacity, reproduction and survival. We also used sensitivity analysis to determine which variables are most influential in viability. We conclude that the Mato Grosso Antbird has low risk of extinction in the next 100 years under current conditions (quasi-extinction probability of 0.002). The MVP of Mato Grosso Antbir d was estimated to be 160 individuals. The most influential demographic parameter is the proportion of adult females that reproduce each year. Our results corroborate the classification of the Mato Grosso Antbir d as Least Concern by IUCN criterion. “ ” KEY-WORDS: endemic passerine, extinction probability, minimum viable population, PVA, Thamnophilidae. INTRODUCTION of persistence probability for many species (Diamond et al. 1987, Soulé et al. 1988, Miller & Lacy 2005). Small Population viability analysis (PVA) is a mathematical populations are very unstable and typically have high modeling tool that evaluates extinction risk and survival extinction probability due to demographic variability probability over time to direct conservation management (Brito & Grelle 2006). For example, the loss of more (Boyce 1992, Akçakaya & Sjögren-Gulve 2000, Keedwell than 20 bird species in the Bogor Botanical Garden (Indonesia) was due to small population sizes (Diamond 2004). This is important, because attempts to conserve small populations are often expensive, difficult, and et al. 1987). The MVP is also useful in estimating the usually offer only small chances of success (Soulé 1987, minimum reserve size that is necessary to the persistence Boyce 1992, Keedwell 2004). The minimum viable of the species (Grumbine 1990, Reed et al. 2002, Leech population (MVP) is the number of individuals that et al. 2008). Information about MVP is very important since habitat fragmentation and loss are key factors a population should contain to minimize the risk of extinction over a specified time interval (Shaffer 1981, influencing the distribution and abundance of threatened Boyce 1992). MVP is based on estimates of area occupied species (Lindenmayer & Lacy 2002). by the species of interest, its demographic trends and time More than one hundred bird species from Pantanal (Soulé 1987). Habitat loss, environmental uncertainty, are on threatened species lists (Tubelis & Tomás 2003) and will require some degree of management to secure demographic stochasticity, genetics (founder effects, genetic drift, inbreeding) and their interactions are the their persistence (Lindenmayer & Lacy 2002). The main important parameters of a PVA (Shaffer 1981). factors associated with bird extinctions are habitat loss, Population size is the most important determinant degradation, fragmentation and excessive harvesting Population viability analysis of the Cercomacra melanaria Nóbrega et al. (Marini & Garcia 2005), and more than one of these factors maximum-recorded age of a reproducing individual was may be affect the population viability (Temple 1986). Fire eight years and mean annual survival was 83% (Nóbrega and prolonged dry season are also factors that can impact 2009). Pairs may attempt more than one nest during each the breeding activities of birds (Duca et al. 2009, Duca breeding season and 50% of females produced at least & Marini 2011). The Mato Grosso Antbir d (Cercomacra one offspring each year (Nóbrega 2009). Nest predation melanaria, Thamnophilidae) is the Pantanal's onl y endemic is the most important cause of nest failure (68% of failed nests) (Pinho et al. 2006). Considering that the breeding bird (Silva et al. 2001). It was formerly considered as “Vulnerable” and a high priority species for conservation season of Mato Grosso Antbird start at the beginning by some authors (e.g., Brown-Jr. 1986, Olson et al. 1998, of the rainy season (Pinho et al. 2006), we expected Silva et al. 2001), but this species has been evaluated as that prolonged dry seasons should impact the breeding “Least Concern” since 1988 (BirdLife International 2016). activities (e.g., Duca et al. 2009, Duca & Marini 2011). Like most Neotropical bird species, only some aspects of the life history of the Mato Grosso Antbird are known Simulation model (Pinho et al. 2006, Bernardon et al. 2014) and therefore more information is required to determine its conservation Simulations were carried out using the program Vortex status in the Pantanal of the Mato Grosso. v. 9.95, which generates individual-based population models that include deterministic and stochastic processes The Mato Grosso Antbir d is a forest understory insectivore usually found near water, from the Bolivian (demographic, environmental and genetic) (Lacy 2000, Chaco and northern Paraguay to the Brazilian Pantanal Miller & Lacy 2005). The simulations included 500 (Ridgely & Tudor 1994, Sick 1997, Zimmer & Isler iterations for each scenario (Chapman et al. 2001, Brito 2003, Pinho et al. 2006). It is almost always found in & Fonseca 2006, Brito & Grelle 2006) and a 100-year time horizon for population dynamics (Walters et al. pairs in association with vines (Cissus spinosa, Cissus sicyoides) and shrubs in seasonally flooded areas ( Ridgely 2002, Miller & Lacy 2005). Initial population size (2500 individuals) was set at half that of the estimated carrying & Tudor 1994, Sick 1997, Silva et al. 2001, Zimmer & Isler 2003). In this study, we carry out a population capacity of the study area (e.g., Brito & Figueiredo 2003, viability analysis for the Mato Grosso Antbird. We used Brito & Fonseca 2006, Brito & Grelle 2006) (Table 1). We assume quasi-extinction when the population size simulations to examine persistence probability and the minimum viable population. falls below 50 individuals (e.g., Burgman et al. 1993) since the extinction probability of such population was high (13%) in the minimum viable population scenario METHODS (see section “Minimum Viable Population” below). Study area Scenarios The Mato Grosso Antbir d was studied in a 100 ha Basic scenario: Most data for this scenario come from forested area at the Retiro Novo Ranch (16 22'00''S; Nóbrega (2009) (Table 1). For survival estimates, we used 56 data from Myrmotherula fulviventris (Thamnophilidae) 17'57''W) in the Poconé Pantanal in Mato Grosso state, Brazil. The area is mostly savanna (see Nascimento (Greenberg & Gradwohl 1997) because its survivorship & Cunha 1989, Cunha & Junk 2001, Arieira & Cunha was estimated in a study conducted over 14 years, and 2006) and has a cold dry season (April – September) and this is a well-studied passerine with similar life history a warm rainy season (October–March). Average annual to the Mato Grosso Antbird. The consideration of the survivorship from M. fulviventris in the basic scenario temperature is 25.8 C, October is the hottest month o o (mean 34.0 C) and July is the coldest (mean 16.2 C). makes it an optimistic scenario because obligate mixed -1 Rainfall varies between 1000–1400 mm yr (mean 1250 flock following species typically have higher survivorship -1 mm.yr ), 80% of which falls between November–March than for antbird species feeding alone or in pairs (Jullien (Allem & Valls 1987). & Clobert 2000). The carrying capacity was estimated at 5000 individuals and details about this estimate are Life history attributes of Mato Grosso Antbird available in Nóbrega (2009). Prolonged dry seasons were included as catastrophe (Table 1). The Mato Grosso Antbir d is a monogamous and territorial The basic scenario is the one that matches the species, with territories of about 0.32 ha (Bernardon situation of Mato Grosso Antbird population in the study area, because most of the parameters were developed 2007). Clutch size is two eggs and rarely one egg (mean 1.9 eggs), and both sexes begin reproduction at one using data collected there (see Nóbrega 2009). In all year old (Pinho et al. 2006, Bernardon et al. 2014). The scenarios that follow, specific variables were modified to Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. Table 1. Summary of input parameters of Mato Grosso Antbird, Cercomacra melanaria, used in basic scenarios using computer program VORTEX. Source Parameters Value Reproduction and survival Correlated Number/type of catastrophe1 (long dry season) Reproduction Monogamous ab Age first reproduction females and males 1 Maximum reproductive age 8 ab Maximum young per year 2 Sex ratio at birth (%) 50 Reproductive females (%) 50.1 ab Environmental variation in reproduction (%) 9 ab Adult males breeding (%) 100 -1 -1 Number of offspring female year Yes ab 1 young (%) 19 ab 2 young (%) 81 First year mortality (%) 23.5 Adult mortality (%) 25 ab Environmental variation in death rate (%) 11 Catastrophe Long dry season Frequency (%) 10 Impact on reproduction (% reduction) 25 Impact on survival No Carrying capacity, K 5000 Environmental variation in K (%) 100 (2) a b c d e Sources: Nóbrega (2009), Bernardon (2007), Duca et al. (2009), Greenberg & Gradwohl (1997), INMET (2008). assess different assumptions about uncertain parameters. assume that the severity of fire results in a 25% reduction All the other variables not specifically mentioned were in reproductive success. The two catastrophes (fire and maintained at their value listed in the basic scenario. prolonged dry seasons) may both cause habitat loss Minimum Viable Population scenario (MVP): to and influence breeding season length, reducing nesting estimate MVP, we changed the initial population size success during the catastrophes. parameter in Vortex and modeled different scenarios with MVP with two catastrophes: here we estimate MVP initial population sizes of 1250, 625, 312, 156, 100, and with two catastrophes – fire and prolonged dry season. 