Variation in Mesopredator Abundance and Nest Predation Rate of The Endangered Strange-Tailed Tyrant (Alectrurus Risora)

Ibis (2023), 165, 1201–1216 doi: 10.1111/ibi.
13202
Variation in mesopredator abundance and nest

predation rate of the endangered Strange-tailed
Tyrant (Alectrurus risora)
MELANIE BROWNE,*1 CONSTANZA PASIAN,1 ALEJANDRO G. DI GIACOMO,2 MARIO S. DI BITETTI3,4,5 &
S. DI GIACOMO1
ADRIAN
1

Laboratorio de Biologıa de la Conservacio n, Centro de Ecologıa Aplicada del Litoral (CECOAL-CONICET),
Corrientes, Argentina
2
Departamento de Conservacio n, Aves Argeninas/Asociacio n Ornitolo
gica del Plata, Matheu 1246, Buenos Aires,
C1249AAB, Argentina
3
Instituto de Biologıa Subtropical (IBS), Universidad Nacional de Misiones (UNaM) – CONICET, Bertoni 85, Puerto
, Misiones, Argentina
Iguazu
4

Asociacio n Civil Centro de Investigaciones del Bosque Atla ntico (CeIBA), Bertoni 85, Puerto Iguazu , Misiones,
Argentina
5
Facultad de Ciencias Forestales, UNaM, Bertoni 124, Eldorado, Misiones, Argentina
Nest predation is the main cause of nest failure in passerine birds and thus is a key deter-
minant of breeding traits. Accordingly, nest predator community composition and abun-
dance modulate nest predation rates. When top predators are absent, mesopredator
populations may become unchecked, which in turn increases the predation pressure on
their prey species. We evaluated whether nest predator abundance and breeding traits of
an endangered grassland bird, the Strange-tailed Tyrant Alectrurus risora, differed
between two protected areas: Ibera National Park (INP), where top predators have been
extinct for 50 years, and Reserva El Bagual (REB), where Pumas Puma concolor still have
a stable population. We measured abundances of potential nest predator species and dur-
ing the 2015–2018 breeding seasons we monitored Strange-tailed Tyrant nests and set
camera traps to identify nest predators. Mesopredator abundance was higher in INP, and
foxes were identified as the main nest predators. Nest predation rate was also higher in
INP, where the daily nest predation rate and cumulative probability of nest predation
were 0.05 0.006 and 0.81, whereas in REB they were 0.04 0.004 and 0.70, respec-
tively. The 10-fold higher frequency of fox records in INP could explain the higher nest
predation rate observed, suggesting that the extinction of top predators may increase nest
predation rates of endangered grassland birds in protected areas through release of native
mesopredator species. The lower nest predation observed in REB was compensated for
by a higher nest abandonment rate, which resulted in a similar fledgling production in
the two study sites. Clutch size, hatching success and number of nestlings did not differ
between study sites. The low breeding success recorded for this endangered species
despite its populations being in protected areas of high conservation value is alarming
given the global decline that grassland bird populations are facing.
Keywords: breeding biology, camera trap, mesopredator release, nest concealment, nest predators,
subtropical grasslands.
Habitat loss is one of the main drivers of biodiver-

sity decline, and grasslands are among the most
*Corresponding author.
imperilled ecosystems due to their alarming loss
Email: [email protected] and degradation rates (Hoekstra et al. 2005,
© 2023 British Ornithologists' Union.

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1202 M. Browne et al.
Watson et al. 2016). As a result, grassland birds Jimenez et al. 2019, Rees et al. 2019), through
are suffering negative population trends worldwide increasing nest predation rates and reducing breed-
(Azpiroz et al. 2012). Subtropical grasslands are ing success (Rogers & Caro 1998, Ritchie & John-
among the ecosystems suffering the highest rates son 2009, Ortega et al. 2021).
of habitat conversion and, as a consequence, 22 Northeastern Argentina’s grasslands are being
bird species of southeastern South American grass- transformed by intensification of cattle grazing and
lands are globally threatened or near-threatened afforestation (Viglizzo et al. 2011, Azpiroz
with extinction (Azpiroz et al. 2012, Watson et al. 2012). In addition, Jaguars Panthera onca and
et al. 2016). Pumas Puma concolor have been the subject of per-
The main cause of nest failure in many passer- secution due to conflict with livestock manage-
ine birds is nest predation, making it a key modu- ment, resulting in the extinction of the former and
lator of avian life history traits (Ricklefs 1969, a population reduction of the latter in northeastern
Martin 1995). Birds seek to minimize the risk of Argentina’s grasslands (Graham et al. 2005, Quir-
nest failure by selecting both nest-sites that are oga et al. 2014, 2016, Di Bitetti et al. 2016).
concealed from potential predators and nest sub- Simultaneously, the Strange-tailed Tyrant Alec-
strates that impede predation (Martin & trurus risora, an endemic grassland species that
Roper 1988, Schmidt & Whelan 1999, Misenhel- nests in tallgrass tussocks and occurs primarily in
ter & Rotenberry 2000). Nest predation probabil- northeastern Argentina and southeastern Paraguay,
ity may also increase with nest age, as increasing has suffered an estimated 90% reduction in its
adult and nestling activity risk betraying the nest range in Argentina (Di Giacomo & Di Gia-
position to potential predators (Skutch 1949). como 2004, Di Giacomo et al. 2011). This drastic
Hence, birds may respond to an increase in nest range reduction and the threat that habitat conver-
predation risk by switching their nest-site choice in sion poses for its remaining natural habitat has led
favour of better concealed sites and regulating the species to be categorized as globally Vulnera-
their behaviour by reducing activity at the nest ble (BirdLife International 2016).
(Skutch 1949, Fontaine & Martin 2006, Zanette The main goal of this study was to evaluate if
et al. 2011). Breeding traits such as clutch size and differences in the breeding biology of the
fledgling production can also be reduced by an Strange-tailed Tyrant in grasslands of high conser-
increase in nest predation risk (Zanette et al. 2011, vation value are associated with differences in
Dillon & Conway 2018). The influence of nest predator abundance and predation rate. Given
predation on annual breeding success is condi- that one of our study sites has a stable Puma
tioned by breeding season length and renesting population but top predators are locally extinct in
intervals, which determine the number of breeding the other area, we studied the nest predator com-
attempts (Roper et al. 2010). Nest predation may munity to evaluate whether nest predator abun-
therefore regulate multiple breeding traits of birds. dance increased where top predators were absent
Nest predation is also modulated by nest preda- and whether this variation in nest predator abun-
tor community composition and predator abundance could be associated with differences in the
dance (Schmidt 2003, Roos et al. 2018). Habitat Strange-tailed Tyrant’s breeding biology. We first
fragmentation, the extinction or suppression of top evaluated differences in the abundance of poten-
predators and the introduction of exotic species tial nest predators, including carnivorous mam-
may increase nest predator densities (Roos mals, raptors and snakes, in both study areas.
et al. 2018, Saggiomo et al. 2021). The mesopre- Next, we compared the breeding biology of the
dator release hypothesis states that the decline or Strange-tailed Tyrant in the two areas, to deter-
extinction of top predator populations may trigger mine whether its breeding phenology, breeding
an increase in mesopredator abundance (Soule parameters (i.e. clutch size, hatching success,
et al. 1988, Crooks & Soule 1999). As mesopreda- number of nestlings and fledgling production),
tor abundance increases, the predation rate on nest predation rate, daily nest survival rate and
their prey is also expected to increase. For exam- nest concealment differed between them. We
ple, mesopredator release has been reported to predicted a higher abundance of mesopredators
negatively impact bird populations, sometimes with negative effects on the breeding biology of
even leading to local extinctions (Soule the Strange-tailed Tyrant in the study area where
et al. 1988, Crooks & Soule 1999, Schmidt 2003, top predators are extinct.