50 individuals. Initial population sizes as above in the MVP scenario. Mortality scenario: we reduced the mortality rate from basic scenario to the one found by Nóbrega (2009). Sensitivity analysis Therefore, the annual mortality was reduced from 25% to 17% for adults (≥ 1 year old) and from 23.5% to 15.9% Sensitivity analysis was used to test the robustness of for juveniles (< 1 year old). some parameters and to assess which parameters have the Fire scenario: in this scenario we add fire as a se cond greatest influence on MVP ( e.g., Miller & Lacy 2005, catastrophe. Due to the lack of official fire recor ds in the Brito & Fonseca 2006, Duca et al. 2009). We changed region, we spoke with residents of the area and estimated mortality rates and number of reproductive adults up and this catastrophe with an annual probability of 10%. We down by 10%, 15% and 20% (Fig. 1). Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. Figure 1. Extinction probability with variation of 10%, 15% Figure 2. Relationship between initial population size and and 20% in adult and juvenile mortality, and variation of extinction probability for the Mato Grosso Antbird (Cercomacra -10%, -15%, -20% on percentage of females breeding and melanaria) in the Pantanal of Poconé in scenarios with one on percentage of males in breeding pool for the Mato Grosso (basic scenario) and two catastrophes. Antbird (Cercomacra melanaria) population in the Pantanal of Poconé. The remaining variables are as in the basic scenario. Table 2. Results summaries of PVA of the Mato Grosso Antbird, Cercomacra melanaria, using program VORTEX. Population growth rate (r) Final population size Scenario Extinction probability ± SD Deterministic Stochastic ± SD Basic 0.002 0.064 0.051 ± 0.198 3858.61 ± 1195.41 Mortality 0.002 0.163 0.152 ± 0.179 4787.05 ± 376.12 Fire 0.002 0.055 0.042 ± 0.198 3571.59 ± 1297.42 RESULTS initial population size had a greater extinction probability than the basic scenario, and again the reproductive females have the greatest impact on extinction probability. Population viability However, in this scenario adult mortality rate was also The Mato Grosso Antbir d population has a low risk of important when reaching 10% greater mortality than in the basic setting (EP = 0.214). Additionally, the becoming quasi-extinct in the next 100 years (Extinction proportion of adult males that were reproductive as well Probability = 0.002, basic scenario) (Table 2). Sensibility as juvenile mortality was also important but their effects analysis showed that the percentage of reproductive on extinction probability were weaker than reproductive females had the greatest impact on extinction probability, and mortality was less important (Fig. 1). females and adult mortality (EP = 0.052 and EP = 0.102, respectively) (Fig. 3). Minimum Viable Population (MVP) DISCUSSION Extinction probability (EP) was higher than 0.05 when the initial population size was 50 individuals (Fig. 2). Population viability With population sizes greater than 100 individuals, the population was viable (EP = 0.036). Thus, MVP of Mato Demographic and environmental processes combined Grosso Antbird with one catastrophe was approximately with the influence of catastrophes indicate that the Mato 100 individuals. With two catastrophes, MVP was Grosso Antbird has low chance of becoming extinct around 160 individuals with low extinction probability under the prevailing conditions and a tendency towards (EP = 0.042) and positive stochastic population growth positive population growth. Even when using pessimistic rate (Table 2, Fig. 2). levels of proportions of reproductive adults and adult and Sensibility analysis with MVP (160 individuals) as Revista Brasileira de Ornitologia 25(3): 2017 Population viability analysis of the Cercomacra melanaria Nóbrega et al. all populations of Mato Grosso Antbird higher than 150 individuals were viable. Other studies have similar results (e.g., Soulé et al. 1988, Thomas 1990, Hamilton & Moller 1995, Leech et al. 2008) and estimates of MVP based on pessimist scenarios are more