Strange-tailed Tyrant nest predation 1203
medium- and large-sized mammal populations at

METHODS
our study sites. We used Reconyx HyperFire
HC500 cameras baited with perforated tuna cans
Study area
to attract carnivorous species, as described by Di
The study was conducted in Portal San Nicolas Bitetti et al. (2009). Cameras were set to take
(28.1282°S, 57.4346°W; 16 780 ha) in Ibera three pictures per trigger, with a picture interval
National Park (INP), located in the province of of 1 s and a quiet period of 3 min. We replaced
Corrientes, and in Reserva El Bagual (REB; bait, batteries and memory cards every 10 days.
28.1869°S, 58.8550°W; 3300 ha), located in the Cameras were simultaneously active in both sites
province of Formosa, Argentina (Fig. 1). Jaguars and sampling was divided into two stages, with
have been extinct in Corrientes for over 50 years half of the stations active during March–April and
and Pumas are mostly absent or extremely rare the other half during May–June.
whereas, in Formosa, Puma populations are still We conducted raptor surveys during the 2017
stable and Jaguars, even though rare, are also pre- Strange-tailed Tyrant breeding season by fixing six
sent (Di Giacomo & Krapovickas 2005, Quiroga and two 1-km-long transects across internal roads
et al. 2014, 2016, Di Bitetti et al. 2016). Both pro- in INP and REB, respectively (Ralph et al. 1993).
tected areas are in the humid Chaco ecoregion Transects were separated by a minimum distance
(Oyarzabal et al. 2018). The landscapes of the of 500 m. We visited each transect on four occa-
study areas are physiognomically similar and con- sions at both sites and recorded all raptor species
sist of open savannas with gradual slopes that and the number of individuals observed within a
result in seasonally flooded grasslands in the low- radius of 100 m from the observer.
lands and drained grasslands in the uplands. We sampled snakes by setting six array traps in
Uplands are dominated by Andropogon lateralis in each study site, during the 2017 breeding season.
INP, and by Imperata brasiliensis and A. lateralis in We built array traps following the design pro-
REB (Carnevali 1994, Di Giacomo & Krapo- posed by Leynaud and Bucher (2005) with 8-m-
vickas 2005). Lowlands are dominated by Paspa- long legs. Array traps were separated by a mini-
lum durifolium in INP and by Paspalum mum distance of 600 m. The traps were active in
intermedium and Sorghastrum setosum in REB (Car- both sites for 42 consecutive days and we con-
nevali 1994, Di Giacomo & Krapovickas 2005). ducted daily visits to check for contents, identify
Both study sites support stable Strange-tailed captured individuals and immediately release
Tyrant populations. them.
We only included in the analysis mammals of
the order Carnivora (the only order with con-
Nest predator surveys
firmed nest predators), and raptor and snake spe-
We surveyed the occurrence of mammals, raptors cies that were confirmed to be potential nest
and snakes in both study sites. To evaluate predators either during our study (i.e. we regis-
medium- and large-sized mammal occurrences tered a predation event) or by bibliographic refer-
(O’Connell et al. 2011), we set 27 camera-trap ences on nest predators of Neotropical grassland
stations in each study area between March and birds (Di Giacomo & Krapovickas 2005, Menezes
July 2015, with a minimum nearest-camera dis- & Marini 2017), hereafter referred to as mammals,
tance of 800 m. The mammal abundance survey raptors or snakes. Four camera-trap stations suf-
did not coincide with the Strange-tailed Tyrant fered technical problems and were discarded from
breeding season because the camera traps were the analyses, resulting in 26 stations in INP and 24
used during that period to detect nest predators in REB. Sampling effort was 1686 camera-trap
(see below). Even though we did not conduct the days in INP and 1834 camera-trap days in REB.
mammal survey during the study species’ breeding We identified photographed mammals to species
season, we considered the data likely to be repre- level and considered records of the same species as
sentative of mammal abundance throughout the independent if more than 12 h elapsed between
year as we used scented bait to attract predators successive pictures. We computed the frequency
and so eliminated possible differences in daily of mammal records as the number of records per
activity patterns. In addition, we have no evidence station divided by the number of days the station
of strong interannual or seasonal fluctuations of was active (Di Bitetti et al. 2009).