appropriated when the goal is to do a conservative analysis. Catastrophes, such as fire and long dry seasons, are known to impact the persistence probability of populations (Cahill & Walker 2000, Bolger et al. 2005, Dawson & Bortolotti 2006). Therefore, catastrophes raise the estimated MVP over that in the basic scenario to approximately 160 individuals. Prolonged drought is important because the Mato Grosso Antbird begins reproduction with the onset of the rainy season (Pinho et al. 2006). Thus, due to its limited breeding season (Hau et al. 2008), a long dry season shortens the time interval over which reproduction may occur, thereby decreasing annual fecundity. If the Figure 3. Extinction probability with variation of 5% and 10 frequency of dry years increases, population viability % in adult and juvenile mortality, and variation of -10% and will subsequently decline (Bolger et al. 2005, Duca et al. -5 % on percentage of females breeding and on percentage 2009). of males in breeding pool for the Mato Grosso Antbird The influence of fire is more direct, as it reduces (Cercomacra melanaria) population in the Pantanal of Poconé survival probability, causes habitat loss and degradation using the alternative scenario of MVP of 160 individuals and fire catastrophe. The remaining variables are as in the basic and may cause nest failure (Cahill & Walker 2000). scenario. Fire was associated with reduced reproductive success in Falco sparverius and Aceros cassidix (Cahill & Walker juvenile mortality, extinction probability remains low. 2000, Dawson & Bortolotti 2006). This is likely to be a These results corroborate the classification of the Mato catastrophe that may increase its frequency in the Poconé Grosso Antbird as “Least Concern” by IUCN criterion area because of the common practice of burning fields (BirdLife International 2016). every year (Harris et al. 2005). As indicated by the sensitivity analysis, the When including the catastrophic events we percentage of reproductive females is the most important can see the population sensitivity to the presence of parameter for extinction risk as in other bird studies such factors. Catastrophes affect population viability (França & Marini 2010, Duca et al. 2009) and those of because they affect other parameters determining the other vertebrates (Goldingay & Possingham 1995, Reed continuity of the population persistence. The sensitivity et al. 1998, Brito & Grelle 2006, Brito et al. 2008). Adult analysis of scenarios with two catastrophes indicate mortality was also indicated by the sensitivity analysis that reproductive parameters are most relevant in the as the second most important parameter to population extinction process and should be prioritized in the viability. Theses results suggest that management strategies definition of management strategies for Mato Grosso (e.g., nest protection - Duca et al. 2009) should be first Antbird population. Studies carried out in the Cerrado directed at female reproduction to improve persistence in Biome suggest that management strategies should be this population, and that future studies should prioritize directed towards increasing reproduction rates rather than estimates of fecundity and mortality, and their variances. to manipulate other factors, such as habitat availability We highlight that our results are indicating a (Duca et al. 2009, França & Marini 2010). Also, adult tendency of the Mato Grosso Antbird population in the survival is clearly important because the impact of study area, but the conclusions should be viewed with stochastic variation in the size of the effective population caution because there are some uncertainties in the input can be eased by the presence of adults able to replace died parameters of the model. breeders (Goldingay & Possingham 1995, Walters et al. 2002). Therefore, providing refuge or other means of Minimum Viable Population (MVP) escaping catastrophes are important management options (Duca et al. 2009). The population size of Mato Grosso Antbir d must The Mato Grosso Antbir d population in the current remain above 50 individuals to be viable (> 95% chance demographic and environmental settings, including to persist in the next 100 years). In a pessimist scenario catastrophes, is not vulnerable to extinction. 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Journal

Ornithology ResearchSpringer Journals

Published: Sep 1, 2017

Keywords: endemic passerine; extinction probability; minimum viable population; PVA; Thamnophilidae

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