National Park, INP), located in the province of Corrientes, and

Reserva El Bagual (REB), located in the province of Formosa, northeastern Argentina.
Figure 1. Location of the study sites, Portal San Nicola s (Ibera

M. Browne et al.
1204
Nest searches and monitoring Breeding parameters

We conducted searches for Strange-tailed Tyrant We defined clutch size as the number of eggs laid,
nests during four consecutive breeding seasons in discarding nests that were predated or abandoned
September–December 2015–2018. In both pro- before laying was completed. We assessed hatching
tected areas we concentrated searches in grassland success only for nests that were found during incu-
patches of ~ 60 ha. We found and monitored bation, that survived until the nestling phase and
270 nests (98 in INP, 172 in REB). The study for which the number of chicks remained constant
species is sexually dimorphic and has a socially on two consecutive visits. We estimated fledgling
polygynous mating system (Di Giacomo production as the number of chicks that success-
et al. 2011). We identified territories by observing fully fledged per nest (i.e. per breeding attempt)
the flight displays of males and then focused on considering both successful and unsuccessful nests.
female behavioural cues for nest searches as only
females construct the nest, incubate and provision
Statistical analysis
nestlings (Di Giacomo et al. 2011). We located
nests during the nest building, incubation or Potential nest predator frequency of records
building stages and georeferenced them with a To compare mammal frequency of records
Garmin GPS device (Garmin International, Ola- between study sites, we used zero-inflated general-
the, KS, USA). We checked nest contents every ized linear models (GLMs) with c distribution
2–4 days (i.e. we concentrated nest visits near to when a mammal species was recorded in both
hatching and fledging dates) until nest fate was study sites. Zero-inflated models comprise a zero-
determined (Martin & Geupel 1993). When nest inflation model that describes the probability of
activity concluded, we identified the nest sub- observing an extra zero that is not generated by
strate species. the conditional model using a logit link function,
We set Reconyx HyperFire HC500 camera and a conditional model that describes the fre-
traps (Reconyx, Holmen, WI, USA), previously quency observed using a c distribution with an
cleaned with alcohol to reduce human scent, dur- inverse link function. In this way, the zero-inflated
ing both the incubation and nestling periods, at a model indicates whether the species had a higher
minimum distance of 1 m from the nest to iden- probability of being recorded in a given study site,
tify nest predators. We monitored 71 nests in INP whereas the conditional c model accounts for dif-
and 29 nests in REB. Camera traps were pro- ferences in the frequency observed among sites
grammed to take three pictures per trigger, with a considering the number of zeros that are expected
picture interval of 1 s and no quiet period. according to the error distribution. The response
variable for the zero-inflated GLM with c distribu-
tion was the frequency of mammal records at each
Nest concealment
station and the explanatory variable was study site.
We measured nest concealment of 39 nests in INP When a mammal species was only recorded in one
and 29 nests in REB by photographing 13-cm- study site, we applied a GLM with binomial distri-
wide discs (i.e. the maximum outer diameter of bution and a logit link function, where the
nests) in 2017 and 2018. Discs consisted of a grid response variable was the presence/absence of the
of 1-cm2 squares and were placed above the nest mammal predator species at each station, in order
and on its side to determine overhead and lateral to test whether the difference in probability of
nest concealment, respectively (Davis 2005). We occurrence was statistically significant. We also
took photographs of the discs at 1 m from the nest checked for differences in the frequency of raptor
in both orientations (Supporting Information, records between study sites with a GLM with
Fig. S1). If the position of the nest was altered by binomial distribution and a logit link function. The
the predator, we did not take photographs. We response variable was the presence/absence of the
estimated nest concealment as 1–p (where p is the raptor predator species per transect over all four
proportion of exposed squares) such that well- visits. We could not subject the data for snakes to
concealed nests received the highest scores statistical analysis because only three captures were
(Davis 2005). made.

Breeding parameters and nest initiation date registered, i.e. 5 September), (3) nest age
We tested for differences between study sites in (days elapsed since the laying of the first egg at
hatching success using a generalized linear mixed day 0) and (4) study site (i.e. INP and REB). We
model with binomial distribution and logit link included year and a linear trend for time of breed-
function. The response variable was the success or ing to account for interannual and within-year
failure of hatching for each egg, and we included changes in weather patterns and predator commu-
nest ID as a random effect to account for the fact nities (Dinsmore & Dinsmore 2007). We included
that eggs were clustered within nests. As data pre- nest age to account for the expected increase in
sented under-dispersion, we analysed the influence predation risk following hatching (Skutch 1949).
of the study sites on clutch size, number of nes- We used the Akaike information criterion cor-
tlings and number of fledglings, using linear rected for small sample sizes (AICc) to select the
models with a normally distributed error (McDo- best fit models from the suite of candidate models
nald & White 2010). We used a chi-squared test constructed by combining the variables that had
of homogeneity to determine if the nest initiation more support than the null-hypothesis model of
pattern (i.e. the percentage of nest attempts constant survival, S(.) in MARK notation (Burn-
throughout the breeding season) and nest substrate ham & Anderson 2002). If more than one model
use differed between study sites. had a ΔAICc less than 2, we selected the one that
had the lowest number of parameters and calcu-
Nest survival and predation rates lated Akaike weights (wi) to measure the relative
We estimated daily nest survival rate (DSR) and support of the models considered (Arnold 2010).
daily nest predation rate with the nest survival
model in Program MARK (White & Burn- Nest concealment
ham 1999). We excluded from these analyses 24 We used Mann–Whitney U tests to test for differ-
nests that were abandoned before egg-laying ences in nest concealment between successful and
began. Daily nest predation rate (DSRp) was esti- predated nests from the overhead and lateral views
mated considering successful and abandoned nests between study sites and between successful and
as not predated and calculating daily nest preda- predated nests.
tion rate as 1 – DSRp. We calculated nest initia- Unless otherwise specified, we performed all
tion dates by backdating to the egg-laying stage analyses with R 3.6.2 (R Core Team 2019) and
when nests were found during incubation and suc- used the package glmmTMB for a zero-inflated
cessfully hatched. For nests that were found and GLM with c distribution (Brooks et al. 2017). We
failed during incubation or nestling stages, we esti- set significance levels at P < 0.05 and express
mated nest age by assuming that the observed results as mean standard deviation (sd).
period was halfway between the end of laying and
hatching or by estimating nestling age, respec-
RESULTS
tively, and backdating accordingly (Dominguez
et al. 2015). The egg-laying period was considered
Nest predator surveys
to last 3 days, the incubation period 17 days and
the nestling period 12 days (Di Giacomo We did not record top predators in INP, but
et al. 2011). We calculated nest survival and nest recorded Pumas three times at three different
predation probabilities as DSRt and 1 – DSRpt, camera-trap stations in REB. We recorded four
respectively, where t was 32 days, which is the mammal nest predator species in INP and five in
length of the Strange-tailed Tyrant’s nest cycle (Di REB (Table 1). The frequency of mammal records
Giacomo et al. 2011). per camera trap per day was significantly higher in
We used Program MARK to test the effects of INP than in REB; and INP had a higher probabil-
several factors that could affect daily nest survival ity of mammal presence than REB (Table 1). The
and predation rates. For analyses of both nest sur- nest predator with the highest number of records
vival and predation rates, we built candidate was the Crab-eating Fox Cerdocyon thous, which
models with different combinations of four nest was more frequently recorded and had a signifi-
variables: (1) year, (2) a linear trend for the time cantly higher probability of presence in INP than
of breeding (days elapsed since the start of the in REB (Table 1). The Pampas Fox Lycalopex gym-
breeding season, defined as the earliest egg-laying nocercus was the second most frequently observed

predator and was also significantly more frequently All snake records occurred in INP, where two
recorded in INP than in REB (Table 1). The fre- Patagonia Green Racers Philodryas patagoniensis
quency of records for both fox species taken and one Lichtenstein’s Green Racer Philodryas
together was 10 times higher in INP than in REB, olfersii were captured. Average body mass for iden-
with values of 0.30 and 0.03, respectively. The tified predators is given in Supporting Information
Maned Wolf Chrysocyon brachyurus and Geoffroy’s Table S1.
Cat Leopardus geoffroyi were only recorded in REB
(Table 1). The probability of presence and abun-
Breeding parameters
dance of Crab-eating Racoon Procyon cancrivorus
and Lesser Grison Galictis cuja did not differ sig- The nest initiation pattern of Strange-tailed
nificantly between study sites (Table 1). Tyrants differed between sites (v2 = 27.7,
The overall frequency of raptor records was P = 0.001). It peaked 1 month earlier in INP (in
greater in REB than in INP, with two species iden- the second half of September) than in REB (in the
tified in INP and seven in REB (Table 1). The fre- second half of October, Fig. 2). Similarly, nest ini-
quency of records of raptor species did not tiation ended in early December in INP but in late
significantly differ between sites, yet they tended December in REB (Fig. 2).
to be less frequently recorded in INP than in REB, Breeding parameters did not differ between
except for the Long-winged Harrier Circus buffoni study sites. Clutch size averaged 2.94 0.41 (range
and Chimango Caracara Milvago chimango, which 2–4, n = 71) in INP and 2.88 0.42 (range 2–4,
were only recorded in INP (Table 1). n = 120, t = 1.10, P = 0.27) in REB. Hatching
National Park
Table 1. Potential nest predator frequency of records in Reserva El Bagual (REB; top predators present) and Ibera
(INP; top predators absent), Argentina, and summary statistics.
Species Common name REB INP z(a) P(a) z(b) P(b)
Mammals
Cerdocyon thous Crab-eating Fox 2.0 (47) 13.6 (354) 2.08 0.04 3.63 2.79e-4
Chrysocyon brachyurus Maned Wolf 0.7 (17) 0 (0) 1.56 0.12
Lycalopex gymnocercus Pampas Fox 0 (0) 5.7 (147) 3.11 1.89e-3
Leopardus geoffroyi Geoffroy’s Cat 0.5 (11) 0 (0) 0.71 0.48
Galictis cuja Lesser Grison 0.1 (3) 0.2 (5) 1.47 0.14 0.29 0.77
Procyon cancrivorus Crab-eating Racoon 1.1 (27) 0.3 (8) 1.45 0.15 1.42 0.16
Total 4.4 (105) 19.8 (514) 4.11 3.95e-5 1.99 0.04
Raptors
Rupornis magnirostris Roadside Hawk 0.5 (1) 0 (0) 0.003 0.99
Circus buffoni Long-winged Harrier 2.0 (4) 0.5 (3) 0.003 0.99
Buteo albicaudatus White-tailed Hawk 2.5 (5) 0 (0) 0.003 0.99
Heterospizias meridionalis Savanna Hawk 1.0 (2) 0 (0) 0.003 0.99
Milvago chimango Chimango Caracara 0 (0) 2.3 (14) 0.003 0.99
Milvago chimachima Yellow-headed Caracara 1.0 (2) 0 (0) 0.003 0.99
Falco femoralis Aplomado Falcon 1.0 (2) 0 (0) 0.003 0.99
Falco sparverius American Kestrel 4.0 (8) 0 (0) 0.003 0.99
Total 12.0 (24) 2.8 (17) 2.14 0.03
Mammal frequency of records is expressed as the number of records per station divided by the number of days the station was
active and raptor frequency of records as the number of records per transect; total number of records are expressed in parentheses.
In the models, the response variables are: mammal frequency of records expressed as the number of records per station divided by
the number of days the station was active for mammals present in both sites and as presence/absence per camera station for mam-
mal species present in one study site and presence/absence per transect for raptors. Models applied: c zero-inflated GLM for Crab-
eating Fox, Lesser Grison, Crab-eating Racoon and overall mammal frequency of records. z(a) represents z value of conditional c
model, P(a) represents P value of conditional c model, z(b) represents z value of zero-inflated model, and P(b) represents P value of
zero-inflated model. For the remaining mammal species which occurred in only one study area and for raptors, a binomial GLM test
(z, z value; P, P value) tests whether the difference in probability of occurrence between study areas is statistically significant. High-
est frequencies are highlighted in bold when significant.

Nest survival and predation rates

When analysing the variables that determined nest
survival, we discarded site and year from model
building because they performed worse than the
null-hypothesis model, and built candidate models
with time of breeding and nest age variables. The
most parsimonious model included only the variable
nest age (Table 2). DSR decreased with nest age
(b = 0.05; 95% CI –0.06 to 0.03). The average
DSR was 0.95 0.004 (n = 246) and the cumula-
tive probability of nest survival was 0.18. The DSRs
were 0.94 0.006 (n = 95) and 0.95 0.005
(n = 151), with cumulative probabilities of 0.15
and 0.20, in INP and REB, respectively.
When considering the factors that influenced
daily nest predation rate, models that included
time of breeding, nest age and site performed bet-
ter than the null-hypothesis model. We discarded
year from model building because it performed
worse than the constant survival model. The most
parsimonious model, and the only one with a
Figure 2. Percentage of nest attempts by female Strange- ΔAICc less than 2, included the variables time of
tailed Tyrants during the breeding seasons of 2015–2018 in breeding, nest age and site (i.e. INP or REB,
Reserva El Bagual (REB) and Ibera National Park (INP,
Table 3). Nest predation rate increased as time of
n = 270 nests), Argentina. Error bars are standard errors.
breeding and nest age advanced (b = 0.01; 95%
CI –0.02 to 0.003 and b = 0.05; 95% CI –0.07
success averaged 0.87 0.17 (n = 39) in INP and to 0.03, respectively) and was higher in INP than
0.91 0.18 (n = 59, z = 1.54, P = 0.13) in REB. in REB (b = 0.44; 95% CI –0.80 to 0.08;
Number of nestlings averaged 2.55 0.61 (range Fig. 3). The daily nest predation rate was
1–4, n = 49) in INP and 2.66 0.65 (range 1–4, 0.05 0.006 (n = 95) and 0.04 0.004
n = 73, t = 0.91, P = 0.37) in REB. We observed
(n = 151), with cumulative values of 0.81 and
brood reduction in two nests in INP and in eight
0.70, in INP and REB, respectively.
nests in REB. Fledgling production was 0.52 1.10
chicks per breeding attempt in INP (range 1–4,
n = 95) and 0.64 1.16 per breeding attempt in
REB (range 1–4, n = 151, t = 0.39, P = 0.39). Only Table 2. Support for models predicting daily survival rate of
Strange-tailed Tyrant nests throughout four breeding seasons
21% and 27% of nests where egg-laying began suc- from 2015 to 2018 in Ibera National Park and Reserva El
cessfully fledged nestlings in INP and REB, respec- Bagual, Argentina (n = 246).
tively. The causes of nest failure were predation and
abandonment. Nest predation was responsible for Model ΔAICc k wi
88% and 75% of nest failure once egg-laying began
S (T + Nest age) 0.0a 3 0.67
in INP and REB, respectively, whereas 12% and
S (Nest age) 1.4 2 0.33
25% were abandoned in INP and REB, respectively. S (T) 19.2 2 0.00
Nest substrate use differed between study sites S (.) 25.5 1 0.00
(v2 = 180.6, P = 0.0005). The Strange-tailed S (Site) 26.1 2 0.00
Tyrant was observed to build its nests on A. lateralis S (Year) 29.5 4 0.00
(n = 88), P. durifolium (n = 1), Schizachyrium spp.
AICc, Akaike’s information criterion corrected for small sample
(n = 1) and on an unidentified species (n = 3) in size; ΔAICc, difference in AICc relative to the best model; k,
INP and on A. lateralis (n = 27), P. intermedium number of parameters; T, linear term for time of breeding; wi,
(n = 37) and I. brasiliensis (n = 108) in REB. model weight. aAICc = 1063.7.

Table 3. Support for models predicting daily nest predation Harrier predated two nests and the Roadside Hawk
rate of Strange-tailed Tyrant nests throughout four breeding Rupornis magnirostris and the Barn Owl Tyto alba
seasons from 2015 to 2018 in Ibera National Park and
Reserva El Bagual, Argentina (n = 246).
predated one nest each (Table 4).
Model ΔAICc k wi Nest concealment

S (Nest age + T + Site) 0.0 a
4 0.78 Nest concealment did not differ between study
S (Nest age + T) 3.6 3 0.13
sites, with overhead nest concealment being
S (Nest age + Site) 5.3 3 0.05
S (Nest age) 5.8 2 0.04 0.94 0.12 and 0.93 0.12 (U = 389, P = 0.89,
S (T + Site) 24.2 3 0.00 n = 58) and lateral nest concealment being
S (T) 31.9 2 0.00 0.67 0.22 and 0.66 0.18 (U = 399, P = 0.76,
S (Site) 38.8 2 0.00 n = 58) in INP and REB, respectively. Overhead
S (.) 40.7 1 0.00
nest concealment did not differ between successful
S (Year) 43.3 4 0.00
and predated nests, with values of 0.92 0.16 and
AICc, Akaike’s information criterion corrected for small sample 0.94 0.09, respectively (U = 341.5, P = 0.91,
size; ΔAICc, difference in AICc relative to the best model; k, n = 58; Fig. 4a). Lateral nest concealment was
number of parameters; T, linear term for time of breeding; wi, greater on successful nests than on predated ones
model weight. aAICc = 891.2.
(U = 237.5, P = 0.04, n = 58; Fig. 4b), with values
of 0.74 0.19 and 0.63 0.20, respectively.
Nest predators
DISCUSSION
Fifteen (21%) of the 71 nests monitored with cam-
era traps in INP were successful, six (9%) were
Nest predator abundances, nest
abandoned and 50 (70%) were predated. We were
predation, nest concealment and
able to identify the nest predator in 27 predation
breeding phenology
events, a predator identification efficiency of 54%.
In addition, a nest containing four nestlings was In accordance with the mesopredator release
observed to experience a snake visit, resulting in at hypothesis (Soule et al. 1988, Crooks &
least one predated chick and two successfully Soule 1999), we found the frequency of fox
fledging the nest. In REB, 10 (34%) of the 29 records to be 10 times higher in INP, where top
monitored nests were successful, four (14%) were predators were absent, than in REB, where Pumas
abandoned and 15 (52%) were predated. We iden- were detected. We identified foxes as the main
tified the nest predator on seven occasions, giving nest predators of the Strange-tailed Tyrant, as was
a predator identification efficiency of 47%. found in other study systems (Rogers & Caro 1998,
The main nest predators in INP were foxes, with Ritchie & Johnson 2009, Ortega et al. 2021). Nest
11 predation events by Crab-eating Foxes and predation rate was high in both study areas but
three by Pampas Foxes, accounting for 50% of pre- increased with fox abundance.
dation events (Table 4). The Patagonia Green The mesopredator release hypothesis is not,
Racer predated nine nests, the Chimango Caracara however, the only possible explanation for the
predated three nests, and the Crab-eating Racoon increase in fox abundance observed in INP. The
and an unidentified snake predated one nest each mesopredator release hypothesis is not always sup-
(Table 4). We observed a Geoffroy’s Cat visiting a ported in manipulative experiments, and climatic
nest and, even though no apparent predation took factors (i.e. bottom-up effects) may be more influ-
place, we considered it as a potential nest predator. ential than top-down regulation in some study sys-
In addition, we found a predated nest with its origi- tems (Castle et al. 2021). Differences in food
nal position altered (i.e. fallen) and another nest availability or habitat quality could be driving the
that was destroyed by the nest predator. Even observed differences in fox abundance. The Crab-
though these nests were equipped with camera eating Fox and the Pampas Fox differ in their habi-
traps we were unable to identify the predator, but tat preferences (Di Bitetti et al. 2022), which may
it was probably a mammal. The main nest predator explain their different abundances between sites;
identified in REB was the Maned Wolf, with three and their generalized and opportunistic diets vary
predation events (Table 4). The Long-winged between study sites (Di Bitetti et al. 2009). These

Figure 3. Variation in daily nest predation rate of Strange-tailed Tyrant nests during (a) incubation and (b) nestling stage in the pres-
ence/absence of top predators in Reserva El Bagual (REB) and Ibera National Park (INP), respectively, throughout the breeding sea-
sons of 2015–2018 (n = 246 nests), Argentina. Solid lines are mean nest predation rate values; dashed lines are standard error
values.
aspects, and lack of replication of study areas, pre- et al. 2014, Suraci et al. 2016), and further studies
clude us from concluding that the higher abun- should be conducted to test possible bottom-up
dance of foxes in INP when compared with REB effects on fox abundance.
directly results from the lack of apex predators in Strange-tailed Tyrants had a more synchronized
the former. However, our results are in accordance breeding pattern in INP than in REB, with an ear-
with the hypothesis that the extinction of top lier peak and a shorter breeding season. The
predators may trigger negative cascading effects on shorter breeding season could translate to fewer
lower trophic levels, including endangered grass- nesting attempts, thus decreasing the chance of
land birds (Ritchie & Johnson 2009, Ripple producing a successful brood (Zanette

Table 4. Percentage of nest predation events in Reserva El

Bagual (REB; seven predation events), where top predators
are present, and Ibera National Park (INP, 28 predation
events), where they are not.
Species Common name REB INP
Mammals
Cerdocyon thous Crab-eating Fox 0 11
Chrysocyon Maned Wolf 3 0
brachyurus
Lycalopex Pampas Fox 0 3
gymnocercus
Leopardus geoffroyi Geoffroy’s Cat 0 0
Galictis cuja Lesser Grison 0 0
Procyon cancrivorus Crab-eating Racoon 0 1
Total 3 15
Raptors
Rupornis magnirostris Roadside Hawk 1 0
Circus buffoni Long-winged Harrier 2 0
Buteo albicaudatus White-tailed Hawk 0 0
Heterospizias Savanna Hawk 0 0
meridionalis
Milvago chimango Chimango Caracara 0 3
Milvago chimachima Yellow-headed 0 0
Caracara
Falco femoralis Aplomado Falcon 0 0
Falco sparverius American Kestrel 0 0
Tyto alba Barn Owl 1 0
Total 4 3
Snakes
Philodryas Patagonia Green Racer 0 9
patagoniensis
Philodryas olfersii Lichtenstein’s Green 0 0
Racer
Unidentified snake 0 1
Total 0 10
et al. 2000). A possible explanation for the differ-

ences observed in nest initiation is a shift in
arthropod availability, which was previously
reported to coincide with the Strange-tailed Figure 4. Average (a) overhead and (b) lateral nest conceal-
Tyrant’s breeding season (Zermoglio et al. 2016). ment of successful and predated Strange-tailed Tyrant nests
In the present study we did not investigate if dif- (n = 58) found during the 2017 and 2018 breeding seasons in
Reserva El Bagual (REB) and Ibera National Park (INP),
ferences in arthropod availability between sites
Argentina. Error bars are standard errors.
correlated with the differences observed in the
phenology of the breeding season. As breeding
synchronization has been hypothesized to result The frequency of snake records did not differ
in the dilution of predation pressure (Verhulst & between study sites, but nonetheless the Patagonia
Nilsson 2008), the earlier peak and higher breed- Green Racer was responsible for a third of nest
ing synchronization observed in INP may be an predation events and was the second main preda-
adaptive response to the increased nest predator tor in INP. Other studies have also identified
abundance and nest predation rate. A higher snakes as important nest predators (DeGregorio
breeding synchronization and earlier breeding of et al. 2014, Menezes & Marini 2017, Jefferies
prey species could be unforeseen consequences of et al. 2021). We were unable to identify the nest
the mesopredator release phenomenon. predator in ~ 45% of nest predation events, and

this could have been due to either a delay in cam- should consider multiple breeding attempts, clutch
era trap triggering or an inability of camera traps replacement and genetic variation to evaluate
to detect small predators (Ribeiro-Silva more accurately the impact of the increased nest
et al. 2018). Moreover, the camera trap’s heat sen- predation rate observed in INP on the Strange-
sor may fail to activate in the presence of cold- tailed Tyrant. Also, as overall breeding success was
blooded snakes. These factors may have caused low, the population viability of this endangered
snake predation events to be under-represented in species should be assessed considering other life
our study. history traits.
Nest concealment is key for avian predation The Strange-tailed Tyrant’s nest survival and
avoidance, and Strange-tailed Tyrants had a rela- nest predation rate were both modulated by nest
tively high overhead nest concealment value of age, whereas nest predation rate also varied
0.94. This may explain why the higher frequency between study sites and was dependent on time of
of raptor records at REB did not translate into a breeding. The finding that nest predation rates dif-
higher predation rate when compared with INP. fered between protected areas while nest survival
On the other hand, as also observed in other did not could be due to the higher nest abandon-
endangered grassland passerine species, lateral nest ment rate observed in REB. A potential cause for
concealment was lower on predated nests than on nest abandonment is adult predation by raptors.
successful ones (Repenning & Fontana 2016, The higher raptor abundance detected in REB
Browne et al. 2021). This pattern, in which better could explain the higher nest abandonment rate
concealed nests tended to be more successful, detected in this protected area, where raptor pre-
could indicate a strong effect of visually oriented dation events on adult Strange-tailed Tyrant indi-
terrestrial predators on nest fate. The study species viduals were registered (Di Giacomo &
could also be engaging in antipredator behaviour Krapovickas 2005). Nest abandonment could also
(i.e. flight-song displays or coordinated misdirec- be caused by nest flooding, which was previously
tion) aimed at deterring visual predators as reported to be the main cause of abandonment of
reported for other tyrant flycatchers (Pereyra & Strange-tailed Tyrant nests in REB but was not
Morton 2010, Gulson-Castillo et al. 2018). observed in INP (Di Giacomo et al. 2011). Both
nest survival and predation rates were influenced
by nest age, which is in accordance with the
Nest survival, nest predation and
expected increase in predation rate as nest activity
breeding parameters
increases (Skutch 1949). The observed increase in
Nest predation rate was higher in INP than in nest predation rate as the breeding season
REB and overall breeding success of Strange-tailed advanced could be due to a predator functional
Tyrants was low. Increased predation is more response, mediated either by the development of a
threatening when population sizes are small, as is search image or by prey switching (i.e. consump-
the case with endangered species, potentially tion rate varies according to prey availability), or
accentuating inbreeding and decreasing genetic to a synchronization in prey–predator breeding
variation (Frankham 1995). Nonetheless, natural timing (Roos 2002, Verhulst & Nilsson 2008,
selection may favour reduced clutch sizes and Randa et al. 2009). Future studies should assess
increased breeding attempts to maximize breeding variations in predator abundance and behaviour
success when predation rates are high (Roper throughout the breeding season.
et al. 2010). Strange-tailed Tyrant females have Nest substrate use differed between study sites.
been reported to achieve an average of two nest Female Strange-tailed Tyrants were not observed
attempts per breeding season (range 1–4) because to build their nests on Paspalum stands in INP,
of nest failure, but double-brooding (i.e. two suc- whereas in REB they did. Interspecific competition
cessful attempts per breeding season) was only for nesting sites between the study species and the
rarely recorded (Di Giacomo et al. 2011). In addi- Black-and-white Monjita Xolmis dominicanus could
tion, 17 days were reported to elapse between be taking place in INP. The Black-and-white Mon-
nesting attempts (Di Giacomo et al. 2011). Given jita uses Paspalum as a nest substrate but does not
that the breeding season was observed to end ear- occur in REB. This larger tyrant could be limiting
lier in INP, females may have fewer renesting the breeding territory available for the Strange-
opportunities there than in REB. Future studies tailed Tyrant in INP (Sætre et al. 1999, Carrete

et al. 2006). Nest substrate choice and interspecific particularly of foxes, stabilize at lower levels under
competition deserve further study to better under- the renewed presence of a top predator, 50 years
stand the driving factors of nest substrate use. after its extinction in the Ibera Wetlands.
Some Strange-tailed Tyrant breeding parameters
were similar between sites, with no differences
We thank the Administraci on de Parques Nacionales, the
found in clutch size, hatching success and fledgling Direccion de Parques y Reservas de Corrientes and the
production. These values were similar to those Direccion de Recursos Naturales for granting us permission
previously reported for the study species (Di Gia- to work in Ibera National Park (permit number: IF-2018-
como et al. 2011). The lack of differences in these 53 280 705-APN-DRNEA#APNAC) and Alparamis S.A.
breeding parameters could imply that they are not and Aves Argentinas for allowing us to work in Reserva El
responsive to increased nest predation risk or that Bagual. We also thank A.C. Giese for her help in data anal-

ysis, M. Alvarez Correa, D.G. Barcos, F. Bres, C. Bruzzese,
trade-offs in adaptive responses are involved.
B. Cortes, H.H. Correa, A.B. Fari~
na, N.V. Got, A. Moreira,
Future studies should consider variations in egg G. Reyes Benz, F. Sanchez Gavier and F.N. Zarza for their
size and clutch mass as possible responses to assistance in the field and the Associate Editor, Alexandre
increased nest predation risk (Fontaine & Millon, and two anonymous reviewers for providing many
Martin 2006). helpful comments to improve the manuscript.
CONCLUSIONS AUTHOR CONTRIBUTIONS

The majority of the Strange-tailed Tyrant’s distri- Melanie Browne: Data curation; formal analysis;
bution range is on private lands dedicated mainly funding acquisition; investigation; methodology;
to afforestation or livestock production. Previous visualization; writing – original draft. Constanza
studies have indicated that high levels of grazing Pasian: Data curation; funding acquisition; investi-
would not sustain populations of endangered grass- gation; methodology; writing – review and editing.
land species, such as our study species (Di Gia- Alejandro G. Di Giacomo: Data curation; investi-
como et al. 2010). Our findings are also gation; resources; writing – review and editing.
discouraging, as we found the breeding success of Mario S. Di Bitetti: Conceptualization; methodol-
an endangered species in protected areas to be rel- ogy; resources; writing – review and editing.
atively low. A population viability model of the Adrian S. Di Giacomo: Conceptualization; funding
Strange-tailed Tyrant should be conducted to bet- acquisition; methodology; project administration;
ter orientate conservation efforts and guarantee the supervision; writing – original draft.
persistence of one of the last populations of this
threatened species.
CONFLICT OF INTEREST
If the differences in breeding parameters
STATEMENT
reported here are indeed the result of mesopreda-
tor release, the recent reintroduction of Jaguars in The authors declare no competing interests.
the Ibera Wetlands provides a promising scenario
for endangered birds breeding in the protected
ETHICAL NOTE
area (Donadio et al. 2022). The restoration of top
predators has been found to reduce mesopredator None.
abundances (Jimenez et al. 2019). Indeed, two fox
predation events by the Jaguars released in INP in
FUNDING
January 2021 have been registered (Q. G omez
pers. comm.). Mesocarnivore control has been This study was supported by grants from the Neo-
reported to increase nest success and bird popula- tropical Grassland Conservancy, the Neotropical
tions (Palmer et al. 2019). In addition, our results Bird Club, the Association of Field Ornithologists,
illustrate the negative effect the release of native the American Society of Mammalogists, Agencia
predators, often considered to be less harmful than Nacional de Promoci on Cientıfica y Tecnol ogica
alien predators, may have on native prey species, (PICT 2014–3397, PICT 2018–3407) and CONI-
such as endangered grassland birds (Salo CET (PUE 22920180100001 CO). The funding
et al. 2007). Likewise, we expect nest predation sources had no involvement in study design, data
rates to decrease as mesopredator populations, collection, analysis and interpretation, writing of

the manuscript, nor in the decision to submit the thous) and pampas foxes (Lycalopex gymnocercus). J.
article for publication. Mammal. 90: 479–490.
Di Bitetti, M.S., De Angelo, C.D., Quiroga, V., Altrichter, M.,
Paviolo, A., Cuyckens, E. & Perovic, P. 2016. Estado de
Data Availability Statement conservacio n del jaguar en la Argentina. In Medellın, R.A.,
Antonio, J.T., Heliot, Z., Cha vez, C. & Ceballos, G. (eds) El
All scripts and files needed to reproduce our ana- jaguar en el Siglo XXI. La Perspectiva Continental. Mexico:
lyses will be archived in Mendeley Data upon Fondo de Cultura Econo mica, Universidad Nacional
Auto noma de Me xico, Instituto de Ecologıa, pp. 447–478.
acceptance.
Di Bitetti, M.S., Iezzi, M.E., Cruz, P., Cirignoli, S., Varela, D.
& De Angelo, C. 2022. Enemies or good neighbors? No
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Variation in Mesopredator Abundance and Nest Predation Rate of The Endangered Strange-Tailed Tyrant (Alectrurus Risora)

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Variation in Mesopredator Abundance and Nest Predation Rate of The Endangered Strange-Tailed Tyrant (Alectrurus Risora)

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Ibis (2023), 165, 1201–1216 doi: 10.1111/ibi.

Variation in mesopredator abundance and nest

Habitat loss is one of the main drivers of biodiver-

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

medium- and large-sized mammal populations at

© 2023 British Ornithologists' Union.

National Park, INP), located in the province of Corrientes, and

© 2023 British Ornithologists' Union.

Nest searches and monitoring Breeding parameters

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

Species Common name REB INP z(a) P(a) z(b) P(b)

© 2023 British Ornithologists' Union.

Nest survival and predation rates

© 2023 British Ornithologists' Union.

Model ΔAICc k wi Nest concealment

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

Table 4. Percentage of nest predation events in Reserva El

Species Common name REB INP

et al. 2000). A possible explanation for the differ-

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

CONCLUSIONS AUTHOR CONTRIBUTIONS

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

© 2023 British Ornithologists' Union.

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